MBL/WHOl
THE NAUTILUS
Volume 102
1988
AUTHOR INDEX
Absalao, R. S.
Adams, W. F
Askew, T. A.
AUFFENBERG, K
AUFFENBERC, T
BlELER, R.
Boss, K. J.
bouchet, p
Collins, T. M
D'AsARO, C. N
DAtiilio, A
DuShane, H.
Emerson, VV. K.
Gerberich, a. G.
Gordon, M. E.
Haimovicl M.
Harasewych, M. G.
Hayek. L. C
Hershleb, R.
Kabat, a. R.
KiLGOUR, B V\'.
36,
M.
B.
99 Lee. H. G.
125 Mackie. G L
89, 92 McKiNSTRY. D M
40. 154 McLean, J. H
40 \1endl. W.
1 MlKKELSEN, P
115 Moore, D. R.
149 Myers, B. W.
102 o'sullivan, j.
1.34 Pawlik. J. R.
106 Petit, R. E.
30 Pip, E.
110 POLACO, O. J
125 POMPONI, S. .\.
123 RoMER, N. S.
82 Sage, W E , III
, 92 DOS Santos Cruz, R. L.
56 Smith, DC
56 Snyder, MA
164 Thompson, F. G.
73 Vermeij, G. J
78, 154
73
127
99
129
1
131
102
47
47
130, 164
65
129
92
131
36, 110
99
155
54
78
102
NEW TAXA PROPOSED IN VOLUME 102 (1988)
GASTROPODA
Perotrochus charlestonensis Askew, new species (Pleurotomariidae)
Macrarcnc difiilala McLean. Absalao and Santos Cruz, new species (Turbinidae)
Nerita fortidentala Wrmeij and Collins, new species (Neritidae)
Alvania colombiana Romer and Moore, new species (Rissoidae)
Faiartia (Murexiella) shaskyi DAttilio and Myers, new species (Muricidae)
Metula crosnieri Bouchet, new species (Buccinidae)
Metula africana Bouchet, new species (Buccinidae)
Latirus martini Sn\der, new species (Fasciolariidae)
Vasum Stephana Emerson and Sage, new species (Turbinellidae)
Axi'lella Petit new name (Cancellariidae)
Terebra imitatrix Auffenberg and Lee, new species (Terebridae)
Hypselostoma holimanae Thompson and Lee, new species (Pupillidae)
CEPHALOPODA
Eledone gaucha Haimovici, new species (Octopodidae)
89
100
102
131
106
150
151
54
36
130
154
78
82
rHE NAUTILUS
Volume 102, Number 1
February 16, 1988
ISSN 0028-1344
A quarterly devoted
to malacology.
Marine Biological Laboratory
LIBRARY
FEB 2 4 1988
Woods Hole, Mass.
EDITOR-IN-CHIEF
Dr. M. G. Harasewych
Division of Mollusks
National Museum of
Natural History
Smithsonian institution
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ASSOCIATE EDITOR
Dr. R. Tucker Abbott
American Maiacoiogists, Inc.
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CONSULTING EDITORS
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Department of
Living Invertebrates
The American Museum of
Natural Historv
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Mr. Samuel L. H. Fuller
1053 Mapleton .Avenue
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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 I'niversity
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Dr. Aurele La Rocque
Department of Geology
The Ohio State L'niversitv
Columbus, OH 43210
Dr. James H. McLean
Department of Malacology
Los Angeles County Musetim of
Natural History
900 E.xposition Boulevard
Los Angeles, CA 90007
Dr. Arthur S. Merrill
% Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Donald R. Moore
Division of Marine Geologv
and Geophysics
Rosenstiel School of Marine and
Atmospheric Science
University of Miami
4600 Rickenbacker Causeway
Miami, FL 33149
Mr Richard E Petit
P.O Box 30
North Myrtle Beach, SC 29582
Dr. Edward J. Petuch
Department of Geology
P"lorida .Atlantic L'niversitv
I?6ca Raton,' "Fl' 33431
Dr. G. Alan Soleiri
Department of Invertebrates
Field Museurri o£ Natural Historv
Chicago, IL 60605
Dr. David H. Stansber\
Museum of Zoology
The Ohio State Universitv
Columbus, OH 43210
Dr. Ruth D Turner
Department of Mollusks
Museum of Comparative Zoolog\
Harvard Universit\
Cambridge, MA 02138
Dr. Geerat J. V'ermeij
Department of Biology
University of Mar\ land
College Park, MD 20740
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Division of Biology and Living
Resources
Rosenstiel School of Marine and
Atmospheric Science
Universit) of Miami
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Miami, FL 33149
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THEt^NAUTILUS
CONTENTS
Volume 102, Number I
February 16, 1988
ISSN 0028-1344
Rudiger Bieler
Paula \1. \likkelsen
Aiiatoni) and reproductive biology of two western Atlantic
species of N'itrinellidae, with a case of protandrous
hermaphroditism in the Rissoacea
Helen DuShane
Geographical distribution of some Epitoniidae (Mollusca:
Gastropoda) associated with fungiid corals
30
\^ illiam K. Emerson
Salter E. Sage, HI
A new species of Vastim (Gastropoda: Turbinellidae) from
off Somalia
36
Kurt Auffenberg
Troy Auffenberg
Density, spatial distribution, activity patterns, and biomass
of the land snail, Geophoriis bothropoma Moellendorff
(Prosobranchia: Helicinidae)
40
Notice
46
Marine Biological Laboratory
LIBRARY
FEB 24 1988
Woods Hole, Mass.
THE NAUTILUS 102(l):l-29, 1988
Page 1
Anatomy and Reproductive Biology of Two Western Atlantic
Species of Vitrinellidae, With a Case of Protandrous
Hermaphroditism in the Rissoacea
Kiidiger Bieler
Smithsonian Marine Station
at Link Port
5612 Old Dixie Highway
Ft Pierce, FL 3-1946, USA
Paula M. Mikkel^en
Harbor Branch Oceanographic
Institution
5600 Old Dixie Highway
Ft. Pierce, FL 34946, USA
ABSTRACT
Two western Atlantic vitrinellids, Cyclostremiscus beauii
(Fischer, 1857) and Circulus texanus (Moore, 1965) new com-
bination are redescribed based on a study of live snails from
the burrows of the stomatopod crustacean Lysiosquilla scabri-
cauda (Lamarck, 1818). Anatoms and reproductive biology are
emphasized, with the first recorded description of spawn and
larval development in the famiK'. Synon\mies are given, and
a lectotype is selected for Cy. beauii. Literature data concerning
anatom) are reviewed for the marine near-planispiral risso-
aceans; Vitrinellidae (including Cyclostremiscus and Circulus,
with Circulidae as a synonym) is considered distinct from Tor-
nidae. Unusual morphological aspects of Cy. beauii are dis-
cussed, including external ciliation patterns, pallial tentacles
(which are functionalK and morphologically different from
each other), and stomach morpholog>' (with a large posterior
chamber). Protandrous sequential hermaphroditism in Cy. beauii
is inferred (for the first time in the Rissoacea) from morpho-
logical evidence of sex change correlated with size.
Key words: Cyclostremiscus; Circulus; Vitrinellidae; Torni-
dae; Rissoacea; systematics; anatomy; hermaphroditism.
INTRODUCTION
Despite the great number of nominal species assigned to
the poorK defined, cosmopolitan, marine rissoacean fam-
ily \ itrinellidae Bush, 1897, little is known about the
biology of its members. Their small size, mostly unknown
habitats, poor representation in collections, and frequent
confusion with other small-shelled members of groups
such as Cyclostrematidae Fischer, 1885, Skeneidae Thiele,
1929, Turbinidae Rafinesque, 1815, and Tornidae Sacco.
1896, may account for the lack of attention paid to this
family, and why authors who have attempted revisions
(e.g., Trvon, 1888; Bush, 1897; Melvill, 1906; Pilsbry &
Olsson, 1945; Pilsbry & McGinty, 1945a,b, 1946a,b, 1950;
Laseron, 1958; Moore, 1964; Adam & Knudsen, 1969)
relied almost entireK' on shell characters to distinguish
ta.xa on all taxonomic levels.
Aside from Pilsbry and McGinty 's (1945a, 1946b)
sketches of crawling animals of several nominal genera.
and studies on the gross morphology of Cochliolepis
parasitica Stimpson, 1858 (by Moore, 1972) and C.
albicerata Ponder, 1966 (by Ponder, 1966), published
information on anatomy is available for only one species
of this family, Circulus striatus (Philippi, 1836) from
the eastern Atlantic. Data on Ci. striatus, extensively
presented by Fretter (1956) and later summarized by
Fretterand Graham (1962, 1978), were based on material
which Fretter and Graham (1978:228) described as "the
only live specimens . . . obtained by Fretter (1956) from
the stomach of the starfish Astropecten . . . dredged on
sandy bottoms 28-30 m deep in the Gulf of Gascony."
Thus, the present concept of vitrinellid anatomv' is
based mainly on Fretter 's description of a single species,
Circulus striatus, the tvpe species of the name-bearing
genus of the nominal family Circulidae (see Discussion,
below). Discussions of phylogenetic relationships (or s\n-
onymies) between families such as Vitrinellidae and Tor-
nidae have demonstrated the need for anatomical and
reproductive data for these groups (e.g., Fretter, 1956;
Tavlor & Sohl, 1962; Moore, 1965; Golikov & Starobo-
gatov, 1975; Boss, 1982; Graham, 1982; Ponder, in press).
Studies on two species of western Atlantic N'itrinellidae
are reported herein. Populations of Cyclostremiscus
beauii (Fischer, 1857) and Circulus texanus (Moore, 1965)
were discovered in Florida in sand-flat burrows of the
stomatopod crustacean Lysiosquilla scabricauda (La-
marck, 1818). These burrows are U-shaped and extend
up to 1.5 m into the sediment, with horizontal distances
of 6-7 m between the two openings; a pair of stomatopods
inhabits each burrow system, maintaining it over long
periods of time (Serene, 1954; R. B. Manning, personal
communication). Cyclostremiscus beauii, with a shell
diameter of 6-8 mm, is one of the largest vitrinellids, a
fact that facilitated detailed studv of its morphology and
anatomy and allowed a test of Fretter s hypothesis that
some of the characters found in the much smaller (2 mm)
Circulus striatus are size-related (Fretter, 1956:380).
Special emphasis was placed upon characters of the
reproductive svstem, for which Moore (1964:18) noted
"Nothing is known of the reproduction of the famiK
Page 2
THE NAUTILUS, Vol. 102, No. 1
except that the animals are dioecious, and that the male
is provided with a penis." Data presented herein suggest
that Cy. beauii is a protaiidrous sequential hermaphro-
dite. Only incomplete data are available for Circulus
texanus, as animals of this species were collected only
twice, both times before the actual beginning of this
study. However, since gross anatomy, spawn, and de-
veloping eggs were observed for this species, available
data are presented here.
MATERIAL AND METHODS
Samples were taken from stomatopod burrows in shal-
low-water sand flats in the Indian River lagoon just inside
the Ft. Pierce Inlet, St. Lucie County, eastern Florida
(27°28.3'N, 80°17.9'W) using a stainless steel bait pump
("yabby pump") in conjunction with sieves of 1-2 mm
mesh. Depths at low tide ranged from less than 0.5 m
to supratidal, wherein the water level lay several centi-
meters below the level of the sand.
Living snails were maintained in Bnger bowls of sea-
water at room temperature (24 °C). Carmine and fluo-
rescein sodium particles were used to observe ciliary
action and currents produced by the animals. For gross
dissections, shells were cracked and animals subsequently
relaxed using magnesium sulfate crystals ("epsom salts").
Other anaesthetic chemicals (7% magnesium chloride in
distilled water, menthol crystals) were tried but pro-
duced little or no effect with gradual addition, or too
strong an effect resulting in retraction. Methylene-blue/
basic-fuchsin and neutral red were used to better delin-
eate tissues and organs in gross dissections. Structures
and organs were measured following in part the outline
given by Davis and Carney (1973: fig. 4A). Terminology
of the nervous system is after Davis et al. (1976).
For histological sectioning, animals were relaxed as
above and fixed in either glutaraldehyde-formalin so-
lution (4% formalin, 2.5% glutaraldeh\de in 0.1 M phos-
phate buffer, pH 7.2) or 5% buffered formalin (Humason,
1962:14). Shells were either broken and removed, or dis-
solved in a 1% solution of ethylene diamine tetraacetic
acid (EDTA, adjusted to pH 7.2). Specimens were
embedded in paraffin, sectioned at 5-7 ^m and stained
with Alcian Blue/ Periodic- Acid-Schiff (PAS), counter-
stained with Harris' Hematoxylin/Eosin (Humason, 1962:
125, 269, 298). Staining reactions described in the text
refer to this method unless otherwise rioted. Photomi-
crographs of sections were taken with a Zeiss Photomi-
croscope-3.
Radulae and jaws were extracted by dissolving the
surrouiitling soft tissue in a solution of lO'^ sodium hy-
droxide. Spermatozoa were prepared for SEM by placing
a drop of concentrated sperm in seawater onto a coverslip
placed in a covered petri dish containing droplets of 25%
glutaraldeliyde, and passing the coverslip through an
etlianol series ending in acetone, and then critical-point
drying the sample. Whole animals were fixed, passed
through an ethanol series, transferred to amyl acetate,
and critical-point dried. These, together with air-dried
shells, radulae, jaws, and opercula were coated with gold/
palladium, and scanned using a Zeiss Novascan-30. Fig-
ures 1-3 were photographed using a Hitachi S-570 scan-
ning electron microscope. Radular terminology is after
Bandel (1984:3).
Protoconch and teleoconch diameters were recorded
as the greatest dimension perpendicular to the columellar
axis. Teleoconch height was the greatest dimension par-
allel to the columellar axis, measured from the apex to
the base of the aperture. Umbilical diameter was the
greatest distance between the columellar lip and the most
prominent portion of the umbilical wall, measured in
ventral \ iew . Teleoconch w horls were counted from the
protoconch Il-teleoconch line to the farthest extent of
the periphery (= the point on the outer lip used for
greatest shell diameter). The number of protoconch whorls
was determined b\ the method of Ta\ lor (1975:10; sum-
marized by Jablonski & Lutz, 1980:332, fig. 4).
Cited repositories are (* indicates location of voucher
material):
ANSP — Academy of Natural Sciences of Philadelphia,
PA.
CAS — California Academy of Sciences, San Francisco.
*IRCZM — Indian River Coastal Zone Museum, Harbor
Branch Oceanographic Institution (HBOI), Ft. Pierce,
FL.
MCZ — Museum of Comparative Zoology, Harvard Uni-
versity, Cambridge, M.A.
MNHN — Museum National d'Histoire Naturelle, Paris.
*RSMAS — Rosenstiel School of Marine and Atmospheric
Sciences, University of Miami, FL.
UNC-IMS — Institute of Marine Sciences, University of
North Carolina, Morehead City, NC.
*USNM — National Museum of Natural History, Smith-
sonian Institution, Washington, DC.
RESULTS
Rissoacea Gray, 1847
= Truncateilacea Gray, 1840, "submission to be made to ICZN
to suppress this name" (Ponder, 1985:15).
Vitrinellidae Rush, 1897
Cyclostremiscus Pilsbry & Olsson, 1945:266.
Type species: VitrineUa panamensis C B. Adams, 1852
(by original designation).
Cyclostremiscus beauii (Fischer, 1857)
(figures 1-52; tables 1, 2)
Adeorbis Beauii Fischer, 1857a:173 [nomen nudum].
Adcorlns Reauii Fischer, 1857b:286, pi. 10, fig. 12 [Guade-
loupe].— Bu.sh, 1897:104.
Cyclostrema Beam [sic].— A. Adams, 1866:251, pi. 255, fig. 25
[after Fischer, 1857b].
Cyclostrema bicarinata Guppy, 1866:291, pi. 17, figs. 5a, b
["Miocene" (Lower Pliocene), Jamaica].
R. Bieler and P. M. Mikkelsen, 1988
Page 3
Skenea sulcata "Bush" Simpson, 1887:61 [nomen nudum, see
Moore, 1964:131],
Adeorlris Beaui.—DaW, 1889:150; 1892:345.
Adeorbis Beam var bicarinata —DaW. 1903:1595 ["Oligo-
cene," Jamaica].
"C.irculus" Incarinatus. — Woodring, 1928:439, pi. 37, figs. 10-
12 [neotype designation].
"Adeorhis" heaui. — Woodring, 1928:440.
C.irculus stirophorus M. Smith, 1937:67, pi 6, figs. 2a, b [Pho-
cene, Florida].
Cyclostrema angulata. — Hertlein & Strong, 1951:110 [West
Indies].
Cyclostremiscus {Ponocyclus) heaui Incarinatus. — Pilsbry, 1953:
427, pi. 55, figs. 1-le.
Vitrinella iSolariorbis) fceaui.— Abbott, 1954:138.
Vitrinella beaui. — Wells et ai. 1961:267.
Cyclostremiscus beaui. — Moore, 1964:131-135. — Morris, 1973:
138, pi. 40, fig. 19.— Porter, 1974:143.— Emerson & Ja-
cobson, 1976:64, pi. 18, fig. 21 —Abbott & Dance, 1982:
58, text-fig.
Cyclostremiscus {Ponocyclus) beaui. — Warmke & Abbott, 1961:
60, pi. 11, fig. b.— Humfrey. 1975:76, pi. 3, figs. U-lla.
Cyclostremiscus (Ponocyclus) beauii. — Abbott, 1974:85, text-
fig. 786.— Rios, 1975:38, pi. 10. fig. 131; 1985:41, pi. 16,
fig. 177.— Yokes & Yokes, 1983:15, pi. 25, figs. 4-4a.
Material examined: Lectotype (designated herein): 10.5
mm, MNHN unnumbered (Guadeloupe). Neotype of
Cyclostrema bicarinata. 7.4 mm, USNM 115621 (Plio-
cene, Jamaica); other material: 60 specimens: NORTH
CAROLINA: 1 specimen with dried animal, BEVER-
IDGE Sta. I (ex Astropecten). UNC-IMS. FLORIDA: Ft.
Pierce Inlet: 10 March 1987, 1 male, 3 unsexed; 2-3 May
1987, 1 male; 24 June 1987, 3 males; 3 August 1987, 6
males; 31 August 1987, 7 females, 1 1 males; 1 4 September
1987, I female; 27 September 1987, 3 females, 2 males.
Peanut Island, Lake Worth Inlet: 11 August 1987, 1 fe-
male, 1 male. Boynton Beach: 1 shell, ANSP 277740.
Miami: 1 shell, EOLIS Sta. 311, USNM 449192. Fowey
Light: 2 shells, EOLIS Sta. 187, USNM 449193; 1 shell',
EOLIS Sta. 129, USNM 449194; 1 shell, EOLIS Sta. 142,
USNM 449195; 1 shell, EOLIS Sta. 170, USNM 449196;
1 shell, EOLIS Sta. 355, USNM 449198. Turtle Harbor:
2 shells, EOLIS Sta. 61, USNM 449197. Sand Key: 1 shell,
EOLIS Sta. 162, USNM 449199. Kev West: 1 shell, EO-
LIS Sta. 63, USNM 449200. Drv Tortugas: 1 shell, USNM
61114; 1 shell, USNM 271949. Cape San Bias; 1 shell,
USBF Sta. 2402, USNM 323914. CARIBBEAN: St. Mar-
tin; 1 shell, ANSP 20621. Jamaica: 1 shell, USNM 426872;
1 shell, USNM 442372. SOUTH AMERICA: CoveSas,
Colombia: 1 shell, USNM 364409.
Description
Teleoconch (figures 1-3): Shell large for family [usu-
ally 2-3 whorls, 6-8 mm diameter (x = 6.9, n = 56);
height: X = 3.8, n = 33; umbilical diameter: x = 1.5, n =
30; largest specimen (female): diameter 11.5 mm, height
6.2 mm, umbilicus diameter 2.4 mm, teleoconch whorls
3%], nearly planispiral. Opaque-whitish, with 5-8 strong,
concentric ribs on apical side, above peripheral keel.
Widely-spaced irregular pustules between ribs (figures
Table 1 . .Anatomical characters and character states, and hab-
itat types of species in the vitrinellid-toriiid complex
1) Projecting snout bilobed, lateral extensions: (a) absent; (b)
present.
2) Cilia along cephalic tentacles: (a) absent; (b) present
3) Terminal stiff setae on cephalic tentacles: (a) absent; (b)
present.
4) Number of pallial tentacles.
5) Pallial tentacles: (a) all finger-shaped; {b) upper finger-
shaped and lower paddle-shaped.
6) Upper pallial tentacle: (a) naked; (b) with motile cilia; (c)
with stiff setae; (d) with distinct motile cilia and/or stiff
setae.
7) Lower pallial tentacle: (a) naked; (b) with motile cilia; (c)
with stiff setae; (d) with distinct motile cilia and/or stiff
setae.
8) Gill filaments: (a) projecting from aperture in crawling
position; (b) not projecting from aperture in crawling po-
sition.
9) Anterior foot margin: (a) straight or only weakly indented;
(b) cleft.
10) Posterior foot margin: (a) simple and rounded or weakly
indented; (b) cleft.
11) Operculum nucleus: (a) concentric; (b) subcentral.
12) Number of whorls on operculum.
13) Eyes: (a) distinctly developed; (b) lack nerve supply.
14) Osphradium: (a) small, ciliated groove; (b) distinctly de-
veloped, paralleling ctenidium.
15) Penis: (a) simple, without glandular processes, recurved;
(b) with glandular area, directed straight back; (c) with
several, finger-like processes.
16) Habitat: (a) under rocks; (b) under scales of annelid Poly-
odontes lupina (Stimpson); (c) ? from stomach of starfish;
(d) in burrows of stomatopod Lysiosquilla scabricauda (La-
marck); (e) sandy mud bottom; (f) under large boulders
on well-ox\genated sand\ mud.
5-8) on first whorl, becoming more dense and regularly
spaced on second whorl. Fields of pustules on body whorl
intersected by smaller concentric ridges, 4-6 between
major ribs, added first between suture and first major
rib, subsequently between peripheral ribs. Strong cords
forming sharp peripheral and basal keels, separated by
wide flat area, inclined ventrally and marked by growth
lines, spiral cords, and pustules. Base widely umbilicate,
with 0-5 concentric ribs between basal keel and umbi-
licus; larger specimens showing decreasing number of
ribs with increasing size. Umbilical wall often with 2-4
narrow ribs. Outer lip sinuous, with shallow sutural sinus.
Microstructure (figure 4) of 3 layers: 2 thick cross-la-
mellar layers and 1 thin homogenous outer layer thick-
ened to form spiral ribs.
Protoconch (figures 9-11): Diameter 0.40-0.48 mm
(f = 0.45, n = 32). Protoconch I (prior to hatching)
smooth, of about 1 whorl (diameter ~ 0.23 mm). Pro-
toconch II (after hatching, before settling) of an addi-
tional whorl, sculptured with irregular, more-or-less con-
centric markings (figure 11). Total protoconch of 2 whorls,
rather high-spired (spire angle = 50°).
Page 4
THE NAUTILUS, Vol, 102, No. 1
Table 2. Summary of anatomical and habitat data for species in the vitrinellid-tornid complex. Characters and character states
are listed in table 1. Sources of data listed under each species. ** = type species of genus; * = synonym of type S[)ecics of genus.
*' = c according to VUmre. 1964:22; d according to Moore. 1964:159 W] biological data from Florida, except for Cochliolepis
parasitica (also South Carolituil. Co. alhiccrata (-\e« Zfalaiid ' Cirrulus sirialiis (iucrnseN. English Channel
Characters
Species
12 3 4 5 6
8 9 10 11 12 13 14 15 16
Croup 1
**Vitrinella helicoidea C. B .\dunis, 1850
Pilsbry & McCinty, 194.5a: pi. 2, fig. 5; 1946b: 13.
*Vitrinella praecox Pil.sbr\ & McGinty, 1946
Pilsbry & McGinty, 19'45a: pi. 2, fig. 4; 1946b:14.
Teinostoma carinacallus Pilsbr\ & McGinty, 1946
Pilsbry & McGinty, 1946b:17, pi. 2, fig. 6b.
Teinostoma lerema Piisbrv & McGintv, 1945
Pilsbry & McGintv, 194.5a:6, pi. 2, fig. la.
Teinostoma parvicallum Piisbrv 6< McGint>, 1945
Pilsbry & McGinty, 194.5a:4. pi 2, fig. 2.'
**Pleuromalaxis halcsi (Piisbrv & McGinty, 1945)
Pilsbry & McGinty, 1945a: 10, pi. 2, fig. 8.
**Cochliolepis parasitica Stirnpson, 1858
Stimpson, 1858:307ff.; Moore, 1972:100ff.
Cochliolepis allncerata Ponder, 1966
Ponder, 1966.38, pi 5
*Circultts striatus (Philippi, 1836)
Fretter, 1956:369ff ; Fretter & Graham, 1978:227£f.
Circtilus texanus (Moore. 1965)
This paper
Cyclostremiscus beaitii (Fischer, 1857)
This paper
Cyclostremiscus pentagonus (Gabb, 1873)
Bush, 1897:127, pi. 22, figs. 6, 12a-g [as Skenea trilix].
Group 2
**Tomura hicaudata (Piisbrv & McGintv, 1946)
Pilsbry & McGinty, 1945a: pi. 2, fig.' 9; 1946b:15.
*Parviturhoides interruptus (C. B. .\dams, 1850)
Moore, 1962:695ff., fig. IB; 1964:21, 156ff.;
1972:106ff., figs. 5, 6.
Group 3
**Tornus subcarinatus (Montagu, 1803)
Woodward, 1898:140ff., pi. 8, figs. 1-3, 5-7; Fretter &
Graham. 1978:229ff; Graham, 1982:144ff.
a b
a b
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b 2
b 2
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External anatomy and organs of the mantle cavity (fig-
ures 12- 18, 20-25, 27, 34-37): Living animal trans-
lucent vellovvish-orange, with buccal mass, anteriormost
gill tilaments, and tip of penis rose-pink; digestive gland
brownish-orange. Base of ctenidial lamellae on osphra-
(iial side pigmented white; central area of osphradium
vvliitf Hanked bv brown lines on each side, resulting in
pattern of parallel lines on left body side (figure 12).
Long snout terminating in pair of muscular "lips," sep-
arated by vertical slit with mouth opening. Lips sepa-
rated Irom remainder ot snout bv strongiv ciliated groove.
Serial sections revealed 2 narrow longitudinal bands of
cilia, situated in grooves on each side, beginning shortly
behind tip of snout. Two slender, De.xible, cephalic ten-
tacles reaching appro.ximately twice length of snout when
fullv extended. Left tentacle fitting into shallow notch
formed bv ventral shell keel (figure 14). iMack eves on
outer expanded bases of cephalic tentacles, each equipped
with spherical, transparent lens.
Ciephalic tentacles exhibiting elaborate pattern of mo-
Figures 1-11. Cyclostremiscus heauii. specimens from Ft. Pierce Inlet, Florida (SEM) (figures 1-3, USNM 846323). 1. Shell,
apical view (8.0 mm diameter). 2. Shell, umbilical view (5.0 mm diameter) 3. Shell, apertural view (4.1 mm diameter) 4.
Microstructure of lateral body wall; fracture surface parallel to growing edge 5. Teleoconch sculpture, apical view. Circled numbers
indicate location of sculptural details in figures 6-8. Stars indicate location of sutures. 6. Detail of teleoconch sculpture, first whorl.
7. Detail of teleoconch sculpture, second whorl. 8. Detail of teleoconch sculpture, third (= body) whorl. 9. Protoconch, apical view.
,\rro\v indicates sculptural line between protoconchs I and II. 10. Protoconch, lateral view. 11. Sculpture of protoconch I (left,
smooth) and protoconch II (right, sculptured) Scale bars: 4, 5, 8-10 = 0.1 mm; 6, 7, 11 = 10 fim.
R. Bieler and P. M. Mikkelsen, 1988
Page 5
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Page 6
THE NAUTILUS, Vol. 102, No. 1
R, Bieler and P. M. Mikkelsen, 1988
Page 7
lilc cilia and stiff bristles. Ventral tip of each tentacle
with L-shaped, heaviK -ciliated groove surrounding
smooth area, forming tactile pad (figure 25, tp). Nu-
merous stiff bristles (lost during fixation, therefore not
evident in histological or SEM preparations) distal to and
just behind pad; additional bristles sparsely distributed
over distal quarter of tentacle. T\\ o longitudinal ciliated
tracts, situated in grooves, extending from region of pad
on ventral side. Near tentacle base, innermost groove
sloping dorsally toward dorsal midline; outermost groove
ending proximal to expanded tentacle base (figure 20).
[•fattened base of tentacle facing snout surface covered
b\ additional, irregular, parallel tracts of cilia (figure 20).
Third longitudinal ciliated groove on dorsal side of ten-
tacle extending from flattened, triangular area just below
eye to near tentacle tip (figure 18). Ciliated grooves usu-
alK lined b\ narrow bands of brown pigment. Cilia fulK'
retractable into grooves. C^ilia in grooves with distinct
movement pattern (right tentacle: dorsal row — toward
tentacle base, inner and outer ventral rows — toward tip;
left tentacle: opposite directions). (These movements are
easily mistaken for direction of ciliary beating, and there-
fore current flow. However, further microscopic obser-
vation with the aid of carmine and fluorescein sodium
particles revealed lateral beating of the individual cilia
across the tentacle, away from the snout, indicating that
the apparent ciliary movement actualK reflects the con-
duction of nervous impulses along the tentacle.) In cross-
section, each tentacle usually with 3 nerves, 1 larger
blood sinus, and several smaller blood spaces in central
area, especially around nerve cords (figure 24).
Foot (figures 12-14) elongate (just reaching posterior
shell margin in crawling animal), flattened, densely and
finely ciliated, with anterolaterally recurved corners;
broadh rounded posteriorly and slightK indented at pos-
terior terminus. Anterior pedal mucous gland (figure 14,
amg) present, opening at center of transverse slit across
anterior, leading edge of foot. No posterior mucous gland
or metapodial tentacles. Locomotion by ciliary action.
Operculum (figures 17, 34-37) corneous, circular, mul-
tispiral, with about 7 whorls and small internal knob,
supported by finely ciliated opercular lobes (figure 17,
ol) on dorsal side of foot. Lobes simple, unpigmented,
without tentacles. Lemon-shaped muscle scar on internal
surface of operculum (figure 35).
Mantle edge (figures 12, 13) somewhat scalloped, cor-
responding to spiral ribs of shell. Large monopectinate
ctenidium, attached along entire length to inner surface
of mantle, originating on posterior left, curving over
dorsum, terminating just above right eye where last few
gill filaments protrude from aperture. Numerous (< 180)
gill filaments (figures 16, 21) flattened, elongated leaflets,
almost finger-like when contracted, forming tapered
blades when extended. Filaments longest in central part
of gill, decreasing somewhat in size toward both ends.
"Supporting rods" lacking; filaments well supplied with
blood spaces and muscles (transverse muscle bands giving
extended filaments ladder-like appearance; figure 16).
Both sides of blade carrying wide band of cilia off-center,
closer to left (osphradial) side. Along right side in same
relative position, each filament with longitudinal row of
small embedded crystals (figure 16, cr). Narrow rim clear,
somewhat thinner on "crystal" side, bearing continuous
band of cilia. Filaments draining into large efferent bran-
chial vessel leading to heart (figure 12). Filaments hardly
reacted to direct physical stimuli, contracting rapidly
when either cephalic tentacle or finger-shaped pallial
tentacle (see below) touched.
Whitish hypobranchial gland (figures 12, 13) paral-
leling entire length of rectum, most conspicuous poste-
riorly. Osphradium (figures 12, 22, 23) paralleling almost
entire left side of gill, comprised of wide central area
and two strongK developed, heavily ciliated, lateral zones.
Central area w ith irregular chevron-like pattern of tracts
of shorter cilia. Osphradial ganglion (figure 12, osg) very
conspicuous at point about 'A of total length from mantle
edge. Mantle cavity ending immediately behind poste-
rior end of ctenidium.
Two pallial tentacles (figures 13, 27) arising from just
inside right mantle edge; upper (= most dorsal) tentacle
finger-shaped, unciliated, somewhat closer to mantle edge,
curling into shallow sutural sinus, directed dorsally. Low-
er tentacle paddle-shaped, ciliated at edges (except on
narrow stalk) and also across its broad surface; narrow
band at tentacle edge unciliated. Single stiff bristles some-
times occurring at tip. Lower tentacle originating at point
on mantle edge ventral and slightly more interior to
upper finger-shaped tentacle; directed anterolaterally.
Stimulation with forceps or needle caused immediate
contraction of upper tentacle (followed by contraction
of cephalic tentacles and exposed ctenidial filaments).
Lower tentacle showed little response to touch and can-
not markedly contract (compare tentacles in figure 27).
Lower tentacle observed to regulate and enhance water
Figures 12-17. Cyclostremiscus beauii. 12. Male, left side, in crawling position (shell removed). Penis reflected anteriorly, out
of mantle cavitv . 13. Female, right side, in crawling position (shell removed). Mantle slightly reflected to show relative insertion
points of pallial tentacles. 14. Crawling animal, ventral view. 15. Diagrammatic cross-section through mantle cavitv of male, at
level of osphradial ganglion. 16. Gill filament. 17. Diagrammatic view of closed operculum, as seen when animal is retracted,
showing position of opercular lobes. Scale bars: 12-14 = 1.0 mm; 15 = 0.5 mm; 16 = 1 mm.
ag, albumen gland; amg, anterior mucous gland; br, immobile bristles; cit, ciliary tract; com, columellar muscle; cont. connective
tissue; cr, crvstals; ct, ctenidium; dg, digestive gland; ebv, efferent branchial vessel; es, esophagus; he, heart; hg, hypobranchial
gland; in, intestine; ki, kidney; Ipt, lower pallial tentacle; mc, mantle cavity; me, mantle edge; ol, opercular lobe; op, operculum;
OS, osphradium; osg, osphradial ganglion; ov, ovary; pe, penis; pvd, pallial vas deferens; re, rectum; sh, shell; sn, snout; st, stomach,
ten, cephalic tentacle; upt, upper pallial tentacle.
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THE NAUTILUS, Vol. 102, No. 1
H. Bieler and P. M. Mikkelsen, 1988
Page 9
tlow , by paddle positioning and by strong ciliary action,
it'spccti\el\-, in or out of right side of mantle cavity.
Alimentary system: Mouth opening between pair of
muscular lips into large buccal mass (figure 42; length
1..') mm in specimen 8.5 mm shell diameter). Radula
protrmling from short radular sac (figure 42, ras) ex-
tending somewhat behind and to left of buccal mass.
Paired jaws (figure 41) each crescent-shaped, appro.x-
imately 0.5 x 0.16 mm (in specimen 7.8 mm shell di-
ameter), composed of interlocking diamond-shaped ele-
ments 14 /im in length. \ar\ ing slightK in shape across
surface of jaw.
Radula (figures 28-33) taenioglossate, with about 100
rows (max. 138; n = 7), length == 1.3 mm, width ~ 0.2
mm. Rachidian tooth (figure 31) wider than long (0.40
/im w ide. n = 8), with acute posterior corners projecting
laterally, and concave front. Main cusp narrowly-trian-
gular, unserrated, with 4-5 flanking cusps on each side
(number of flanking cusps var\ing within single radula,
apparently by splitting and fusion), decreasing laterally
in size; base with 1 strong basal denticle per side, midway
between posterior corners and central ridge. Lateral tooth
ifigures 32, 33) with asymmetrical cutting edge, strongly
indented at front edge, bearing large, narrowK -trian-
gular, unserrated main cusp and highly variable number
of flanking cusps (3-6 inner, 7-8 outer) decreasing in
size laterally. Base of lateral tooth with broad central
ridge; basal platform long, blade-like. Apex of inner mar-
ginal (figures 30, 33) with short, stout main cusp, and
12-14 subequal, inner and outer flanking cusps. Stalk
long, blade-like, with robust supporting ridge. Apex of
outer marginal (figures 28, 30) with sharp, undulating
inner margin (sometimes finely incised into closely-ad-
hering flanking cusps), and smooth, rounded outer mar-
gin.
Esophagus opening widely into buccal cavity, without
esophageal pouches. Salivary glands (figures 19, 26, 42)
narrow, tube-like, empt\ ing into buccal mass at its lateral
mid-point, extending posteriorly along esophagus. Posi-
tion of salivar\ ducts relative to circumesophageal nerve
ring highly variable: some or all passing through ring
(n = 3; figure 49), stopping just anterior to ring (n = 4),
or extending past without going through ring (n = 2).
Relati\ e lengths of sali\ ary glands vary. Anterior part of
esophagus, in cross-section (figures 26, 43), bearing 2
muscular, longitudinal, dorsal folds, their lower exten-
sions coiling upwards to form .semi-isolated compart-
ments, .interior esophagus thus divided into strongly cil-
iated dorsal food channel (figure 26, dfc), larger ventral
channel, and two small lateral pockets. Posterior to nerve
ring (figure 42), ventral projections into main channel
increasing gradualK in size, but without clear distinction
between anterior and mid-esophageal regions. Food
channel remaining dorsal. Dorsal folds and ventral pro-
jections gradually decreasing in size, with posterior
esophagus as a simple muscular tube.
Stomach (figures 12, 38-40) amber in color, slightly
translucent in li\'ing animal, approximately 4 x 1.5 mm
(in largest specimen, 11.5 mm shell diameter), lying on
exterior surface of visceral coil, appressed to the surface
of digestive gland just posterior to heart and kidney,
encompassing approximately Vi total length of digestive
gland. Stomach consisting of two continuous chambers
(figure 39) differing in function: anterior chamber ('/a of
total stomach length) containing gastric shield, working
end of cr\stalline style, and all openings into stomach;
remaining -/3 forming large sorting and storage chamber.
Esophagus entering stomach on left side at junction of
anterior and posterior chambers. From this point, series
of folds extending transversely across stomach, poste-
riori) into posterior chamber, and anteriorU toward in-
testinal opening. Opening to digestive gland lying to
right of esophageal opening, between it and gastric shield.
Gastric shield (figures 39, 40), with cup-like lateral wing
upon which crystalline style rotates, protruding into an-
terior chamber, and central longitudinal portion with
flattened lateral expansion that cradles style, positioning
it against cup-like grinding surface.
Style sac and intestine, at anterior end of stomach,
usualK partially obscured b\ connective tissue and kid-
ney. Style sac (figure 39, ss) narrow, finger-shaped, ap-
proximately '/3 length of stomach, not communicating
directly with intestine [Johansson's (1940:1) group 3, re-
vised after Mackintosh (1925)]. Style completely trans-
parent, rod-shaped (length 1.8 mm, diameter 0.35 mm),
rotating within style sac by action of denseK packed cilia
on style sac walls, protruding into anterior chamber of
stomach through flesh\ tube-like structure above gastric
shield. Channel extending betw een digestive gland open-
ing and intestinal opening at anterior terminus of anterior
chamber.
Figures 18-27. Cyclostremiscus beauii, specimens from Ft. Pierce Inlet. Florida (light micrographs or critical-point dried SEM
preparations). 18. Head-foot (male), with mantle edge reflected posteriorly, dorsal view (SEM). 19. Sagittal section of male through
buccal mass and esophageal region. 20. .\nterior view of snout and left cephalic tentacle, showing pattern of ciliated tracts on
ventral surface (SEM). 21. Tip of gill filament ^SEMl. 22. Osphradium, anterior to osphradial ganglion (SEM). 23. Osphradium,
cross-section through osphradial ganglion 24. Cephalic tentacle, cross-section. 25. X'entral tip of cephalic tentacle, showing tactile
pad (SEM). 26. Esophagus, cross-section through anterior section, with salivary glands. 27. Pallial tentacles: upper, finger-shaped
tentacle (retracted), and lower, paddle-shaped tentacle (SEM). Scale bars; 18 = 0.5 mm; 22 = 50 ^m; 19, 20, 23, 24, 26, 27 = 0.1
mm; 21, 25 = 20 ^m.
bm, buccal mass; bs, blood space; car, cartilage; cit. ciliar\ tract; cont. connective tissue; df, dorsal folds of esophagus; dfc, dorsal
food channel of esophagus; es, esophagus; leg, left cerebral ganglion; Ipt, lower pallial tentacle; me, mantle edge; ne, nerve; os,
osphradium; osg. osphradial ganglion; pe, penis; ra, radula; rs, receptaculum seminis; sag, salivary gland; sn, snout; tp, tactile pad;
upt, upper pallial tentacle.
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THE NAUTILUS, Vol. 102, No. 1
R. Bieler and P. M. Mikkelsen, 1988
Page 11
Posterior chamber (figures 39, 47, pch) with irregular
longitudinal folds along its left side leading from esoph-
ageal opening, and well-defined longitudinal groove on
its right side leading toward gastric shield. Central area
\\ ith series of ciliated transverse folds.
Proximal portion of intestine (figure 39, in) consisting
of 3 histologicalK separate sections: (1) slightly bulging
section immediately adjacent to stomach, containing
large typlilosole, leading into second section via small
loop; (2) very muscular, ball-shaped section passing un-
der style sac to join third section; and (3) moderately
ridged intestine proper, initially quite narrow, dilating
slightly as it passes, without further loops or undulations,
toward rectum in mantle cavity. Anus set back from
mantle edge. No special ciliated tracts from anus to man-
tle edge and exterior.
Observed flow of particles {figure 39, smalt arrows):
Large and small food particles enter stomach through
esophagus. Counter-clockwise whirlpool at esophageal
opening preliminariK' sorts particles according to size.
Smaller particles pass laterally to right in groove toward
gastric shield and style. Large particles move into pos-
terior chamber along left series of longitudinal folds,
passing to far posterior terminus of stomach. From there,
particles pass into central area where peristaltic action
manipulates and returns large particles to anterior cham-
ber. ConcurrentK . smaller particles separate and/or break
oft, and follow transverse folds toward right longitudinal
groove, and then anteriorly within groove toward gastric
shield. Large particles in central area of sorting chamber
continue moving anteriorly to region of esophageal open-
ing, where they pass rapidly by ciliary action directly to
intestinal opening. Small particles entering area of gastric
shield are manipulated by clockwise-rotating style against
cup-shaped flange of gastric shield. Resultant particles
move directly left into opening of digestive gland, guided
by cuticularized lateral folds near style sac opening. Un-
acceptable particles and material returning from diges-
tive gland are shunted via longitudinal groove toward
intestinal opening. Ball-shaped proximal section of in-
testine probably serves as a pellet compressor.
Live specimens fed on single-celled algae and detritus
scraped from laboratory- aquaria walls. Fecal pellets oval,
0.33 X 0.19 mm (n = 5), round in cross-section, with
rounded ends.
Renopericardial system: Two-chambered heart and
surrounding kidney visible on left surface of visceral coil,
posterior to ctenidium and hypobranchial gland (figure
12). Kidney large, with nephridial gland on its outer wall.
Kidney opening at posteriormost end of mantle cavity,
without conspicuous ciliated tract associated with open-
ing. No gonopericardial or renogonadial ducts observed.
Nervous system: (iircumesophageal ganglia (figures 42,
43, 49) moderately concentrated. RPG ratio [Davis et
ai, 1976:263; defined as length of pleurosupraesophageal
connective/(length of connective + length of right pleu-
ral ganglion + length of supraesophageal ganglion)] av-
eraged 0.49 (n = 7). Cerebral ganglia connected by nar-
row commissure, each separated from pleural ganglia by
constriction. Tentacular nerves with distinct swellings at
their bases. Pedal ganglia each with paired connectives,
connecting anteriorly to cerebral ganglia, posteriorly to
cerebropleural junctions. Pedal commissure very short,
narrow. Subesophageal ganglion somewhat smaller than
supraesophageal ganglion; connective to its pleural gan-
glion much shorter. Statocysts (figure 44, stc) about 110
)xm diameter. Buccal ganglia small, conspicuous, joined
by commissure passing beneath esophagus at posterior
end of buccal mass.
Highly-vacuoled connective tissue surrounding nerve
ring, as well as other organs and areas throughout body.
Reproductive system: .Animals in male phase (for dis-
cussion of sex change, see below) distinguished from
functional females by smaller size and by dark-orange,
rather than creamy-orange or beige, gonadial coloration.
Penis large (size at rest: length 2.5-3.0 mm, width at
midlength 0.4 mm), muscular, somewhat flattened (fig-
ures 12, 18, 42, 44) arising just behind and slightly right
of bases of cephalic tentacles, coiling counter-clockwise
back into mantle cavity. Fully extended penis may be
longer than cephalic tentacles {e.g., 4.4 mm vs. 3.0 mm).
Subcentral penial duct terminating in opening on slightly
hooded tip, which, unlike remainder of penis, is uncil-
iated. Long, closed pallial vas deferens extending from
penis, along right side of body, to prostate in posterior
part of mantle cavity (figure 44). Vas deferens relatively
wide (diameter ~ 0.2 mm), tubular, forming prominent,
somewhat undulating ridge that differs from surrounding
tissue by its shiny, unciliated surface and by its white
appearance caused by heavy internal ciliary action. Pros-
tate orange, egg-shaped, 0.65-0.90 x 0.35-0.48 mm (n =
5), lying ventral to rectum, connected to right pallial
vvall, its lumen communicating with mantle cavity by
slit (~ 0.35 mm length) at its base (figures 44, 48). Vis-
ceral vas deferens much narrower, passing through pos-
terior mantle wall, leading along inner coil of visceral
mass, where widened, extensively-coiled portion func-
tions as vesicular seminalis (figure 47, vs) before reaching
Figures 28-37. Cyclostremiscus beauii. radula and operculum, specimens from Ft. Pierce Inlet, Florida (SEM). 28. Outer marginal
teeth. 29. Radula, whole mount 30. Inner marginal teeth. 31. Rachidian teeth. 32. Lateral teeth. 33. Tips of lateral (left) and
inner marginal (right) teeth 34. Operculum, outer surface (2 1 mm diameter). 35. Operculum, inner surface (1.7 mm diameter).
36. Operculum, oblique lateral view of inner side (2 2 mm diameter). 37. Opercular peg, oblique lateral view. Scale bars: 28, 30-
33 = 10 Mm: 29, .37 = .50 fim.
rac, rachidian teeth.
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THE NAUTILUS, Vol. 102, No. 1
r\ipig ^^®9
stc sti
pispec es
PP9
mpg
R. Bieler and P. M. Mikkelsen, 1988
Page 13
testis. Testis simple, elongated sac, yellow to bright or-
ange, along inner right side of visceral coil, totalling 50-
60% of its length (figure 44).
Spermatozoon (56-62 nm total length, n = 3) with
slightly twisted head comprising elongated nucleus and
pointed acrosome (~ 14% of total length), very long,
narrow midpiece (« 58%) and long tail (* 28%). Atyp-
ical sperm not found.
Female reproductive tract (figures 13, 46) adjacent to
rectum along right side of mantle cavity. Female open-
ing, about level with anus, situated on muscular papilla
hanging freel\ in mantle ca\it\ at distal end of capsule
gland. From there, closed sperm duct leading posteriorly,
initially forming very thick-walled muscular vagina (fig-
ure 52), with lumen almost filled with large gland cells.
Sperm duct giving rise to small, ball-shaped, dorsally-
situated sperm pouch w ith weakK muscular walls. Thin-
walled ducts leading from sperm pouch and muscular
vagina joining shortly before opening into slit-like lumen
of massive capsule gland. \'agina-t>pe muscular wall
gradualK disappearing, while thin-walled part forms
sperm channel ("sperm groo\e " in "ventral channel of
authors), in communication with gland but partially sep-
arated by lateral fold (figure 46, cross-section B, If). An-
terior part of capsule gland w ith large, turquoise-staining
gland cells; remaining capsule gland staining dark blue
in sections. [M least in the anterior part, the gland cells
are arranged in a comple.\ pattern (see Ponder, in press).
A detailed histological description was hampered by the
extremely strong staining reaction of the glands.] Pos-
teriorly, glandular mass continuing, with communicating
lumina, as albumen gland, .\lbumen gland massive, thick-
walled, with narrow, slit-like lumen, pinkish-orange in
living animals, staining turquoise in sections; folded as
S-shaped loop on right side of animal, partK in parallel
with posterior part of capsule gland. Posteriormost part
of albumen gland pressing against, or, in large females,
extending beyond posterior mantle wall. Sperm channel
separating at junction of capsule and albumen glands,
forming closed o\iduct. penetrating posterior mantle wall,
and forming large, widened, non-glandular coil, con-
taining (from sections and dissected specimens) both
sperm and eggs. At its posterior end, coiled oviduct giving
rise to 2 proximal sperm pouches, as inconspicuous, sub-
equal, ball-shaped sacs, closeK' adjacent, and partialK'
hidden under la\ers of connective tissue and kidnev.
In ripe females, unoriented sperm, oltcn in large quan-
tities, found throughout length of sperm channel and in
anterior sperm pouch (functional bursa copulatrix).
Packed oriented sperm, with heads embedded in walls,
found in the two posterior pouches (receptacula seminis).
Visceral oviduct very thin-walled, leading from coiled
oviduct to ovary. Ovar\ situated at right side of visceral
mass, structure not grape-like, extending over central
80% of digestive gland, covering half to all ot right side
of coil (figure 46).
(During copulation, sperm are apparently deposited
in the \agina, temporariK stored in the bursa copulatrix
and then stored in the two posterior receptacula. Fertil-
ization occurs in the anterior part of the coiled oviduct,
after which fertilized eggs pass through the ciliated lu-
mina of the albumen and capsule glands where they are
surrounded by capsular and mucous material. Eggs ap-
parently enter the mantle cavity through the vagina, as
no other female opening was found.)
Sex and size relationships (figure 53): Specimens less
than 7.6 mm shell diameter (n = 25) were all males.
Specimens of shell diameter greater than 8.6 mm (n =
6) were all fully functional females without male repro-
ductive structures. Specimens 7.8-8.5 mm shell diameter
(n = 6) were "transitional" in appearance (figure 45);
visceral and pallial reproductive organs were clearly fe-
male, however, with visible remnant of pallial vas def-
erens and "wound" (= penis scar) at attachment site of
penis. Pallial vas deferens of smallest "transitional" spec-
imen present as unciliated duct, partialK opened at prox-
imal end, extending between penis scar and distal end
of capsule gland. No positive connections observed. This
specimen also with nearly transparent gonad containing
small amount of whitish ovarian tissue in early part of
coil, and with no sperm in the female system (it appar-
ently had not yet mated in its female phase). Largest
"transitional" specimen with fully-developed ovary, faint
longitudinal marking on integument in position of vas
deferens, and weak penis scar. Simultaneous possession
of both ovarian and testicular tissues not observed.
"Transition" somewhat correlated with date of col-
lection (figure 53). Collections made between March and
mid-August exclusively male. Fully developed females,
"transitional" individuals, and males encountered in each
of four collections in later part of August and September.
Figures 38-43. Cyclostremiscus beauii. 38. Left lateral view of visceral coil, showing position of stomach. Dashed line indicates
location of dorsal incision. 39. Stomach, dorsal view, opened at incision shown in figure .38. .\rrows inside stomach indicate flow^
of particles, 40. Gastric shield. 41. Right jaw, inner side, posterior end up. showing details of jaw elements. 42. Head with mid-
dorsal incision show ing buccal mass, circumesophageal nerve ring, and base of penis. Connective tissue and minor nerves removed.
43. Central nervous s\stem, left side, oblique lateral view. Scale bars: 39, 42 = 0.5 mm; 41, 43 = 0.1 mm
bg. buccal ganglion; bm, buccal mass; cpc, cerebropedal connective; cs, crystalline style; ct, ctenidium; dg, digestive gland; dgo,
opening of digestive gland; es, esophagus; eso. opening of esophagus; gon, gonad; gs, gastric shield; in. intestine; ino. opening of
intestine; ki. kidney; leg, left cerebral ganglion; Iplg. left pleural ganglion; Ipg. left pedal ganglion; mo, mouth; mpg. metapodial
ganglion; on, optic nerve; os, osphradium; pch. posterior chamber; peb, base of penis; pipe, pleuropedal connective; plspec,
pleurosupraesophageal connective; ppg, propodial ganglion; pvd, pallial vas deferens; ra, radula; ras, radular sac; rcg, right cerebral
ganglion; rplg, right pleural ganglion; sag, salivary gland; sbeg, subesophageal ganglion; speg, supraesophageal ganglion; ss, style
sac; st, stomach; stc, statocyst; stl, statolith; tns, swelling of tentacular nerve; ty, typhlosole.
Page 14
THE NAUTILUS, Vol. 102, No. 1
spc'^ spc
Figures 44-46. Cyclostremiscus beauii. reproductive system (semi-diagrammatic) 44. Male phase 45. "Transitional female
phase. 46. Female phase, with three cross-sections (A, B, C) through female glands. Small arrows indicate reflected organs
ag, albumen gland; an, anus; be. bursa copulatrix; eg, capsule gland; co, coiled oviduct; cont. connective tissue; ct, ctenidium; hg,
hypobranchial gland, If. lateral fold, ov, ovary; pe, penis; pr, prostate gland; prom, opening of prostate gland into mantle cavity;
R. Bieler and P. M. Mikkelsen, 1988
Page 15
Mating and larval development unknown. Protoconch
morphology suggests planktonic veliger stage (see Dis-
cussion). Sex change within single individual not ob-
served.
Habits and habitat: To our knowledge, this is the first
record of habitat type for this species, i.e., within the
burrows of thestoniatopod L(/sio.sq(n7/a scabricaiida, oth-
er published records (,see synonym) ) refer only to empty
shells. Wells et al. ( 1961 ) recorded Cyciostremiscus beatiii
from the stomachs of Astropecten articutatus (Say, 1825),
however it is unclear whether the material was alive
when swallowed b> the starfish. Another specimen from
gut contents of A. articutatus, in the UNC-IMS collec-
tions, from 24 m depth off North Carolina, contained
dried animal tissue and is the only other verified live-
collected specimen known to the authors. Although bur-
rows of other local species, e.g., callianassid shrimps,
polychaetes, hemichordates, and sipunculans, were also
sampled, these vitrinellids were not collected in associ-
ation with an\- burrower but Lijsiosquilla during this
stud\ .
The snails probabK feed on algae, bacterial films and
detritus in the burrows. They are capable of handling
larger items in their alimentary tract, as evidenced by
various shell pieces and foraminiferan shells found in the
stomach. From the absence of glandular esophageal
pouches and the presence of a crystalline style in the
stomach, it can be inferred (Yonge, 1930) that free pro-
teolytic enzymes, capable of digesting animal matter,
are not present in this species and would not be expected
in this group.
Individuals were almost invariably collected in groups
of more than one animal per burrow sample; the max-
imum number encountered in one burrow sample was
seven. Cyciostremiscus heauii was twice found crawling
openly on sand or seagrass in the vicinity of Lysiosquilla
burrow openings. In captivity, the animals were active
crawlers and were not distracted by light; dark, sheltered
areas in the tank (provided by black plastic film con-
tainers) were not preferred. During resting periods, all
specimens attached themselves to the wall of their con-
tainer, just above the water level, by means of highly
\ iscous mucus produced by the anterior pedal gland. This
behavior was not correlated with food availability or
water qualitv'. It might reflect tidal rhythm, but material
was not sufficient to test that hypothesis.
Geographical distribution: Western Atlantic, from
North Carolina to Brazil (Ceara and Alagoas; teste Rios,
1985:41). Recorded from the Pliocene of Jamaica and
Florida (see synonymy).
Taxonomic remarks: Fischer (1857b;286), in the orig-
inal description of Adeorbis heauii (in his earlier pub-
lication, the name occurred only as a nude list name;
1857a;173), did not give an indication of the number of
specimens in the original lot. Moore (1964T32) men-
tioned a "holotype" in the Paris Museum. However, the
single A. fceauii-specimen in that type collection (MNHN
unnumbered, vidi) is much smaller than the dimensions
given by Fischer (teleoconch diameter 6.3 mm, height
3.5 mm, protoconch diameter 0.42 mm, umbilical di-
ameter 1.1 mm; protoconch whorls 2, teleoconch whorls
2%; vs. 10.5 mm maximum shell diameter in the original
description). The specimen is therefore considered the
only remaining syntype of a formerly larger lot, and is
here selected as lectotype.
Woodring (1928:440) pointed out the clo.se resem-
blance between Guppy's (1866) Cyctostrerna bicarinata
and Adeorbis beauii Fischer, but separated the two be-
cause the latter "has a more strongly sculptured base."
Smith (1937:67) in turn separated his new species CAr-
culus stirophorus from both "Circuhis bicarinatus Gup-
py" and "Circuhis beauii Fischer, ' by differing numbers
of upper and basal spiral ribs. A re-examination of the
neotype (Woodring, 1928) of Cyclostrema bicarinata
(USNM 115621) and the original description and figure
of Circuhis stirophorus Smith revealed that both fall
within the range of variation (partly caused by ontoge-
netic change of basal sculpture) displayed by the Ft.
Pierce population, and both are here synonymized (see
also Moore, 1964:131). The holotype of Circuhis stiro-
phorus could not be located. Although the original author
indicated the type to be in the MCZ collection, it was
never received by that institution (K. J. Boss, personal
communication); it was also not located at the Florida
State Museum, Gainesville (F. G. Thompson, personal
communication).
Cyciostremiscus beauii is the largest vitrinellid in the
western Atlantic. Two Recent Panamic species are very
similar in size, shape and sculpture:
Cyciostremiscus major Olsson & Smith (1951:46, pi.
3, figs. la,b) from Panama differs from Cy. beauii in
having more numerous spiral ribs throughout and a more
rounded periphery (holotype ANSP 187199; figured by
Olsson & Smith, 1951).
Cyclostrema gordana Hertlein & Strong (1951:110, pi.
9, figs. 3, 4, 7; holotype CAS 064803, vidi) from the
Gulf of California, differs conchologically from Cycio-
stremiscus beauii in having a prominent double spiral
rib on the base between the basal keel and the umbilicus
(see Pilsbry, 1953: pi. 55). Hertlein and Strong (1951:
110) gave the following measurements for the "unique
type": maximum diameter 9.7 mm, minimum diameter
7.0 mm, and height 3.3 mm; our examination of the
holotype yielded, respectively, 8.6, 6.8, and 3.9 mm.
Hertlein and Strong (1951:110) compared C. gordana to
"Cyclostrema angulata A. Adams [1850] from the West
Indies ' [following "Pilsbry" (= error for Tryon), 1888:
ps, penis scar; pvd, pallia! vas deferens; pw, posterior pallial wall; re, rectum; rs, receptaculum seminis; sc, sperm channel; spc,
sperm channel; te. testis; va, vagina; vo, visceral oviduct; vs, part of visceral vas deferens serving as vesicula seminalis; vvd, visceral
vas deferens.
Page 16
THE NAUTILUS, Vol. 102, No. 1
Figures 47-52. Cycloslrcmiscus heauii, specinu-iis from Ft. Vwvcv Inlet, Florida (light micrographs or critical-point dried SEM
preparations). 47. Cross-section through \isceral ma.ss (male), at level of ve.siciila seminalis. 48. Section through prostate, showing
R. Bieler and P. M. Mikkelsen, 1988
Page 17
TW4-1
3-
n D
o
D
D
o o
o o
o
n15
10
5i
2 3 4 5 6 7 8 9 10 11 12
Imml
Figure 53. Cyclostremiscus beauii. Plot of maximum shell diameter vs. number of teleoconch whorls (TW). Inset: Histogram
summarizing sex distribution of collections, Ft. Pierce Inlet population. A = collecting period March-July, B = August 1-15, C =
August 16-31, D = September, 1987. Open circles = empty shells and unsexed specimens. Open squares = functional males. Cross-
hatched open squares = "transitional" females with pallial vas deferens and/or penis scar. Solid squares = females without male
structures. Star indicates lectot>pe specimen (MNHN unnumbered).
92, they considered C. angulata a senior synonym of
Cijclostremiscus beauii — see below] and stated that it
differed "principally in the smaller size and more de-
pressed form." However, a re-examination of the C. gor-
dana t%pe showed it to differ principaJK- b\' a strong,
second basal keel surrounding the umbilicus. It definitely
belongs in the genus Cyclostremiscus, as advocated by
Keen (1971),
Tryon (1888), followed by Hertlein and Strong (1951),
considered Cyclostremiscus beauii a synonym of Cy-
clostrema angulata A. Adams, 1850: "C. angulata, A.
Ad. was described as from the Philippines on the au-
thority of Cuming, but as that great collector sometimes
made mistakes, the localit> needs confirmation. There
can be no doubt of the identity with this species of C.
Beaui, Fischer (fig. 63), a West Indian species" (Tryon,
1888:92-93). Tryon 's figures (1888: pi. 32, figs. 64, 65)
may represent Cyclostremiscus beauii. However, as sim-
ilar species are known from the Indo-Pacific, we feel that
the two nominal species should not be synony mized with-
out a re-examination of Cyclostrema angulata type ma-
terial, which could not be located in the British Museum
(Natural History) (London) nor in the Redpath Museum
(Montreal).
The type species of Cyclostremiscus, Vitrinella pan-
amensis C. B. Adams, 1852, and several other, similar
species (see, e.g., Pilsbry & Olsson, 1945) have distinct
axial sculpture which is lacking in Cy. beauii, Cy. major,
Cy. gordana, and a number of other nominal species.
Pilsbry (1953) described the subgenus Ponocyclus in Cy-
lumen and opening into mantle cavity. 49. Section through circumesophageal nerve ring, with saUvary glands. 50. Spermatozoon
(SEM). Arrow indicates junction of niidpiece and tail. 51. Same, detail of two heads (SEM). Arrow indicates junction between head
and midpiece. 52. Cross-section through vagina (left) and rectum (right). Scale bars: 47-49, 52 = 0.1 mm; 50 = 4 ^m; 51 = 2 ^m.
cc, cerebral commissure; cont, connective tissue; ope, cerebropedal connective; dg, digestive gland; es, esophagus; hg, hypobranchial
gland; leg, left cerebral ganglion; Ipg, left pedal ganglion; mc, mantle cavity; mw, mantle wall; pc, pedal commissure; pch, posterior
chamber of stomach; pr, prostate gland; prom, opening of prostate gland into mantle cavity; rcg, right cerebral ganglion; re, rectum;
rpg, right pedal ganglion; sag, salivary gland; te, testis; tm, transverse muscle band; va, vagina; vs, part of visceral vas deferens
serving as vesicula seminalis; vvd, visceral vas deferens.
Page 18
THE NAUTILUS, Vol. 102, No. 1
R. Bieler and P. M. Mikkelsen, 1988
Page 19
clostremiscus \\ itli Adeorbis beauii as the type species,
and distinguished it from Cyclostreniiscu.s by its lack of
axial sculpture. As Pilsbr\ alread\ pointed out in his
description of Ponocijclus (1953:426). there are a number
of species showing intermediate conditions ("weak traces
of axial sculpture") and the name is not used here.
Due to the small number of species studied in the
\ itrinellidae, little can he said about anatomical char-
acters on the generic level. Aside from the large and
elaborately-sculptured shell, the extensive ciliation of the
head-foot, and the paddle-shaped pallial tentacle, Cy-
clostremiscus beauii differs anatomically from Circulus
striatus (see Fretter, 1956), Ci. texaniis (see below), and
Cochliolepis parasitica (see Moore, 1972) in ha\ing the
pallial portion of the intestine straight and uncoiled.
Circulus Jeffreys, 1865 (page 315)
Type species by monotypy: Delphinula duminiji Re-
quien. 1848 (page 64; not "Trochus duminyi Requien"
as usually cited) [= Yalvata? striata Philippi. 1836; see
Jeffre\s, 1865:317, where he used the junior synon> m D.
duminyi only to a\oid secondary homonymy with Tro-
chus striatus Linne, 1767].
Circulus texanus (Moore, 1965) new combination
(figures 54-68; tables 1, 2)
Vitrinella texana Moore, 1964:66 [unpublished dissertation).
Vitrinella texana Moore, 1965:76, pi. 7. figs. 4-6 [Gulf of
Mexico].
? Vitrinella texana "Moore, 1964" — .■Andrews, 1971:73-74, text-
fig. — Andrews, 1977:887 [citing "1965"], text-fig. [poor fig-
ure; = Vitrinella sp., teste Ode, 1987.35].
Vitrinella (Vitrinella) feiana,— Abbott, 1974:83.
Vitridomus texana. — Ode, 1987:37.
Material examined: Ho]ot\pe: empt\ shell, 1.8 mm,
USNM 636311 (Texas). Other material: 15 specimens.
FLORIDA: Ft. Pierce Inlet: 2-3 May 1987, 1 unsexed;
24 June 1987, 14 specimens (4 males, 7 females, 2 un-
sexed, 1 empty shell).
Description
Teleoconch (figures 54-56): Shell small (1.7-1.8 mm
diameter, 0.55-0.65 mm height), with 1V2-1% teleoconch
w horls; almost planispiral, sculptured dorsalK' and ven-
trally with about 18 fine spiral ribs; transparent when
alive, opaque after death. Ribs slightly stronger, more
widely spaced just below suture on dorsum and at pe-
riphery, where about 3 ribs form rounded keel below
lateral midline. Suture impressed. Ventral surface below
keel less convex, often with 30-40 widely-spaced, low
axial ribs which are primarily evident from inside of
body whorl (figure 57). L'mbilicus wide (~ 25% of width).
Outer lip ver>- slightly reflected; some specimens with
one former varix. Aperture at oblique angle to dorso-
ventral axis. Sutural sinus shallow. Periostracum thin,
transparent, with spiral grooves more numerous than on
shell surface.
Protoeonch (figure 58): Smooth, 0.5 mm diameter,
about 2 whorls. No sculptural demarcation separating
protoeonch I and protoeonch II.
External anatomy and organs of the mantle cavity (fig-
ures 64-66): Living animal translucent white with
buccal mass (visible through proboscis) and protrusible
portion of ctenidium rose-pink; digestive gland orange;
ner\e ring area opaque yellow, visible through integu-
ment between eyes. Head with long, extensible snout,
rounded and terminally notched at mouth. Ventral side
of snout ciliated behind mouth opening, with cilia beat-
ing toward mouth. Two long cephalic tentacles w ith im-
mobile bristles on slight]} enlarged tip; motile cilia at
least at tip and along side facing snout. Left cephalic
tentacle fitting into shallow notch formed by shell keel
(figure 65). Eyes black, on slight bulges at base of cephalic
tentacles. Neck region very long, slender. Foot elongate,
flattened, with anterolaterally recurved corners; fineK
ciliated sole broadly rounded posteriorly, extending well
beyond shell (figure 65). Transverse crease on sole about
'/^ of total length from anterior edge. Anterior pedal
mucous gland (figure 65, amg) opening at slit across
entire leading edge; mid- ventral fold evident on posterior
quarter of sole, but presence of posterior mucous gland
not confirmed. Locomotion by ci]iar\ action. Operculum
(figures 61-63) corneous, circular, multispiral {ca. 8
whorls), with small central peg and rounded-triangular
muscle scar on inner surface. Operculum supported by
opercular lobes (figure 65, ol) on dorsal side of foot. Lobes
simple, unpigmented, without tentacles. Epipodial ten-
tacles absent.
Two finger-shaped pallial tentacles (figure 64, Ipt, upt)
arising separately but adjacently from inner mantle edge,
at right side of animal. Upper (= most dorsal) tentacle
unciliated, with immobile terminal bristles, curling into
shallow sutural sinus, directed dorsalK during crawling.
Lower tentacle bearing motile cilia, directed anterolat-
erally. Ctenidium (figures 64, 65, ct) with numerous fin-
ger-shaped filaments, attached for most of its length to
internal surface of mantle; anteriormost filaments darker
Figures 54-63. Circulus texanus, specimens from Ft. Pierce Inlet, Florida (SEM) (figures 54-56, USNM 846324). 54. Shell, apical
view (2.7 mm diameter). 55. Shell, umbilical view (2.7 mm diameter). 56. Shell, apertural view (2.2 mm diameter). 57. Inside
surface of base of bod> whorl, showing shallow radial grooves. 58. Protoeonch. 59. Larval shell, left lateral view (167 /im max.
diameter! 60. Radula. 61. Operculum, inner surface (0.7 mm diameter) 62. Operculum, outer surface (0.8 mm diameter). 63.
Operculum, oblique view of inner surface with opercular peg. Scale bars: 57, 58 = 0.2 mm: 60 = 5 ^m; 63 = 50 ^m.
rac, rachidian teeth.
Page 20
THE NAUTILUS, Vol. 102, No. 1
Figures 64-68. Circulus texanus. 64. Crawling female, dorsal view, drawn as with transparent shell. 65. Crawling animal, ventral
view, drawn as with transparent shell. 66. Head of male, left lateral view, showing penis. 67. Egg mass 68. Two egg capsules,
each with veliger larva shortly before hatching. Scale bars: 64, 65 = 0.5 mm; 66, 67 = 0.25 mm.
ag, albumen gland; amg, anterior mucous gland; br, immobile bristles; eg, capsule gland; ct, ctenidium; dg. digestive gland, ebv,
efferent branchial vessel; ey, eye; he, heart; in, intestine; ki, kidney; Ipt, lower pallial tentacle; ol, opercular lobe; op, operculum;
ov, ovary; pe, penis; re. rectum; sh, shell; sn, snout; ss, style sac; st. stomach; ten, cephalic tentacle; umb, umbilicus; upt, upper
pallial tentacle.
in color, not attaclu'd to tnantle skirt, extending out ot
aperture at right side of head. E.xtent of osphradium not
ascertained.
Alimentary system: Riiduia (figure 60) taenioglossate.
Rachidian tooth wider than long, with acute posterior
corners projecting laterally and concave front. Main cusp
narrow, unserrated, with 5-6 slightly smaller flanking
cusps on each side (number of flanking cusps var\ ing
within single radular rihhon) decreasing laterally in size;
base with 1 weak elevation per side (in position of basal
denticle), midway between posterior corners and central
ridge. Lateral tooth with asymmetrical cutting edge,
strongly indented at front edge, with narrow , unserrated
main cusp and subequal flanking cusps (6-7 inner, 9
outer). Apex of inner marginal tooth with numerous long,
thin, flanking cusps on either side of similarly-sized main
cusp. Apex of outer marginal tooth with long, thin inner
flanking cusps, slightly more robust than tho.se of inner
marginal tooth; cusps at termiiuis subequal in size; outer
margin with 2-3 cusps decreasing in size, remainder of
outer margin smoothly rounded. Jaws composed of ele-
ments approximately 4 ^m in length.
Stomach traii.spinciit, with rotating style often clearly
visible through shell. Posterior part of stomach not mark-
edly elongated. Intestine (figures 64, 65, in) forming wide
loop in posterior part of mantle cavity. Anus just above
anterior end ol capsule gland in females. Fecal material
R. Bieler and P. M. Mikkelsen, 1988
Page 21
as continuous rods of irregular length, rather than pellets,
with no obvious surface sculpture.
Reproductive system: Penis ifigure 66, pe) a posteriorly
directed, counterclockw ise, double coil; base arising pos-
terior to eyes, somew hat right of dorsal midline. Testis
orange.
Ovary (figure 64, ov) filling inner side of early whorls
with grape-like lobes, lighter in color than digestive gland.
Capsule gland and much darker-staining (in metln lene
blue) albumen gland (figure 64, eg, ag) forming compact
mass on right wall of mantle cavity, below rectum, ter-
minating anteriorly at level of anus. Position and number
of sperm pouches not ascertained.
No evidence of se.xual dimorphism associated with
possible sequential hermaphroditism. Males both consid-
erably smaller (1.7 mm diameter at 1% whorls) or larger
(1.8 mm diameter at 1': whorls) than females in same
sample (1. 74-1. 78 mm at I'/s whorls). One copulation
briefly observed, wherein male paused while crawling
over dorsal surface of female, with heads adjacent (male
at left) and at angle of about 30°. Position of penis not
ascertained.
Eggs and larval development: Three weeks after col-
lecting, 7 females and 4 males were placed into glass
bowl with seawater (24 °C). Overnight, 15 egg masses
were laid; each mass contained 1-5 eggs, each in clear,
flattened, circular capsule (egg diameter 1 10 fim: capsule
diameter 170 ^m; n = 20). All eggs found in same de-
velopmental stage, uncleaved, completely round, finely
granular. Soft capsule walls of eggs laid in groups touch-
ing each other; each "mass," even if consisting of single
egg, coated b\ sticky, clear, gelatinous layer (figure 67).
Eighteen additional egg masses were found in glass jar
in which specimens had been kept since collecting. Total
number of masses (with respective numbers of single-
egg-capsules) was: 4 masses with 1 capsule, 4 masses with
2, 13 masses with 3, 9 masses with 4, and 3 masses with
5 capsules.
One mass with 5 eggs followed through development:
From beginning of equal, holoblastic 2-celled stage, 60
minutes elapsed to 4-celled stage, 205 minutes to 8-celled
stage, 385 minutes to 16-celled stage, respectively. Gas-
trula-stage discernible after 1 1 hours; embryos rotating
after 17 hours.
X'eliger larvae (figure 68) fully developed at 104 hours,
intermittently or constantly rotating, with transparent
shell, 2 short veliger lobes, black eye spots (not situated
on tentacles at this point), statocysts and operculum [much
as described for Caecum glabrum by Gotze (1938:108,
text-fig. 33), but with smaller vela]. Periphery of velar
lobes bearing long cilia, underlain by row of shorter cilia.
Capsule size now approximately 150 x 120 fim, greatest
shell diameter 107 /xm, longest dimension of veliger with
extended \ela 133 ^m. During entire process, 3 of 5
embryos developed completely synchronously; other 2
initially about 45 minutes, later several hours, behind.
First veligers hatched after 130 hours and swam actively.
For follow ing 36 hours, veligers observed either swim-
ming or resting on bottom of bowl, with strong ciliary
action on extended vela. No further development noted.
Added cultures of mixed single-celled green algae ap-
parently not excepted; all larvae died within next 2 days.
[From the size difterence between these larvae (170 nm)
and the protoconchs of adult shells (500 nm), a longer
lar\al stage, w ith extensive food intake can be predicted.]
Habits and habitat: Habitat as described for Cyclo-
stremiscu.s hcaiiii. In the laboratory, animals of Circiilns
texanus were very active and fed on normally-occurring
algal and bacterial surface films.
Geographical distribution: Circuhis texanus (type lo-
cality: Mustang Island, near Port Aransas) was previously
reported to be endemic to the northv\estern Gulf of Mex-
ico, and has never been reported alive (Moore, 1964:41;
.\ndrews, 1977:87; Ode, 1987:37). No fossil records are
know n.
Taxonomic remarks: Moore (1965:77) placed this species
in the genus Vitrinclla C. B. Adams, 1850, and pointed
out the differences in shell shape and sculpture in com-
parison to other species. However, members of Vithnella
(based on the type species V. helicoidea C. B. Adams,
1850; see tables 1, 2) are generally smooth-shelled or
weakly sculptured and are usually furnished with a ca-
rina bordering the umbilicus (e.g.. Keen. 1971:377; Ab-
bott, 1974:82). We feel that this species is better placed
in Circuhis sensu lato because it agrees in shell shape
and sculpture with the type species, Circuhis striatus
from the eastern Atlantic. Ode's (1987:37) placement of
this species in VHridomus Pilsbry & Olsson, 1945, cannot
be accepted: members of Vitridomus [a "rather feebly
defined genus (Pilsbry & Olsson, 1952:36), very similar
to Teinostoma H. & A. Adams, 1853] ha\ e the umbilicus
partly covered by a callus.
DISCUSSION
Comparison of the anatomical information on Cyclo-
stremiscus beauii and Circuhis texanus with literature
data revealed that published information is scarce or
misleading, and that genus- as well as family-level taxa
of marine near-planispiral Rissoacea are poorly defined.
Three nominal families, Vitrinellidae Bush, 1897, Tor-
nidae Sacco, 1896, and Circulidae (for authorship, see
below), are currently in use for members of this group,
and the lack of comparative data led Ponder (in press)
to treat all three families as a single group in a phylo-
genetic analysis of the Rissoacea. In the follow ing dis-
cussion, we (a) review the available data on species of
this complex, (b) compare Cy. beauii and Ci. texanus
w ith these data and with other members of the Rissoacea,
(c) suggest a preliminary grouping for the species for
w hich anatomical data are available, (d) demonstrate the
differences between Vitrinellidae and Tornidae, (e) eval-
uate the taxonomic status of the nominal family Circu-
lidae, and (f ) discuss some unusual aspects of the anatomy
and reproductive biology of Cy. beauii. namely ciliation.
Page 22
THE NAUTILUS, Vol. 102, No. 1
tentacle shape, stomach morphology, and hermaphro-
ditism.
(a) Available literature data (see tables I, 2).
Except for Fretter's (1956) study of Circulus striatus,
and Woodward's (189S) and Graham's (1982) work on
Tornus suhcarinatus, most of the available data on the
anatom\ of marine near-planispiral Rissoacea are re-
stricted to descriptions or illustrations of external mor-
phology, especially Pilsbr\ and McGinty's (1945a, 1946b)
sketches of western .\tlantic forms. Some of the few
published observations on the radular structure and gross
morphology of vitrinellid softbodies are misleading (see
also Moore, 1964, 1972). Based on the misconception that
these species are archaeogastropods. Rush (1897:127, 142,
pi. 22, figs. 12a-g) erroneously constructed a rhipido-
glossate radula for Circulus trilix (Bush, 1885) [= Cy-
clostremiscus pentagonus (Gahh, \8S7), fide Moore, 1964;
138]. The "supplementar\ plicated gill" (Stimpson, 1858:
308) of Cochliolepis parasitica uas not found in that
species by Moore (1972:101). Pilsbry (1953:427) men-
tioned a taenioglossate radula for Cyclostremiscus beauii,
and added "This will be figured in a paper now in prep-
aration b\ T.L. McGinty and the author " This work was
apparentl) never published.
(b) Comparison of Cyclostremiscus beauii and Circulus
texanus with other \itrinellids and rissoaceans.
Both Cyclostremiscus beauii and Circulus texanus dis-
play "typical" rissoacean characters (as outlined by Fret-
ter & Graham, 1978:153; Boss, 1982:984; Ponder, 1983,
in press) and agree to a large extent with the previous
concept of Vitrinellidae (Fretter, 1956; Moore, 1972; see
tables 1, 2, Group 1). As in most other vitrinellid species,
the cephalic and pallial tentacles bear immobile bristles
and/or motile cilia. Tracts of motile cilia on the cephalic
tentacles occur in numerous groups in the Rissoacea; for
lack of comparative data, it is not yet clear whether the
arrangement of these cilia in grooves, as in Cy. beauii,
is unusual. Photographs of critical-point dried tentacles
of a freshwater rissoacean, the hydrobiid Tryonia ctath-
rata Stimpson, 1865 (see Hershler & Thompson, 1987:
27, figs. 13-17), show no grooves present in that species.
Cyclostremiscus beauii and Circulus texanus have two
closely-spaced pallia! tentacles on the right side. This
arrangement is present in most studied vitrinellids (the
exception being Cochliolepis albiceratus Ponder, 1966),
in the type species of Tornus (Tornidae, see below), in
Hydrococcus brazieri (T. Woods, 1876) (monotypic Hy-
drococcidae, see Ponder, 1982), and in the rissoid genus
Rissoina Orbigny, 1840, where some species have either
the anterior (right) or posterior (left) pallial tentacle bi-
lobed (Ponder, 1985:78). As described for Cy. beauii, the
function of the upper tentacle, which usually bears im-
mobile bristles, and is held upright and bent around the
aperture, is clearly sensory, while the lower one, often
distinctly ciliated and sometimes broadened, controls and/
or enhances water flow.
Both species have a small central opercular peg which
is probably not homologous with the lateral "neritid-
type " peg in other families of Rissoacea (character con-
sidered plesiomorphic in this superfamily), while the cir-
cular shape of the vitrinellid operculum is considered
derived (Ponder, 1985:5).
The osphradium of Cyclostremiscus beauii is ver) sim-
ilar to those described for Circulus striatus (see Fretter,
1956:372) and for species of the Rissoidae (Johansson,
1939:319, pi. 3, figs. 3, 4, text-fig. 5; Haszprunar. 1985:
476, figs. 7k, 16).
The radulae of the two species studied here ha\ e only
one pair of basal denticles (or elevations) on the rachidian
tooth, a feature known from some other rissoacean fam-
ilies, such as Hydrobiidae (see Bandel, 1984:29, text-fig.
47) and Rissoidae (see Ponder, 1985:10). This character
was hvpothesized to be primitive in the Rissoacea by
Ponder (1985:119).
The position of the salivary glands relative to the cir-
cumcsophageal nerve ring is often used in family- and
superfamiK -level discussions (e.g., Ponder, 1983:236, 258;
in press). However, as a highly variable character in the
populations of Cyclostremiscus beauii studied herein,
further data on additional species are necessarv before
it can be reliabl) utilized to infer phylogenetic relation-
ships of the Vitrinellidae.
The extensive anterior section of the esophagus of Cy-
clostremiscus beauii, with its long, coiled, dorsal folds,
is verv similar to that of Hydrococcus brazieri (see Pon-
der, 1982:77).
The conspicuous ciliated tract leading from the renal
opening to the head of Circulus striatus (see Fretter,
1956:372) is lacking in Cyclostremiscus beauii.
The presence of markedly vacuoled connective tissue
in Cyclostremiscus beauii is a character shared by Cir-
culus striatus and Tornus subcarinatus (see Graham,
1982:147).
The nervous system of vitrinellid species is tvpically
rissoacean and shows moderate (Cyclostremiscus beauii)
to high (Circulus striatus; Fretter, 1956:377) concentra-
tion. The RPG ratio of Cy. beauii, averaging 0.49. lies
within the range of those of members of the rissoacean
family Pomatiopsidae (see, e.g., Davis ii Mazurkiewicz,
1985:45, table 8).
The structure of the male and female reproductive
systems encountered during this stud\ is ris,soacean (e.g.,
Johansson, 1956). As in most other rissoaceans (e.g.. Pon-
der, 1985:6), the albumen and capsule glands form a
single, continuous mass [a character not recognizable from
Fretter's (1956) description and figures of Circulus stria-
tus]. The N'itrinellidae differ from the Rissoidae (e.g..
Ponder, 1985: text-fig. 2) wherein a non-homologous
structure, an expanded part of the upper oviduct, pos-
terior to the bursa copulatrix and/or receptacula seminis,
functions as an albumen gland. This "upper oviduct
gland " of the Rissoidae is homologous with the coiled
part of the visceral oviducts of the Hvdrobiidae (see
Johansson, 1956) and N'itrinellidae [Fretter's (1956) "ren-
al oviduct '].
Most members of the Rissoacea have one bursa cop-
ulatrix and one receptaculum seminis near the posterior
R. Bieler and P. M. Mikkelsen, 1988
Page 23
pallial \\ all, ifquii iiig the sperm to travel the entire length
of the sperm ehannel (vaginal lumen, sperm groove of
the ventral channel of authors) immediately after cop-
ulation. Others have develeoped additional, ilistal sperm
pouches (anterior sperm-storage structure, distal l)lind
sac, sac-hke vestibule, spermatheca, pseudo-bursa of au-
thors), eitlier in addition to [e.g., Ptisillina incon.^pictia
(Alder, 1844); see Johansson, 1939:337. text-fig. 22 (as
Rissoa); Rissoidae], or instead of the pro.ximal bursa cop-
ulatrix [e.g., Htjala vitrea (Montagu, 1803); see Johans-
son, 1949: text-fig. 1; Iravadiidae]. These distal sperm
pouches are not necessarily homologous with each other
and certainly not with the proximal bursa (see, e.g., Sla-
voshevskaya, 1978). Vitrinellids for which such data are
available (CArciilus striatus and Cyclostremiscus beauii)
differ from most other rissoaceans in the presence of two
more-or-less eciually developed, proximal receptacula
seminis which, in position and size, look much like the
two sperm sacs {i.e., bursa copulatrix and receptaculum
seminis) of other Rissoacea {e.g., in the rissoid Lucidestea
Laseron, 1956; see Ponder, 1985:67, text-fig. 3). Whether
the presence of two proximal receptacula seminis is a
distinguishing character for Vitrinellidae cannot yet be
determined, as accessory receptacula have been dem-
onstrated for members of the genus Alvania in the Ris-
soidae (Johansson, 1956; Ponder, 1985). In both Circiilus
striatus (see Fretter, 1956:377) and Cyclostremiscus
beauii, the distal end of the coiled oviduct serves as the
fertilization area, and not as an additional functional
receptaculum as has been reported for some other ris-
soaceans [e.g., Alvania suhsoluta (Aradas, 1847), where
the distal end contained oriented sperm; Johansson, 1956:
380].
The spawn mass of members of the Rissoidae usually
has numerous eggs per capsule in planktonic forms, while
one-egg-per-capsule is characteristic of "direct ' devel-
opment (see Lebour, 1937; Thorson, 1946). This does not
hold true for the spawn of Circiilus texanus, which gen-
erally resembles that of the freshwater rissoacean Bithijn-
ia tentaculata (Linne, 1758), as described and illustrated
by Ankel (1936:164, text-fig. 142B). [The number of eggs
there, however, is larger (4-24; Lilly, 1953:104), and
hatching occurs at the crawling stage]. Jablonski and Lutz
(1980:336; after Taylor, 1975) stated that the Rissoacea
". . . follow one of two developmental pathways: those
that hatch as crawling juveniles from relatively large eggs
(140-320 nm) and those that hatch from relatively small
eggs (60-130 jxm) and spend 2-3 weeks as planktonic
veligers. Both species studied here definitely fall into
the latter category: veliger shells of Circulus texanus
(figure 59) were one full whorl smaller than the final
larval shells as seen in the adult protoconch (figure 58);
the protoconch of Cyclostremiscus beauii (figures 9-11)
shows a distinct line and change of sculpture between
protoconch I and protoconch II, with almost another full
whorl of growth before metamorphosis, suggesting sev-
eral weeks (K. Bandel, personal communication) of
planktonic life. In his unpublished revision of western
Atlantic Vitrinellidae, Moore (1964:18) inferred that
"most, if not all, species appear to have a planktonic
veliger stage of some duration."
The sperm cells of Cyclostremiscus beauii are of the
general type known tor other members of Rissoacea (see
Gotze, 1938; Franzen, 1955). The twisted acrosome and
relative lengths of head, midpiece and tail are similar to
those described by Franzen (1955) for Caecum glabrum
(Montagu, 1803). The head is much shorter than that of
the two rissoids previously studied [Pusillina incon.spicua
(Alder, 1844) (as Rissoa) and Onuba striata (J. Adams,
1797); Franzen, 1955:406-409], and the relative length
of the midpiece much greater than in Hydrolna ulvae
(Pennant, 1777).
Direct communication of the coiled ("renal") oviduct
with the kidney (Fretter, 1956), previously considered
unique for Circulus or the Vitrinellidae, is now also known
for other families of the Rissoacea (Tornidae, Graham,
1982; Truncatellidae, Fretter & Graham, 1962).
Anatomical characters of Circulus striatus, which
Fretter (1956:380) discussed as probably "associated with
small size and body form," are equally expressed in the
much larger Cyclostremiscus beauii, suggesting that
Fretter's hypothesis was incorrect.
(c) Preliminary grouping of "vitrinellid-like" species for
which anatomical data are available.
The species for which sufficient gross morphological
data are available appear to fall into three groups, two
of which are here considered of familial rank and one
comprising species of incertae sedis (tables 1, 2). Mem-
bers of Group 1, currently placed in the genera Vitri-
nella, Teinostoma, Pieurornalaxis, Cochliolepis, Circu-
lus and Cyclostremiscus, are here considered to belong
to the Vitrinellidae (the placement of the species de-
scribed as Cochliolepis albicerata Ponder, 1966, is some-
what doubtful as it has only one pallial tentacle, and the
morphologies of its osphradium and penis are unknown).
Some of the characters attributed to the family Vitri-
nellidae in Moore's dissertation (1964) were based on
Parviturboides interruptus (C. B. Adams, 1850) (tables
1, 2, Group 2), later considered a species of uncertain
systematic position (Moore, 1972:107). This species dif-
fers from the other forms here grouped in Vitrinellidae
by the following anatomical characters: posterior foot
margin with immobile cilia, left cephalic tentacle with
four low swellings on proximal posterior border, penis
with glandular area and directed straight back. Another
species, Tomura bicaudata (Pilsbry & McGinty, 1946)
differs in many head-foot characters (tables 1, 2, Group
2) and is here also considered as incertae sedis. White
(1942:92) advocated the inclusion of "Cyclostrema" bushi
Dautzenberg & Fischer, 1907, in the Vitrinellidae/Tor-
nidae complex and published anatomical descriptions
and illustrations of that species. While some morpho-
logical characters [head with well-developed eyes and
terminally ciliated cephalic tentacles, the well-developed
osphradium (1942: text-fig. 6), and the circular, multi-
spiral operculum (1942: pi. 2, fig. 3)] agree well with the
species listed here as vitrinellids, the glandular pouches
Page 24
THE NAUTILUS, Vol. 102, No. 1
in the esophagus and, most of all, the pair of epipodial
(not pallial) tentacles on the right side of the animal
(White, 1942:90, text-fig. 5), prevent inclusion in \itri-
nellidae. Tornm siibcarinatus (tables 1, 2, Group 3) is
here considered a member of a separate famiK (see be-
low).
(d) Family relationships: Vitrinellidae-Tornidae.
Several attempts have been made to define the key
characters of the family Vitrinellidae [e.g., Moore, 1965:
74, 1969:170, 1972:107ff.; Boss, 1982:991). However, be-
cause of the small number of species fulK studied ana-
tomicall), no single synapomorphy defining the famiK
is clear at this time. The most obvious features of the
Vitrinellidae are: a low-spired, translucent white shell;
long cephalic tentacles equipped with terminal bristles
and, in most cases, motile cilia; two closely-spaced pallial
tentacles on the right side; a large monopectinate gill
often projecting to the right of the head; a large linear
osphradium; a foot with simple or only slightly indented
anterior and posterior margins; a horn\ concentric, mul-
tispiral operculum; and the possession of a non-glandular
curved penis in the male. Some of these features (shell
shape, pallial tentacles, and projecting gill) have led au-
thors (e.g., Ta\ior & Sohl, 1962) to s\nonymize Vitri-
nellidae and Tornidae, based on Woodward s (1898) work
on Adeorbis [= Tormts] .siibcarinatus (Montagu, 1903).
Others (e.g.. .\dam & Knudsen, 1969; Moore, 1972) have
pointed out differences between vitrinellids and Tornus/
toniids in features of the e\ es (functional eyes are lacking
in T. .'iubcarinatiis ) and, or the operculum (oval and pau-
cispiral in T. siibcarinatus) and have separated the two
families. In a redescription of the anatomy of T. sub-
carinatus, Graham (1982:147) saw the opercular shape
as the "single difference between the two nominal fam-
ilies and again advocated synonymy.
Additional anatomical characters of Tornidae that
warrant separation from Vitrinellidae are: (1) the con-
spicuous, elongate osphradium of the latter is not present
in Tornus (Graham, 1982:144, found only "a small cil-
iated groove, which may be a reduced osphradium"), (2)
the attachment of the ctenidial axis to the mantle is short
and the axis hardly supplied with blood vessels, and (3)
the penis of 7". siibcarinatus bears several finger-like
processes which are not known in vitrinellids as delimited
here. [Fretter & Graham (1978:231) described the cte-
nidium of T. siibcarinatus as "partly bipectinate." This
is apparentK in error, as both Woodward (1898) and
Graham (1982) described and illustrated that gill with
only a single row of lamellae.] As T. siibcarinatus lives
in a comparable habitat (under boulders on well-oxy-
genated sand or iiuid) and is of about the same size as
Circulits spp.. the drastic difference in osphradial type
indicates phylogenetic difference rather than specializa-
tion on a low taxonomic level.
Ponder (in press) combined the "tornicl-\ itrinellid-cir-
culid complex" as lamiK 'i'ornidae tor the purpose of
his pin logenetic analysis of the Rissoacea. However, most
of the characters and character states he attributed to
that complex (metapodial tentacle present, esophageal
pouches and glands present, penial glands present, os-
phradium short, posterior end ot foot not simple) do not
occur in N'itrinellidae as understood here, and must refer
to Tornus and other tornid genera studied b> Ponder
(Pseudoliotia Tate, 1898, Scrupus Finlay, 1927; unpub-
lished).
Vitrinellidae and Tornidae fit well into the Rissoacea,
and an independent grouping of these famiilies as Tor-
nacea (e.g., Kuroda et al., 1971; Golikov & Starobogatov,
1975) is not justified. The family Adeorbidae Montero-
sato, 1884, used b\- some authors for members of this
complex, is a s\non> m of Tornidae, since Adeorbis S.
Wood, 1842, is an objective synonym of Tornus Turton
& Kingston, 1830 (see Iredale, 1914:172, 1915:344).
(e) The nominal family Circulidae.
The taxonomic status of the nominal family Circulidae
remains problematic. The family name "Circulidae" was
first used b> Fretter and Graham (1962:642. ".\ppendix
I"), in a list of taxa treated in that publication. The text
reads merely "Circulidae: Circulus striatus (Philippi). '
Fretter and Graham (1962) did not state that they in-
tended to create a new family, did not mention the name
in the main bod\ of the text, even u hen the taxonomic
position of the genus (1962:550, 618) and the composition
of the superfamiK (1962: 622-623) were discussed, and
in fact ne\er used any famiK name but N'itrinellidae
when the\ referred to Ci. striatus in subsequent publi-
cations (Fretter bi Graham, 1978:227; Graham, 1982:
147). Lacking an\ description, definition, or bibliograph-
ic reference, a name thus introduced is not taxonomically
available (ICZN, 1985: Art. 13). "Circulidae" could
therefore be regarded as a nomen nudum.
However, Golikov and Starobogatov (1975:211) ac-
cepted "Circulidae Fretter & Graham, 1962" as a valid
family and stated (1975:218) that "the characteristics of
the latter famiK are found in Fretter (1956:381)," there-
by referring to the summar\ of Fretter s description of
Ci. striatus. This fulfills the requirement of ICZN (1985)
Art. 13(a)ii (Bibliographic Reference to Published State-
ment) and, unless there are earlier such statements that
have escaped us, this makes Goliko\ and Starobogatov
(1975) the authors of Circulidae, with Circulus as the
name-bearing t\ pe. The nominal family Circulidae is,
with anatomical descriptions available for Ci. striatus
and Ci. texanus, much better detined than the X'itrinel-
lidae, where our knowledge ot Vitrinella is based onK
on shell characters and Pilsbry and McGinty's (1945a,
1946b) sketches of crawling animals (tables 1. 2). .\11
available data suggest s>non\in\ of N'itrinellidae and
Circulidae (tables 1, 2; Boss, 1982:991 ). In any case, the
placement of N'itrinellidae and Circulidae in separate
superfamilies, or even separate superorders (Golikov &
Starobogatov, 1975), is highly exaggerated.
(f) Unusual features of Cyclostremiscus beauii.
The stud) of ('yclustrentisciis beauii revealed a num-
ber of features that differ from other members of the
N'itrinellidae (and, in part, from the Rissoacea). Most of
R. Bieler and P. M. Mikkelsen, 1988
Page 25
these features may be related to either its unusually large
size (for a \ itrinellid) or its unusual habitat in stoniatopod
burrows.
Ciliation and tentacle shape: The extensive ciliation of
the gills, cephalic tentacles and lower pallial tentacle
provides eflective respirator) and excretory currents. On
the right side of the head, where in- and outgoing cur-
rents are present, water flow is controlled by the paddle-
shaped tentacle. Low tides and high water temperatures
are likely tc create anoxic conditions in the burrows, and
a large vitrinellid such as Cyclontremiscus heauii may
be more strongly affected by oxygen deficiencies than,
for instance, the smaller Circuhis texanus. and thus would
benefit from an elaborate s\stem to produce and direct
currents. The onl\ other known commensal \itrinellid,
Cochliolepis parasitica, is much smaller, and lives di-
rectly on its host [under the scales of the giant scale worm
Pohjodontcs lupina (Stimpson, 1856); see Stimpson, 1858;
llartman, 1945:10]. Moore (1972:104) did not find cilia
on the gill filaments of Cochliolepis parasitica, suggesting
tliat currents produced by the annelid are sufficient to
supply the snail. All other species studied live under rocks
(table 2).
Stomach: The rissoacean stomach does not normally
have a caecum or an elongated posterior chamber, as
occurs in various other prosobranchs ("it is not possible
to see any trace of it in the . . . Rissoacea"; Fretter &
Graham, 1962:225). OnK a few exceptions are known:
Ponder (1985:78) described the stomach of Rissoina (Ris-
soidae) as "very long due to elongation of posterior cham-
ber," and Ponder (in press) listed the character state
"posterior gastric chamber not small" for the families
Emblandidae, Truncatellidae and Stenothyridae. The
presence of a large posterior chamber in Cijclostreiniscus
beauii, similar in relative shape and organization to that
of Pomatias elegans (Miiller, 1774) (Pomatiasidae), as
described by Graham (1939:90, fig. 6D), is therefore sur-
prising. It may allow Cij. beauii to maintain a regular
supply of food particles by regulating fluctuations in the
rate of food intake (as suggested by Graham, 1939:93,
for Pomatias). especially since feeding must be strongly
affected b> tidal, and therefore water-level, changes
within the stomatopod burrows. A relatively large animal
such as Cy. beauii would be more strongly affected than
smaller vitrinellids, e.g., Circulus texanus, which were
observed to feed in very small pockets of water (personal
observation), thus maintaining a more-or-less continuous
feeding activity, as has been described, e.g., for a mem-
ber of the genus Caecum (Morton, 1975:14).
Hermaphroditism: The most striking result of this study
was the discovery of small functional females in Cy-
clostremiscus beauii with apparently functionless rem-
nants of the male reproductive system. Penial structures
are known to occur in various female rissoaceans (par-
ticularly hydrobiids and rissoids). Thiriot-Quievreux
(1977:779ff.) based her hvpothesis of sequential her-
maphroditism in four species of Ris.soa on the presence
of more-or-less developed penes in immature specimens
and lemales. However, a later in-depth study (Thiriot-
Quievreux, 1982:]67ff.) of these species through annual
cycles shov\ed a seasonal (and geographical) pattern of
penis-size distribution in females, but no evidence of sex
change. The survey did "not support the hypothesis of
a successive hermaphroditism" (1982:167). In Cyclo-
stremiscus heauii, remnants of the male apparatus are
only evident in the smallest functional ieniales (figure
53), with a gradual decrease of male structures with size.
As outlined above, we take this as evidence for a sex
change in this species rather than a simple sexual di-
morphism in shell size. During ontogeny, the male re-
productive system apparently disappears completely and
is replaced by female organs. The anteriormost part of
the pinkish-orange female albumen gland is found in the
same relative position as the prostate (of similar color
and reaction to staining) in the male phase. As no animal
was found with a very early stage of development of the
albumen gland, it cannot be decided whether these or-
gans, or parts of them, are homologous. [Reid (1986), in
a study of Mainwaringia Nevill, 1885, the first reported
case of protandrous sequential hermaphroditism in the
Littorinacea, found both a closed pallial oviduct and an
open prostate in the intersexual and female stages, "sug-
gesting that these structures are not strictly homologous"
(1986:225). In that littorinid group, small penes are re-
tained in the female phase (1986:237), and the pallial
oviduct opens into the also-retained pallial vas deferens,
then serving to carry egg capsules (1986:238).] With this
admittedly small data set for Cy. beauii from only two
localities, and the absence of data on individual devel-
opment and longevity, settlement cues and adult mo-
bility, several scenarios could be constructed. It is, for
example, theoretically possible that functional males ar-
rest growth to prolong the male phase, or that the change
into the female phase occurs only after successful mating
as a male. However, no evidence was found to consider
this sex change as labile, i.e., environmentally mediated,
as is known for other caenogastropod families such as
Calyptraeidae and Stiliferidae (see Hoagland, 1978, for
examples and discussion). The data are further insuffi-
cient to demonstrate either seasonal or partner-induced
change. The sex change in Cy. beauii appears to occur
only once, at a predetermined size (figure 53; about 8
mm shell diameter, 3 teleoconch whorls), and it appar-
entlv affects most, if not all, individuals (as the general
shift in gender over time in the Ft. Pierce population
suggests).
Thus Cyclostremiscus beauii is interpreted as a pro-
tandrous sequential hermaphrodite. The term sequential
(= consecutive, successive) is important here. In a review
of the terms protandry, protogyny and hermaphroditism,
Hoagland (1984:86) defined protandry as "the function-
ing of an organism first as male, then as female, with no
further sex change. The two sexual phases are separated
by a phase in which male primary and secondary sex
characters disappear, and the animal re-differentiates as
a female." This however omits those species which, after
an initial male-only phase, have both male and female
Page 26
THE NAUTILUS, Vol. 102, No. 1
reproductive systems developed and tiiiictional (e.g.,
members of Rissoellidac and Omal(>i;\ ridae; see below),
i.e., protandrous simultaneous hermaphrodites. [Simul-
taneous hermaphrodites, in our understanding, do not
necessariiK use "the same gonad to produce both eggs
and sperm" as defined b% Hoagland (1984:85).]
Although, at this point, we do not know whether pro-
tandrous sequential hermaphroditism is the exception,
rather than the rule, in this family, the occurrence of
hermaphroditism in Cyclostremiscus beauii could be in-
terpreted as an adaptation to its peculiar habitat char-
acterized b\ low densit\ and relative isolation. Protan-
drous sequential hermaphroditism has an ad\antage for
Cy. beauii, as each individual thereby minimizes the age
at which it first reproduces and increases the likelihood
of finding a compatible mate in a small group. Inbreeding
between siblings would also be reduced (see Ghiselin,
1969, for discussion).
Members of the approximately 25 families grouped
under Rissoacea are general!) described as gonochoristic
(see, e.g.. Boss, 1982:984), although possibly derived from
ancestors that were sequential hermaphrodites (Slavo-
shevskaya, 1984). Reported cases of hermaphroditism in
prosobranchs (e.g., Webber, 1977:10; Fretter, 1984:15)
include onl\ two genera that have classicalK been as-
signed to this superfamily, Omalogyra (Omalogyridae)
and Rissoella (Rissoellidac). Fretter (1948) described in
detail the anatomy and reproductive biology of O. ato-
muilPhilippi. 184'l)andfi. Jiap/!a(!a(Alder,'l848). Both
species were found to be protandrous simultaneous her-
maphrodites, with some likelihood of self-fertilization in
O. atomus (1948:612, 621, 630; see also Fretter & Gra-
ham, 1962:381, and 1978:218, 223). [Fretter & Graham,
1964:134, refer (erroneously:') to the .same two species as
being "protandrous con.secutive hermaphrodites. '] Both
families have been subsequently removed from the Ris-
soacea and have been recently placed outside the Cae-
nogastropoda, near (Salvini-Plawen & Haszprunar, 1987)
or in (Ponder & Waren, in press) the Allogastropoda
(= Heterostropha of authors). Thus, Cyclostremiscus
beauii is to our knowledge the only species in the su-
perfamily Rissoacea for which protandrous sequential
hermaphroditism has been demonstrated. However, as
sequential hermaphroditism is not as easily recognized
as simultaneous hermaphroditism, it might be more
widely distributed in the N'itrinellidae and/or other fam-
ilies of this group. The mismatch in descriptions of the
male reproductive system ol Tornus subcarinatus, where
a penis was lacking according to Woodward (1898) and
was found by Graham (1982), might find an explanation
after all.
ACKNOWLEDGEMENTS
For invaluable help in the field and laboratory, we es-
pecially thank William D ("Woody") Lee (SMSLP),
w hose enthusiasm, expert technique and keen eye added
significantly to the progress of this study. Dr. M. G.
Harasewych (USNM) furnished the .scanning electron
micrographs for figures 1-3. Dr Richard S. Houbrick
provided valuable field assistance (he found the largest
temale, without his glasses). Drs. Klaus Bandel (Univer-
sitiit Hamburg, West Germany), Donald R. Moore
(KSM.\S) and Winston F. Ponder (Australian Museum,
S\dne\ ) offered comments on species identifications and
larval shells Dr Ra\mond B. Manning (USNM) was first
to bring the molluscan associates of Lysiosquilla to our
attention and provided information about the stomato-
pods. Dr. Robert Robertson (ANSP), Dr. Patrick Nuttall
and Ms. Kathie Way [British Museum (Natural History)],
Dr. Terrence M. Gosliner (CAS), Dr. Fred Thompson
(Florida State Museum, Gainesville), Dr. Kenneth J. Boss
(MCZ), Dr. Philippe Bouchet (MNHN), Hugh J. Porter
(UNC-IMS), Dr. M. G. Harasewych (USNM), and Fred-
erick J. C'ollier (USNM) provided comments on and or
loans of specimens entrusted to their care. We also thank
the following: Julianne Piraino (SMSLP) for SEM assis-
tance; Patricia Linley (HBOI) for advice on histological
techniques; Tom Smoyer for help with photographic
printing; Dr. Robert Hershler (USNM), Dr. M. G. Har-
asewych (USNM), Richard E. Petit (North Myrtle Beach,
SO, kristen Metzger (HBOI), and Carol Browder (HBOI)
for help with literature acquisition. Drs. Kevin J. Eck-
elbarger (HBOI), M. G. Harasewych (USNM), Robert
Hershler (USNM), and Richard S. Houbrick (USNM) are
thanked for valuable comments on the manuscript. The
senior author gratefully acknowledges financial support
provided by a NATO postdoctoral fellowship, adminis-
tered b\ the German .Academic Exchange Service
(D.\AD), and research facilities provided by the Smith-
sonian Marine Station at Link Port, Ft. Pierce, FL.
This is Smithsonian Marine Station Contribution No.
217 and Harbor Branch Oceanographic Institution Con-
tribution No. 622.
LITERATURE CITED
Abbott, R. T. 1954. .■American seashells. D. Van Nostrand
Company, Inc., Princeton, NJ, xiv + 541 p., 40 pis
Abbott, R. T. 1974. .American seashells: the marine Mollusca
of the Atlantic and Pacific coasts of North .America, 2nd
ed. Van Nostrand Reinhold Company. New York, 663 p.,
24 pis.
Abbott, R. T. and S P Dance. 1982 Compendium of sea-
shells: a color guide to more than 4,200 of the world's
marine shells. E. P Dutton, New York, x -I- 411 [+ 1] p.,
illus.
Adam, W. and J. Knudsen. 1969 Quelques genres de Mol-
lusques prosobranches marins inconnus ou peu connus de
r.Afrique occidentale. Bulletin Institut Royal des Sciences
Naturelles de Belgique 44(27): 1-69.
Adams, .A. 1850. Monographs of Cyc/o«(rf'ma, Marryat, and
Sejmratista, Gra\ . two genera of gasteropodous mollusks.
Proceedings of the Zoological Society of London 185041-
45.
Adams, A. 1866. Monographs of the genera Cyclostrema,
Adeorbis, and Teinostoma. Thesaurus Conchyliorum.
Sowerby, G. B. (ed.) 3:249-262, pis. 255-256.
Andrews, J. 1971. Sea shells of the Texas coast. University of
Texas Press, .Austin, xvii -I- 298 p.. illus.
R. Bieler and P. M. Mikkelsen, 1988
Page 27
Andrews, J. 1977. Shells and shores of Texas. University of
Texas Press, Austin, xx + 365 p., illus.
Ankel, W. E. 1936. Prosobranchia. In: Grimpe, G. and E.
Wagler (eds. ) Die Tierwelt der Nord- und Ostsee, Vol.
I\b. .-Kkademische Verlagsgeselischaft, Leipzig, p. 1-240.
Bandel, K 1984. The radulae of Caribbean and other Me-
sogastropoda and Neogastropoda, Zoologische Verhande-
lingen No. 214, 188 p., 22 pis.
Boss, K. J. 1982. Mollusca. In: Parker, S. P. (ed. in chief).
Synopsis and classification of living organisms. Vol. 1.
McGraw-Hill Book Company, New York, p. 945-1166.
Bush, K. J. 1897. Revision of the marine gastropods referred
to Cyclostrema, Adeorbis. Vitrinella, and related genera;
with descriptions of some new genera and species belong-
ing to the .\tlantic fauna ot .America. Transactions of the
Connecticut .^cadeni) 10:97-144, pis. 22, 23.
Dall. W H 1889. \ preliminary catalogue of the shell-bear-
ing marine mollusks and brachiopods of the south-eastern
coast of the United States, with illustrations of many of
the species. Bulletin of the United States National Museum
37:221 p., pis. 1-74.
Dall, W. H. 1892. Contributions to the Tertiary fauna of
Florida, with especial reference to the Miocene Silex-beds
of Tampa and the Pliocene beds of the Caloosahatchie
River, 11. — Streptodont and other gastropods, concluded.
Transactions of the Wagner Free Institute of Science 3(2):
201-448, 1 map, pis. 13-22. p. 459-473.
Dall, VV. H. 1903. Contributions to the Tertiary fauna of
Florida with especial reference to the Silex Beds of Tampa
and the Pliocene Beds of the Caloosahatchie River in-
cluding in many cases a complete revision of the generic
groups treated of and their American Tertiary species.
Part VI. Concluding the work. Transactions of the Wagner
Free Institute of Science of Philadelphia 3(4):i-xiv, 1219-
1654, pis. 48-60.
Davis, G. M. and W, P. Carney. 1973. Description of On-
comelania lindoensis, first intermediate host of Schisto-
soma japonicum in Sulawesi (Celebes). Proceedings of the
Academy of Natural Sciences of Philadelphia 125(1 ):1-
34.
Davis, G. M., V. Kitikoon, and P. Temcharoen 1976. Mono-
graph on "Lithoglyphopsis" aperta, the snail host of Me-
kong River schistosomiasis. Malacologia 15(2):241-287.
Davis, G. M. and M. Mazurkiewicz. 1985. Systematics of
Cincinnatia winkleyi (Gastropoda: Hydrobiidae). Pro-
ceedings of the Academy of Natural Sciences of Phila-
delphia 137:28-47.
Emerson, W. K. and M K. Jacobson 1976 The American
Museum of Natural History guide to shells: land, fresh-
water, and marine, from Nova Scotia to Florida. Alfred
.\. Knopf, New York, 482 p. text -I- xviii p. index, 47 pis
Fischer, P. 1857a. Etudes sur un groupe de coquilles de la
famille des Trochidae (suite). Journal de Conchyliologie
6:168-176.
Fischer, P. 1857b. Etudes sur un groupe de coquilles de la
famille des Trochidae (fin) Journal de Conchyliologie 6:
284-288, pi. 10.
Franzen, A. 1955. Comparative morphological investigations
into the spermiogenesis among Mollusca. Zoologiska Bi-
drag fran Uppsala 30:399-456, pis. 1, 2.
Fretter, V. 1948. The structure and life history of some minute
prosobranchs of rock pools: Skeneopsis planorbis (Fabri-
cius), Omalogyra atomus (Philippi), Rissoella diaphana
(Alder) and Rissoella opalina (Jeffreys). Journal of the
Marine Biological Association of the United Kingdom 27:
597-632.
Fretter, V. 1956. The anatomy of the prosobranch Circulus
striatus (Philippi) and a review of its systematic position.
Proceedings of the Zoological Society of London 126:369-
381.
Fretter, V. 1984, Prosobranchs. In: Wilbur et al. (eds.). The
Mollusca, Vol. 7, Reproduction Academic Press, New York
and London, p. 1-45.
Fretter, V. and A. Graham. 1962. British prosobranch mol-
luscs: their functional anatomy and ecology. Ray Society,
London, xvi -I- 755 p.
Fretter, V. and A. Graham. 1964. Reproduction. In: Wilbur,
K. M. and C. M. Yonge (eds). Physiology of Mollusca,
Vol. 1. Academic Press, New York and London, p. 127-
164.
Fretter, V and A. Graham. 1978. The prosobranch molluscs
of Britain and Denmark. Part 4 — Marine Rissoacea. Jour-
nal of Molluscan Studies, Supplement 6:153-241.
Gardner, J. 1948, Mollusca from the Miocene and lower Plio-
cene of Virginia and North Carolina. Part 2. Scaphopoda
and Gastropoda. U.S. Geological Survey Professional Paper
199-B, i-iii, 179-310, pis. 24-38.
Ghiselin, M. T. 1969. The evolution of hermaphroditism among
animals. Quarterly Review of Biology 44:189-208.
Gotze, E 1938. Bau und Leben von Caecum glabrum (Mon-
tagu). Zoologisches Jahrbuch (Systematik), Jena 71(1-2):
1-190.
Golikov, A. N. and Y. I. Starabogatov. 1975. Systematics of
prosobranch gastropods. Malacologia 15(l):185-232.
Graham, A. 1939. On the structure of the alimentary canal
of stvle-bearing prosobranchs. Proceedings of the Zoolog-
ical Society of London (B) 109(1):75-112.
Graham, A. 1982. Tornus subcarinatus (Prosobranchia, Ris-
soacea), anatomy and relationships. Journal of Molluscan
Studies 48:144-147.
Guppy, R. J. L. 1866. On the Tertiary Mollusca of Jamaica.
Quarterly Journal of the Geological Society of London
22(l):28i-295, pis. 16-18.
Hartman, O. 1945. The marine annelids of North Carolina.
Bulletin of the Duke University Marine Station 2:1-54, 10
pis., 2 charts.
Haszprunar, G. 1985. The fine morphology of the osphradial
sense organs of the Mollusca. I. Gastropoda, Prosobranchia.
Philosophical Transactions of the Royal Society of London,
B 307:457-496.
Hershler, R. and F. G. Thompson. 1987. North .'American
Hydrobiidae (Gastropoda: Rissoacea): redescription and
systematic relationships of Tryonia Stimpson, 1865 and
Pyrgulopsis Call and Pilsbry, 1886. The Nautilus 101(1):
25-32.
Hertlein, L. G. and A. M. Strong. 1951 Eastern Pacific ex-
peditions of the New Y'ork Zoological Society. XLIII. Mol-
lusks from the west coast of Mexico and Central .America.
Part X. Zoologica, Scientific Contributions of the New York
Zoological Society 36(2):67-120, pis, 1-11.
Hoagland. K. E. 1978. Protandrv and the evolution of en-
vironmentally-mediated sex change: a study of the Mol-
lusca. Malacologia 17(2):365-391.
Hoagland, K. E. 1984. Sententia. Use of the terms protandry,
protogyny, and hermaphroditism in malacology. Ameri-
can Malacological Bulletin 3(l):85-88.
Humason, G. L. 1962. Animal tissue techniques. W. H. Free-
man and Company, San Francisco and London, xv -f
468 p.
Page 28
THE NAUTILUS. \'ol. 102, No. 1
Humfrey, M. 1975. Sea shells of the West Indies; a guide to
the marine molluscs of the Caribbean. Taplinger PubMsh-
ing Company, New York, 351 p., 32 pis.
International Code of Zoological Nomenclature [ICZN]. 1985.
Third edition. Ride, VV. D. L.,C. W. Sabrosky. G. Bernardi,
and R. \' Mel\ille(eds.). International Trust for Zoological
Nomenclature, London, .xx + 338 p.
Iredale, T 1914. On some invalid niolluscan generic names.
Proceedings of the Malacological Societv ol London 11(2):
170-178.
Iredale, T. 1915. The nomenclature of British marine Mol-
lusca. Journal of Conchology 14(ll):341-346.
Jablonski, D and R. A Lutz. 1980. Molluscan larval shell
morphology Ecological and paleontological applications.
In: Rhoads', D. C. and R. A Lutz (eds. ) Skeletal growth
of aquatic organisms. Topics in geobiolog) , Vol. 1. Plenum
Press, New York and London, p. 323-377.
Jeffreys, J. G. 1865. British conchology, or an account of the
Mollusca which now inhabit the British Isles and the sur-
rounding seas. Vol. 111. Marine shells, comprising the re-
maining Conchifera, the Solenoconchia, and Gasteropoda
as far as Littorina. ]. van \'oorst, London, 393 (+ 1) p.,
8 pis. + frontispiece
Johansson, J. 1939. .•Xnatomische Studien ijber die Gastro-
podenfamilien Rissoidae und Littorinidae. Zoologiska Bi-
drag fran Uppsala 18( 1938-40);287-396, pis. 1-11.
Johansson, J. 1940. Ein Beitrag zur Kenntnis des Kristall-
stielsackes der Mollusken und der Flimmerbewegung in
demselben. Arkiv for Zoologi 33B(3):l-8.
Johansson, J. 1949. Dber die weiblichen Geschlechtsorgane
von Hijala vitrea, einer von deni Rissoa-Typus stark ab-
weichenden Form der Gruppe Rissoacea. Arkiv for Zoologi
42A(7);l-6.
Johansson, J. 1956. Genital organs of two Alvania species,
and a comparison with related families (Moll, Pros.). Arkiv
f6r Zoologi 9(16):377-387, pi. 1.
Keen, A. M. 1971. Sea shells of tropical west .America —
marine mollusks from Baja California to Peru, 2nd ed.
Stanford University Press, Stanford, C.A, xiv -t- 1064 p.,
22 pis.
Kuroda, T , T. Habe, and K. Oyama. 1971. The sea shells of
Sagami Bay. Maruzen, Tokyo, xi.x -I- 741 p. [in Japanese],
pis. 1-121, 489 p. [in English], 51 p., index, map.
Laseron, C. F. 1958. Liotiidae and allied molluscs from the
Dampierian Zoogeographical Province. Records of the
Australian Museum 24(11);165-182.
Lebour, M. V. 1937. The eggs and larvae of the British proso-
branchs with special reference to those living in the plank-
ton. Journal of the Marine Biological Association of the
United Kingdom 22:105-166.
Lilly, M. M. 1953. The mode of life and the structure and
functioning of the reproductive ducts of Buhijnia tenta-
culata (L.). Proceedings of (he Malacological Society of
London 30:87-110.
Mackintosh, N. A. 1925. The crystalline style in gastropods.
Quarterly Journal of Microscopical Science 69:317-342,
pis. 20, 21.
Melvill, J. C. 1906. A revision of the species of Cyclostre-
matidae and Liotiidae occurring in the Persian Gulf and
North .Arabian .Sea. Proceedings of the Malacological So-
ciety of London 7(l):20-28, pi 3.
Moore, D. R. 1962. The systematic position of the family
Caecidae (Mollusca: Gastropoda) Bulletin of Marine Sci-
ences of the Gulf and Caribbean 12(4)695-701
Moore, D. R. 1964. The familv X'itritiellidae in South Florida
and the Gulf of Mexico. Ph.D. dissertation. University of
.Miami, Coral Gables, FL, xi + 235 (+ 2) p.
Moore, D. R. 1965. New species of \itrinellidae from Gulf
of Mexico and adjacent waters. The Nautilus 78(3):73-79,
pis. 7, 8.
Moore, D. R. 1969. Cyclostrema miranda Bartsch, a synonym
of Tornus subcarinatus Montagu. The N'eliger 12(2):169-
170,
Moore, D. R. 1972. Cochliolepis parasitica, a nonparasitic
marine gastropod, and its place in the Vitrinellidae. Bul-
letin of Marine Science 22(1):100-112.
Morris, P. A. 1973. A field guide to shells of the .Atlantic and
Gulf Coasts and the West Indies, 3rd ed. Houghton Mifflin
Company, Boston, xxviii -I- 330 p., 76 pis.
Morton, J. 1975. Form and habit in some small gastropods of
New Zealand boulder beaches. The \'eliger 18(1): 1-1 5.
Ode. H. 1987 Distribution and records of the marine Mol-
hrsca in the nortliwest Gulf of Mexico (a continuing mono-
graph) family X'itrinellidae Bush, 1897 (continued). Texas
Conchologist'24(l):21-40.
Olsson, A. A. and M. Smith. 1951. New species of Epitoniidae
and Vitrinellidae from Panama City. The Nautilus 65(2):
44-46, pi. 3.
Pilsbry, H. A. 1953. Vitrinellidae. In: Olsson, A A and A.
Harbison (eds). Pliocene Mollusca of southern Florida with
special reference to those from north Saint Petersburg.
Monographs of the .Academv of Natural Sciences of Phila-
delphia, no. 8, p. 411-438, pis. 49-56.
Pilsbry, H. A. and T. L. McGinty. 1945a. Cyclostrematidae
and Vitrinellidae of Florida— I. The Nautilus 59(1):1-13,
pis. 1, 2.
Pilsbry, H. A. and T L McGint\ . 1945b. "Cyclostrematidae"
and Vitrinellidae of Florida, 11, The Nautilus 59(2):52-59,
pi. 6,
Pilsbry, H. A. and T. L, McGinty. 1946a. "Cyclostrematidae"
and Vitrinellidae of Florida, Part III. The Nautilus .59(3):
77-83, pi. 8.
Pilsbrv, H. A. and T. L. McGinty. 1946b. \itrinellidae of
Florida, Part 4. The Nautilus'60(l):12-18, pi. 2.
Pilsbrv, H. A. and T. L. McGinty. 1950. Vitrinellidae of
Florida: Part 5. The Nautilus 63(3):85-87, pi. 5.
Pilsbry, H A. and A A Olsson 1945. \'itrinellidae and similar
gastropods of the Pariamic Pro\ince. Part I. Proceedings
of the Academy of Natural Sciences of Philadelphia 97:
249-278, pis. 22-30.
Pilsbry, H. A. and A. A. Olsson. 1952. \itrinellidae of the
Panamic Province: II. Proceedings of the .Academv of Nat-
ural Sciences of Philadelphia 104:35-88, pis. 2-13.
Ponder, W. F. 1966. On a subterranean snail and a tornid
from New Zealand. Journal of the Malacological Societ\
of Australia 10:35-40, pis. 4, 5.
Ponder, VV. F. 1982. The anatomv and relationships ol Hy-
drococcus brazieri (T. Woods). Journal of Molluscan Stud-
ies 48:64-79.
Ponder, W. F. 1983. Review of the genera of the Barleeidae
(Mollusca: Gastropoda: Rissoacea), Records of the .Austra-
lian .Museum .35:231-281.
Ponder, W. F. 1985. A review of the genera of the Rissoidae
(Mollusca: Mesogastropoda: Rissoacea). Records ot the
Australian Museum, Supplement 4:1-221.
Ponder, VV, F. In press The rissoacean radiation — a prelim-
inary phylogeny. Malacological Review, Supplement.
Ponder, VV. F. and .\ VV'aren In press. Classification of the
Caenogastropoda and Heterostropha — a list of the family-
R. Bieler and P. M Mikkelsen, 1988
Page 29
group names and higher taxa. Malacological Review, Sup-
plement.
Porter, H. J. 1974. The North CaroHna marine and estuarine
Molhisca — an atlas of occurrence. I'niversity of North Car-
ohnu. Institute of Marine Sciences, Morehead (."ity, NC,
vi + .351 p.
Reid, D. G. 1986. Mainwaringia Nevill, 1885, a littorinid
genus from Asiatic mangrove forests, and a case of pro-
tandrous hermaphroditism. Journal of Molluscan Studies
52:225-242,
Hequien, E. 1848. Catalogue des coquilles de 1 lie de Corse.
F. Seguin Aine, Avignon, 109 (+ 2) p.
Rios, E. C. 1975. Brazilian marine mollusks iconography.
Museu Oceanografico, Funda^ao Universidade do Rio
Grande, Rio Grande, 331 p., 91 pis.
Rios, E. C. 1985. Seashells of Brazil. Museu Oceanografico,
Fundaijao Cidade and L'niversidade do Rio Grande, Rio
Grande, 328 [+ 1] p., 102 pis.
Salvini-Plawen, L. von and G Haszprunar. 1987. The Veti-
gastropoda and the S)stematics of streptoneurous Gastrop-
oda (Mollusca). Journal of Zoology, London 211:747-770.
Serene, R. 1954. Observations biologiques sur ies Stomato-
podes. Memoires de ITnstitut Oceanographique de Nhat-
rang 8:1-93, 10 pis.
Simpson, C. T. 1887. Contributions to the Mollusca of Florida.
Proceedings of the Davenport Academ\ of Natural Sci-
ences 5:45-72.
Slavoshevskaya, L. \". 1978. Peculiarities of the genital s\ stem
of the Rissoacea and their importance for taxonomy in this
superfamily. Malacological Review 11:112-114.
Slavoshevskaya, L. V. 1984. An analysis of the organization
of the Rissoacea (Mollusca: Gastropoda). 2. The reproduc-
tive system. Zoologicheskii Zhurnal 63(3):361-372 [in Rus-
sian].
Smith, M. 1937. Further notes upon Tertiary and Recent
mollusks from Florida together with descriptions of new-
species. The Nautilus 51(2):65-6S, pi. 6.
Stimpson, W. 1858. [On Cochlioleph parasiticus]. Proceed-
ings of the Boston Society of Natural History 6:307-309.
Taylor, D. W. and N. F. Sohl. 1962. An outline of gastropod
classification. Malacologia l(l):7-32.
Ta\lor, J. B. 1975. Planktonic prosobranch veligers of Ka-
neohe Bay. Ph.D. thesis. University of Hawaii, Honolulu,
599 p., illus.
Tliiriot-Quie\reux, C. 1977. Particularite de I appareil genital
de quelques especes de Ri.ssoidae (Mollusca: Mesogastrop-
oda). Compte Rendu de I'Academie des Sciences, Paris,
serie D 285(7):779-781, pi. 1.
Thiriot-Quievreux, C. 1982. Donnees sur la biologic .sexuelle
des Rissoidae (Mollusca: Prosobranchia). International
Journal ol Invertebrate Reproduction 5:167-180.
Thorson, G. 1946. Reproduction and larval development of
Danish marine bottom invertebates, with special reference
to the planktonic larvae in the Sound. Meddelelser fra
Kommissionen for Danmarks Fiskeri- og Havunders0gel-
ser, Serie Plankton, Vol. 4(1), 523 p., illus.
Tryon, G. W. 1888. Monograph of the families Neritidae,
Neritopsidae, Adeorbiidae, Cyclostrematidae, and Lioti-
idae. Manual of Conchology 10:3-160, pis. 1-36.
Yokes, H. E. and E. H. Yokes. 1983. Distribution of shallow-
water marine Mollusca, Yucatan Peninsula, Mexico. Me-
soamerican Ecology Institute Monograph 1, Middle Amer-
ican Research Institute Publication 54, viii -t- 183 p., 50
pis.
Warmke, G. L. and R. T. .Abbott. 1961. Caribbean seashells:
a guide to the marine mollusks of Puerto Rico and other
West Indian Islands, Bermuda and the Lower Florida Keys.
Livingston Publishing Company, Wynnewood, P.\, \ +
348 p., 44 pis.
Webber, H. H. 1977. Gastropoda: Prosobranchia In: Giese,
A. C. and J. S. Pearse (eds. ). Reproduction of marine in-
vertebrates, lY. Molluscs: gastropods and cephalopods. Ac-
ademic Press, New York, p. 1-97.
Wells, H. W., M. J. Wells, and 1 E. Gray. 1961. Food of the
sea-star Astropecten articiilatus. Biological Bulletin 120:
265-271.
White, K. M. 1942. The anatomy of Cyclostrema bushi D.
& F. Proceedings of the Malacological Societv of London
25:89-94, pi. 2.
Woodring, W. P. 1928. Miocene mollusks from Bovvden, Ja-
maica. Part II: Gastropods and discussion of results. Car-
negie Institution of Washington, Publication .395:vii + 564
p., 40 pis.
Woodward, M. F. 1898. On the anatomy of Adeorbis sub-
carinatus, Montagu. Proceedings of the Malacological So-
ciety of London 3:140-146, pi. 8.
Y'onge, C. M. 1930. The crystalline style of the Mollusca and
a carnivorous habit cannot normally co-exist. Nature, Lon-
don 125:444-445.
THE NAUTILUS 102(1 ):30-.35, 1988
Page 30
Geographical Distribution of Some Epitoniidae
(Mollusca: Gastropoda) Associated with Fungiid Corals
Helen DuShane
Research Associate
Natural History Museum
Los Angeles Count)
Los Angeles, CA 90007, USA
mailing address:
15012 El Soneto Drive
Whittier, CA 90605, USA
ABSTRACT
The known ranges of three coral associated epitoniids, Epito-
nium costulalum (Kiener, IS39), Epitonium ulu Pilsbry, 1921,
and Epitonium hulldtttm (Snwcrln, 1844) are reviewed
INTRODUCTION
The family Epitoniidae has had a complicated nomen-
clatural history, with more than 3,000 Recent and fossils
species named. The genus Epitonium is an overwhelm-
ingly large group about which itiucIi has been published,
but information on habitats, ecology, growth patterns,
and reproduction is known for relatively few species.
Some epitoniids are perhaps the only gastropods e.xcept
certain coraiiiophiiidae to have feeding associations with
iuiigiid corals.
The stony corals comprise a large group of animals
showing a great variety of form. Wells ( 1956:F388) listed
1 1 living genera in the family Fungiidae Dana, 1846,
and further stated. "Within the family Fungiidae some
animals are attached as juveniles, but as adults become
detached and live free on rubble substrates. Others re-
main attached throughout their lives." There are 11-12
genera, within w liich are probably 49 species. Fungiids
live in warm seas in water temperatures 22 °-33 °C. The
corallum (skeleton) is rounded or elongate, varying in
length from about 5 cm to 35 cm, and in width from 5
cm to 20 cm Known commonly as "hat' or "mushroom "
corals, living FuiifJia sp|). arc pink, green, off-white, or
pale tan. Distribution of these corals is extensive, from
the east coast of Africa, across the Arabian Sea, the Bay
of Bengal, Western Pacific Ocean and including the
Southern and Flastern Pacific oceans.
Previous reports ol epitoniids associated with Fiingia
are those of Root (1958:8), Robertson (1963:57, 1965:7,
1970:45), Bosch (1965:267), Taylor (1977:254), Kay (1979:
152), Bell (1982:508, 1985:161 ). Sabelli and Taviani (1984:
92). These papers report at least three species of epito-
niids associated w ith species ol Fungia.
A collection of Epitonium spp. made by SCUBA divers
in the northern Red Sea verified that Fungia is "home"
to the tollowing species of epitoniids: Epitonium cos-
tulalum (Kiener, 1839), originally described from an
unknown locality; Epitonium ulu Pilsbr\. 1921, origi-
nalK described from Hawaii; and Epitonium hullatum
(Sowerby, 1844), originalK described from the Philip-
pines. An unidentified Epitonium sp. associated with
Fungia was also reported by Sabelli and Taviani (1984)
from the Red Sea.
The following abbreviations appearing in the text are
defined as follows:
ANSP — Academy of Natural Sciences, Philadelphia. P.\.
GENEVA — Museum d Histoire Naturelle, Geneva, Sw it-
zerland.
NHM, L.A. — Natural History Museum. Los Angeles
County, Los Angeles, CA.
Bratcher (Collection — Twila Bratcher, Los .Angeles, C.\.
Chane)' Collection — Henry Chaney, Redondo Beach, CA.
DuShane Collection — Helen DuShane, Whittier, CA.
Kaiser Collection — Kirstie Kaiser, Park Citv, UT.
RECORDS
Epitonium costulatum (Kiener, 1839)
(figures 1, 2)
Discussion: Epitonium costulatum (Kiener, 1839) as
far as presentK know n lives in warm seas from 10°N to
SCN, from the Red Sea, Bay of Bengal (India and Thai-
land), and the Philippine Islands. This is the first report
of the egg capsules oi E. costulatum found on the un-
derside ol Fungia sp. from the Red Sea. Each egg capsule
was encrusted w ilh bits of white coral-sand and attached
to another by two slender threads (figure 2). The number
of egg capsules per mass is approximately 100.
Hoot (1958) was the first to report this species as an
unidentified eiJitoniid li\ing under Fungia, in the Sulu
Archipelago, Philippines. His six specimens are at the
Academy of Natural Sciences of Philadelphia (ANSP
230639) and were studied b\ Robertson (1963) who later
(1970) compared them with the holot\pe in CJeneva
(original locality unknown) and concluded that Root's
specimens are indeed £. costulatum. The ANSP speci-
H. DuShane, 1988
Page 31
4
Figure 1. Apertural view of Epitoniitm costulatum (Kiener, 1839). Length 28 mm, width 13 mm, DuShane Collection. Straits of
Tiraii, Red Sea Figure 2. Coral-sand encrusted egg mass of £. costulatum. Chaney Collection. Straits of Tiran, Red Sea. Figure
3. Epitonium ulu Pilsbr\, 1921. Length 13.5 mm, width 5.5 mm, Bratcher Collection. Saudi .Arabia, Red Sea. Figure 4. £. ulu
with egg mass. Length of shell 16 mm, width 11 mm, Chaney Collection. Tiran Island, Straits of Tiran, Red Sea. Figure 5.
.Apertural view of Epitonium bullatum (Sowerby, 1844). Length 13 mm, width 9 mm, Kaiser Collection. Thomas Reef, Sinai, Red
Sea. Figure 6. Dorsal view of E. htllatum. Kaiser Collection.
Page 32
THE NAUTILUS, Vol. 102, No. 1
mens have 21-26 costae and range in length from 12 4
to 35.3 mm (Robertson, 1963, 1970).
Robertson (1963:60) stated, "There are some wentle-
traps which seemingly live throughout most ot their post-
larval lives with (relatively) large sea anemones (or cor-
als). Such species are . . . £. aff. costulatum ..." "This
F.pitonitim. the oiiK wentletrap so far found w ith a coe-
leiiterate other than a sea anemone, presurnabK feeds
on Fungia." We now know that £. costulatum lays its
gelatinous egg masses under Fungia on a sandy, rubble
substrate, in depths of 2-30 m. Feeding observations are
lacking.
Subsequent to 1958, other specimens have been col-
lected from various Indo-Pacific localities.
Recent records: Ba\ of Bengal, India. One specimen,
length 31 mm, with 17 costae and 12 whorls. DuShane
Collection.
Raya Island, Bay of Bengal, Thailand. One specimen,
length 34.5 mm, with 23 costae and 1 1 \\ horls. DuShane
Collection.
Phuket Island. S\V Thailand, Collected by fishermen
under "hat" coral, 15-20 m, 5/22/85. Three specimens,
lengths 23.5, 24.5, 25 mm, with 19, 20, 26 costae and 7,
9, 12 whorls. DuShane Collection.
Phuket Island, S\V Thailand. Collected by fishermen
under "hat" coral, 15-20 m, 5/22/85. One specimen,
length 23.5 mm, with 20 costae and 10 whorls. NHM,
L.A. 124505.
Tiran Island (SW), Straits of Tiran, Red Sea (27°57'N,
34°32'E). Collected by Henry Chaney, 10/31/85, SCU-
BA, 2-5 m in rubble under Fungia sp., four specimens,
three specimens, Chaney Collection, lengths 13, 20, 29
mm, with 18, 23, 28 costae and 8, 9, 13 whorls; one
specimen, DuShane Collection, length 28 mm, with 24
costae and 12 whorls. .\\\ C^haney collected specimens
were live-taken, two were pink in color, with gelatinous
egg masses as described above.
Thomas Reef, Sinai, Red Sea (27°59'N, 34°27'E). Col-
lected by Kirstie Kaiser, 10/30/85, SCUBA, 6.1 m in
sand pockets under live, detached Fungia sp., one spec-
imen, length 23 mm, witli 21 costae and 11 whorls, water
temperature 25 °C, live shell attached to coral. Kaiser
Collection.
Little Hiva, Maldive Islands. Collected by Henry Cha-
ney, 8/31/86, SCUBA, 1 m reef sand, under Fungia
repanda Dana, 1846, one specimen, live-taken, length
14 mm, with 20 costae and 11 whorls. DuShane Collec-
tion.
Epitonium ulu Pilsbry, 1921
(figures 3, 4, 7-10)
Discussion: Epiloniuiit ulu l'ilsbr>, 1921, a widely dis-
tributed species, lives in warm seas from 30°N to 5°S,
from the Red Sea, to the Maldive Islands, New Guinea
and Haw aii. It is specific to certain species of the stony
coral genus Fungia. Masses of beige colored egg capsules
are laid on the concave underside of Fungia spp. attached
to both the coral and to adult Epitonium.
Bosch (1965) reported that Fungia .scutaria Lamarck,
1816, from Kaneoke Ba\, Oahu. Hawaii, was infested
with both eggs and adults of Epitonium ulu. The snails
had large amounts of pink tissue containing symbiotic
algal cells and nematoc\sts the\ had ingested.
Taylor (1977) studied the growth rate of Epitonium
ulu feeding on the sea anemone Aiptaaia sp. and found
that the intracapsular development time for this species
varied from five days to several weeks. She found that
after hatching the ju\ eniles added approximately 0.2 mm
of shell length per day.
Bell (1982) observed that £. ulu produced a "mean of
32 capsules per day, each capsule containing 500-600
eggs." She later stated (Bell, 1985) that "embryos com-
plete intracapsular de\elopment in six da\s at 26-27''C
and hatch as planktotrophic veligers. '
Epitonium ulu has been collected recentK from five
localities.
Recent records: Maldive Islands. Robertson (1965) re-
ported an Epitonium sp. living in association with Fun-
gia. He now identifies the four specimens as Epitonium
ulu (personal communication, 1986).
Papua New Guinea. Collected by Tw ila Bratcher, 1980,
from under Fungia sp. (figure 3), 11m, water temper-
ature 28 °(]. length 16 mm, w ith 30 costae and 11 w-horls.
Bratcher Collection, and one specimen, length 12 mm
with 30 costae and 12 whorls. DuShane (Collection.
Tiran Island (SW), Straits of Tiran, Red Sea (27°57'N,
34°32'E). Collected b\' Henry Chaney. 10/31/85. SCU-
BA, 2-5 m, with egg mass, under detached Fungia on
rubble substrate. Five live-taken specimens, all exuding
mucus, were collected. Four specimens, Chanc\ Collec-
tion, lengths 3, 4, 5, 16 mm, with 10, 11, 24, 33 costae
and 9, 10, 12, 12 whorls; one specimen, DuShane Col-
lection, length 6.5 mm, with 21 costae and 11 whorls,
Sinafir Island, Saudi Arabia, Red Sea. (Collected b\
Twila Bratcher, 10/30/85, SCUBA, 9 m, with eggs under
detached Fungia. Two live-taken specimens, lengths 9,
13.5 mm, with 17, 24 costae and 9. 10 whorls. Bratcher
Collection.
Thomas Reef, Sinai, Red Sea (27°59'N, 34°29'E). Col-
lected by Kirstie Kaiser, 10/30/85, SCUBA, 6 m, water
temperature 25 °C. .\ttached with eggs to underside of
detached Fungia. 1\ ing in small sand pockets with sand
and light rubble, l-'our live-taken specimens, lengths 6,
8, 15, 19 mm, with 17, 20, 23, 32 costae and 10, 10, 13,
14 whorls, all exuding mucus when collected. Kaiser
Collection.
All three S(CUB.'\ divers (Bratcher. Chaney, Kai.ser)
collected egg masses (figures 4, 8-10) associated with
specimens of Epitonium ulu from the Red Sea. The eggs
were off-w liite w ith light purple spots. One egg sac con-
tained 4()()-()()0 embr\()s in different stages of de\elop-
ment. Each translucent egg sac w ithin the cluster is pa-
pillose over the entire external surface. The papillae are
softly rounded. From each egg sac three transparent
twisted threads are attached at each end of the oval egg
H. DuShane, 1988
Page 33
8
10
.113 mm
Figure 7. Epitonium ulu Pilsbry, 1921 with egg masses on concave underside of Fungia sp. Figure 8. Egg capsules of £. ulu
that appear to be empty and transparent. Figure 9. Developing, opaque embryos of £. ulu located on the inner surface of the
transparent sac Figure 10. Prehatching veligers of £. ulu. Diameter of a single snail 0.113 mm.
sac, and eventually twist together to form a single knotted
mass. Each connecting thread appears to be many times
the length of the sac itself.
The egg capsules e.xamined under a microscope (300 x )
showed the young capsules in three different stages of
development. Within each capsule development was uni-
form. The first stage capsules appeared to be empty and
transparent (figure 8). The second stage showed devel-
oping, opaque, white embryos that appeared to be lo-
cated on the inner surface of the sac, but not filling the
entire compartment (figure 9). The third stage consisted
of well-developed veligers, thus crowding the capsule
(figure 10). One capsule in the third stage of development
contained 400-600 free swimming veligers of equal size
and development. The diameter of a single veliger was
0.113 mm. The gibbous shell is transparent e.xcept for
the darkened area of the columella. In each embrvonic
shell the animal appeared as a uniform gelatinous mass
with two dark spots.
Epitonium (Globiscala) bullatuni (Sowerby, 1844)
(figures 5, 6)
Discussion: Epitonium {Globiscala) bullatum (Sower-
by, 1844) lives in warm seas from 35°N to 30°S, from the
Red Sea, East Africa, New Guinea, Australia, the Phil-
ippines, and Japan. Kilburn (1985:330-331) reported this
variable species from off Southern Mozambique, and in
addition to living under species of Fungia, "under rocks
in low tide pools, associated with the actinian Fseudac-
tinia flagellifera (Hertw.) on w hich it feeds; on occasion
it ma\ be partly covered by the basal disk of the anem-
one. Juvenile bullatum shelter among coralline algae,
probably feeding on the small anemones that are at-
Page 34
THE NAUTILUS, Vol. 102, No. 1
tached to the fronds." He synonymized Scala (Globis-
cala) papyracea do Boiiry, 1912 from Natal, Epitonium
(Glolmcala) woolacuttae Kersiake, 1958 from South
Queensland and GlobUcala kashiuajimensis .\zuma, 1962
from Japan with £. bullatum (Sosserby, 1844).
Recent records: Thomas Reef, Sinai, Red Sea (27°59'N,
.34°27'E), collected by Kirstie Kaiser, 10/30/85, SCUBA,
6. 1 m, water temperature 25 °C. one live specimen from
under live, detached Fungia. length 13 mm, with 40
almost imperceptible costae, 7 whorls, protoconch whorls
lost, spiral striations weak when viewed under a micro-
scope (10 X ), umbilicate, color white. A second live spec-
imens was eaten by a wrasse (Labridae), before it could
be collected. An egg mass was observed and photo-
graphed with this species but not collected.
New Guinea. Three specimens in the DuShane Col-
lection (ex Withrow Collection), taken in 30 m, lengths
11, 11.5, 13 mm, with 8, 9, 9 whorls.
Epitonium sp.
An unidentified species of £pi7o;!nu>i was reported taken
from under Ftingia paunwtensis (Stutchbur\-, 1833) off
Saudi Arabia in the Red Sea b\ Sabelli and Taviani
(1984). Three specimens in varying growth stages were
collected with an egg mass at a depth of 2 m. The authors
described their specimens as umbilicate with a very thin
shell, 22 obsolete costae and interspaces crossed by fine
spiral grooves. This is similar to the description of Epi-
tonium ulu, but comparison with E. ulu is needed.
The extensive paper by Jousseaume (1911) treating 58
species of epitoniids from the Red Sea is of little help in
identifying species from that area. Unfortunately, his
figures are too small to be of value, and some of his
descriptions are vague.
CONCLUSIONS
With an upper surface exposed to the light and a pro-
tected concave under surface, species of the genus Fun-
gia serve as host for several species of epitoniids with
very delicate shells, thin, fragile walls, and numerous
costae that are easiK shattered. Whether the wentletraps
feed on the mucus from the host polyps or the polyps
themselves is unknown. Taylor (1975) concluded that
Epitonium ulu can, under laboratory conditions, feed
on the sea anemone Aiptasia. All other evidence indicates
that E. ulu deposits eggs only under Fungia. The coral
is used as the spawning site, the eggs being laid in ge-
latinous masses, each capsule connected to another by
two threads. The ability of these epitoniids to locate and
utilize the underside of F'ungiidae requires further study.
An increasingly high degree of specificity is being found
w ith certain epitoniids seemingK having permanent as-
sociations with Fungia corals.
ACKNOWLEDGEMENTS
My thanks to Robert Robertson for encouragement, and
for reading the manuscript and making useful sugges-
tions; to William K. Emerson and James H. McLean for
reading the manuscript and offering advice; to the divers,
Twila Ikatcher, Henry (^hane\ , and Kirstie Kaiser, who
returned (rom the Red Sea with epitoniids that formed
the backbone of this paper; to the personnel of the .\llan
Hancock Foundation Library, Universit>- of Southern
California, for help in locating several rare volumes; to
Da\id Nhilliner and Bertram Draper for the excellent
photographs; and to Kirstie Kaiser for the drawings.
LITERATURE CITED
Azuma, M, 1962. Descriptions of five new species of Japanese
Epitoniidae. Venus 22(2);130-136, text figs. 1-6.
Bell, J. L. 1982. Larval development and metamorphosis of
the prosobranch mollusc, Epitonium ulu, associated with
a solitary coral. Pacific Science 16:508.
Bell, J. L. 1985. Larval growth and metamorphosis of a proso-
branch gastropod associated with a solitar\ coral. Pro-
ceedings of the Fifth International Choral Reef Congress,
Tahiti 5:159-164, 6 figs.
Bosch, H. F. 1965. .\ gastropod parasite of solitary corals in
Hawaii. Pacific Science 19:267-268, 1 fig.
Boury, E. de. 1912. Description de Scalidae nouveaux ou peu
connus. Journal de Conch\ liologie 60t2):87-108, 1 pi.
Dana, J. D. 1846. Zoophytes. United States Exploring Ex-
pedition during the vears 1838-1842 under the command
of Charles Wilkes, L .S.N., Vol. 7, p. 1-741 Philadelphia.
Dana, J. D. 1849. Descriptions of fossils. United States Ex-
ploring Expedition, 1838-1842, under the command of
Charles Wilkes, U.S.N. Geology. By James D. Dana. \'ol.
X, .Appendix, p. 681-729, pars. Atlas, xxi, pis. pars. Phila-
delphia, 1849.
Jousseaume, F. P. 1911[1912]. Faune malacologique de la
Mer Rouge. Memoires de la Societe Zoologique de France
24:180-246, pis. .5-7.
Kay, E. A. 1979. Hawaiian marine shells Reef and shore
fauna of Hawaii. Section 4: Mollusca. Bernice P. Bishop
Museum Special Publication 64(4). Bishop Museum Press,
Honolulu, Hawaii, 653 p., 2 figs.
Kersiake, J. 1958. A new Epitonium from eastern .Australia.
Proceedings of the Royal Zoological Society of New South
Wales for 1956-57, p.' 157-158, 1 fig.
Kiener, L. 1838-39. Species general et iconographie des co-
quilles vivantes; Genre Scalaire. Rousseau, Paris 7:1-22,
pis. 1-7 (text: 1839; plates: 1838).
Kilburn, R. N. 1985. The family Epitoniidae (Mollusca: Gas-
tropoda) in southern Africa and Mozambique, Annals of
the Natal Museum 27(l):239-337, 171 figs.
Lamarck, de M. de. 1816. Histoire naturelle des animaux sans
vertebres, Paris 2:1-568.
Pilsbry, H. A. 1921. Marine mollusks of Hawaii. Proceedings
of the Academy of Natural Sciences of Philadelphia 72:
360-382.
Quelch, J. J. 1886, Scientific results of the vo\age of Chal-
lenger. Report on the reef corals collected b\ H.M.S. Chal-
lenger during the years 1873-76. London. Zoology 16:1-
203.
Robertson, R. 1963. Wentletraps (Epitoniidae) feeding on sea
anemones and corals. Proceedings of the Malacological
SocietN of London 35(2&3):51-63, pis. 5-7.
Robertson, R 1965 Coelenterate-associated prosobranch gas-
tropods .•Viinual Reports of the .American Malacological
Union for 1965, 32:6-8.
H. DuShane, 1988
Page 35
Robertson, R. 1970. Review of the predators and parasites of
stony corals, with special reference to symbiotic proso-
branch gastropods. Pacific Science 24(l):43-54.
Root, J. 1958. Rapa rapa in the Snlu Sea. Hawaiian Shell
News 7:7-8,
Sabelli, B. and Marco Taviani.
Fungia-associated epitonid
20(l-4):91-94,
Sowerby, G. B., II. 1844 (July)
of Scalaria, collected by Mr. H. Cuming, to be figured in
the fourth part of Thesaurus Conchyliorum. Proceedings
of the Zoological Societ) of London, pt 12:1-38.
Stuchbury, T. 1833. .\\\ account of the mode of growth of
young corals of the genus Fungia. Transactions of the
Linnean Society of London 16:493-498, pi. 32, figs. 6a, b.
1984 Red Sea record of a
sic]. Bollettino Malacologico
Descriptions of new species
Taylor, J B 1975 Planktonic prosobranch veligers of Ka-
neolie Bay Pli D. dissertation, I'niversity of Hawaii, Ho-
nolulu, 599 p.
Taylor, J. R. 1977. Growth rate in juvenile carnivorous proso-
branchs (Mollusca: Gastropoda) of Kaneohe Bay, Oahu,
Hawaii. Proceedings Third International Coral Reef Sym-
posium Rosenstiel School ut Marine and Atmosplieric Sci-
ence, University of Miami, Miami, Florida 3(l):253-259,
8 figs.
Wells, J. W. 1956. Treatise on invertebrate paleontology. Part
F. Coelenterata. Geological Society of America and Uni-
versity of Kansas Press, xvii -I- 498 p., 2,700 figs.
THE NAL TILLS 102(1 ):36-.39, 1988
Page 36
A New Species of Vasum (Gastropoda: Turbinellidae)
from off Somalia
William K. Emerson
Waller E. Sage, III
Department of ln\ertebrates
American Museum of Natural History
New York, NY 10024-5192, USA
ABSTRACT
Vasum stephanti new species is described from moderately
deep water off Cape Gardafui (Ras Asir), Somalia and com-
pared with congeners.
INTRODUCTION
In the past few years, deep-sea commercial fisheries op-
erations off the northeastern coast of SomaUa have re-
sulted in the discovery of several new or otherwise in-
teresting species of moilusks. Lorenz (1987:11) described
P.seudosiiunia wieseonim new species from the region
off (^ape Ras Hafun in about 300 m, and recorded the
presence oi Festilyria j estiva (Lamarck, 1811), S(romi>us
oldi Emerson, 1965, and Cijpraea broderipii Sowerby,
1832, from the trawl samples. Waller (1986:39-46) de-
scribed Somalipecten crannwrorurn new genus, new
species, from "off Somalia, depth 150-300 m ", obtained
from Taiwanese fishermen, and also provided a list of
associated species. Another species recently described
from off Somalia is Vulutocorbis rosavittoriae Rehder,
1981.
Through the good offices ot John Bernard, our atten-
tion was called to an umiamed species of Vasum trawled
in Somalian waters. We take pleasure in describing this
new species in honor of Adolphe Stephant, w ho obtained
the specimens from Danish shrimpers and generously
provided Mr. Bernard with the data and specimens. Oth-
er species of moilusks reported by Mr. Stephant (in Hit.,
August 19, 1987) to have been taken during these trawl-
ing operations include h'cstihjria jcstiva, titwrnbits oldi,
S. piicatus (Roding, 1798), Phalium microstoma (von
Martens, 1901), P. bituberadosnm (von Martens, 1903),
Ficus investigatoris (E. A. Smith, 1894), Cytnatium ran-
zanii (Bianconi, 1851), Bufonaria fernandesi Beu, 1977,
Vasum crosseanum (Souserbie, 1875), Tudicula zartzi-
barica Abbott, 1958, Metula bosu:ellae Kilburn, 1975,
CucuUaca labiata (Lightfoot, 1786), and Chlamys tonn-
sendi (Sowerb>, 1895).
SYSTEMATICS
Family Turbinellidae Swainson, 1840
Subfamily Vasinae H. & .\. .\dams, 1853
Genus Vasum Roding, 1798
Remarks: See Abbott (1959) and \okes (1966) for re-
views of this subfamily.
Vasum stephanti new species
(figures 1-6)
Diagnosis: Similar to Vasum tubiferum (.-^nton, 1839)
in general appearance, but differs in having a more tri-
angular outline, three equalK well-de\ eloped columellar
plaits (in place of three major, plus one or two minor
plaits), a nearly uniformly milk-w hite shell w ith a white,
glazed aperture and parietal wall (compared to an or-
ange-brown to yellowish shell with the parietal wall a
light tan w ith \er\ large splotches of chestnut to purple-
brown), and in the presence of two or three rows of spines
at base of shell (instead of one row).
Description: Shell moderately large for genus, attaining
108-f- mm in length. Solid, heavy, turbinate, and strongly
spined. Spire ele\ated with a short, smooth, bulbous nu-
cleus of 1'2 whorls (figures 5, 6). Postnuclear whorls 7
(adult specimens lack complete spires), the body whorl
with 7 to 8 well-developed, curved to strongly recurved,
flaringly grooved and terminally open, subsutural spines.
Figures 1-6. V'a.sinn stephanti new species. 1, 2. Paratype, AMNll 225988. 3, 4. Holot\pe, AMNH 225987. 5, 6. Paratype,
AMNH 225989 (details of spire). All from type locality: off Cape Gardafui, Somalia, see text; figures 1-4 approximately x %.
figures 5, 6, x 2.
W. K. Emerson and W. E. Sage, III, 1988
Page 37
Page 38
THE NAUTILUS, Vol. 102, No. 1
A row of similar but much shorter spines below the first
row, followed by 5 coarse spiral cords and intervening
raised lines. Base of shell with 2 to 3 spiral rows of
moderateK developed, groove spines and lower surface
with weakK developed spiral lines. Parietal wall thick-
ened, slightK raised, glazed. (Columella with 3 plicae,
posterior 2 better developed; first posterior plica semi-
bifid in 1 specimen. Outer lip moderateK thin, slightK'
reflected, crenulated. Umbilicus funnel-shaped and in
most specimens widely open. Base color of shell milky
white, spire stained buff. CJolumella glazed. Aperture
white, with a slight bluish tinge. Periostracum moder-
ateK thick, tannish brown, and somewhat foliaceous. Soft
parts not seen. Operculum brown, corneous, unguiculate,
apicalK nucleate, filling most of the aperture with foot
fulK u ithdrawn. Inner surface marginally thickened on
basal and abcolumellar sides, central area depressed and
with irregular concentric rings; outer surface scabrous.
Type locality: 13-16 km east, 80-96 km south of Cape
Gardafui (Ras Asir), Somalia, trawled by shrimp fisher-
man in 183 to 220 m, December, 1986.
Range: Known only from the type localit\ and in the
Gulf of .'Kden off the Bari coast of Somalia.
Material examined: Holotype, AMNH 225987, 102.36
mm, ex E. Schelling Collection (figures 3, 4); paratype
2, AMNH 225988 (figures 1, 2), paratype 5, AMNH
225989 (figures 5, 6), paratype 8, AMNH 225990, ex J.
Bernard Collection; paratypes 1, 3, 4, 7, 9 J. Bernard
Collection; paratypes 6, 10 A. Stephant Collection, all
from the t\pe locality; referred specimen, H. Lee Col-
lection, Alula, Bari coast, Somalia (.see Table 1).
Remarks: As noted above, Vasum stephanti new species
most closely resembles in shell morphology the endemic
Philippine (Cuyo-Palawan group) species V. tubiferum
(Anton, 1839:70; Kobelt, 1876:155, pi. 9, fig. 3; Abbott,
1959:20, pi. 4, fig. 1; Spring.steen & Leobrera, 1986:105,
pi. 28, fig. 6). Anton's taxon and the closeK related V.
turbinellus (Linne, 1758:750; Abbott, 1959:17, pi. 1, figs.
2, 3), which ranges from East Africa to the western Pa-
cific, are inhabitants of shallow water, as are the other
four Indo-Pacific species assigned to Vasum {sensu stric-
to) by .Abbott (1959). One of these, V. rhinoceros (Gme-
lin, 1791 ), from Kenya and Tanzania, is somewhat similar
but has a lower-spired, heavier shell with massive nod-
ules, thickened and reflected outer lip, and a brown-
blotched to light yellow parietal wall (Abbott, 1959:21,
pi. 4, figs. 3, 4). Strongly spino.se specimens with im-
mature outer lips of the Brasilian V. cassiforme (Kiener,
1840), cited from low tide to 60 m (Rios, 1985:115), are
superficially similar to the new species. (See Abbott &
Dance, 1982:209, 210 for polychrome illustrations of these
taxa.) The Australian Altivasurn Hedle\, 1914, and sev-
eral species of Indo-Pacific Tudicula H. & A. Adams,
1864 are known to occur in moderate depths (to 220 m).
Some of the spinose species originalK assigned to Tu-
dicula (e.g., T. zanziharica Abbott, 1958, Iron) the west-
ern Indian Ocean, and T. rasilistorna Abbott, 1959), from
Table 1. Vasum stephanti new species. Shell measurements
in mm and number of whorls; width measured including spines,
n = 12. Spires incomplete e.xcept for 5, 10
l.rliUtl]
Wiiltli
= \\1
Paratype 1
Parat) pe 2
Holotype
Paratype 3
Parat) pe 4
Paratype 5
Paratype 6
Paratype 7
Paratype 8
Paratype 9
Referred
Parat\|)c II)
107 68
106.75
102.36
97.72
97.10
95.53
86.22
78.75
73.68
66.87
63.47
56,92
97 SS
97.11
82.70
82.08
84.19
77.93
73.47
64.10
57.18
67.25
52.78
,55 79
6
6
6
6
6
8'/4
6
5
5
5
5
4'/4
off Queensland, .\ustralia, may prove to be referable to
Vasum when the soft anatomy is known. The weakK
spinose V. crosseanum (Souverbie, 1875), from the In-
dian Ocean, appears to be closely related to T. rasili-
storna. The development of long spines in the species
described herein may reflect the deeper-water habitat.
ACKNOWLEDGEMENTS
We are grateful to Edward T. Schelling, Shalimar, Flor-
ida, and John H. Bernard, Crossville, Tennessee, respec-
tively, for generously donating the holotype and two
purat) pes to the American Museum of Natural Histor\-.
John H. Bernard, Harr\ G. Lee, Jacksonville, Florida,
and Adolphe Stephant, Lorient. France, kindK lent spec-
imens. We thank each of these collectors for pro\ iding
useful information. R. Tucker Abbott and M. G. Hara-
sewych oftered helpful suggestions and critically re-
viewed the manuscript. Stephen Butler, AMNH, con-
tributed the photographs.
LITERATURE CITED
Abbott, R, T. 1958. .\ new Recent species of Tudicula from
Zanzibar (Gastropoda: X'asidae). Notulae Naturae of the
Academy of Natural Sciences of Philadelphia 305:1-4.
.\bbott, R. T. 1959. The family Vasidae in the Indo-Pacific.
Indo-Pacific MoUusca 1(1): 15-32.
Abbott, R, T. and S P. Dance. 1982. Compendium of sea-
shells, E, P. Dutton, New York, ix -(- 411 p.
.•\dams, H and .-^ ,\dams. 1864. Descriptions of new species
of shells chiefl) from the Cumingian collection. Proceed-
ings of the Zoological Society ot London, for 1863:428-
435 (published April, 1864).
Anton, H. E. 1839. Verzeichniss der Concliylien welch sicli
in der Sammlung von Herrmann Eduard Anton befinden.
Halle, xvi -I- llOp
Gmelin, J. F. 1791. Caroli a Linne Systema naturae per regna
tria naturae, 13th ed Leipzig, Vol. 1, pt. 6, cl. 6. Wrmes,
p. 3021-3910.
Hedley, C. 1914 Report on the MoUusca obtained 1)\ the
F.I.S. "Endeavour " from the Great .\ustralian Bight and
from north and south of Gabo Island. Biological results of
F".I.S. "Endeavour ". Sydney, 2:65-74.
W. K. Emerson and W. E. Sage, III, 1988
Page 39
kiener, L. C;. 1840[-41]. Species general et icoiiographie des
coquilles vivantes. . . . Paris, Genre Turbinelle 6(59-71):
1-50, 1841; pis. 1-21, 1840 (Vasum cassijormc (Kiener)
dates from the citation on plate 9 of "Turhinclla ca.ssi-
formis \'alene." [= Valenciennes]; the text was pnhlislied
in 1841).
Kobelt, H. C 1876. Systematisches Conchylien-Cabinet von
Martini und Chemnitz. ... Niirnberg. Purpur.schnecken,
Band 3, Abt. 3a, pt. 251:121-164.
Linne. C, von. 1758. Systema naturae per regna tria naturae,
10th ed. Stockholm, \ol. 1, Regnum animale, 284 p.
Lorenz, F , Jr. 1987 Description of a new Ovulidae species
from Somalia (Gastropoda: Ovulidae). La Conchiglia, Rome
19(214-215):11-12.
Rios, E. C. 1985. Seashells of Brazil. Fundagao Sociedade do
Rio Grande, Funda^ao Universidade do Rio Grande, Mu-
seo Oceanografico. 329 p., 102 pis.
Souverbie, S. M. 1875. Description d une espece nouvelle
appartenant au genre Titrhinella. Jonrnal de (>'onchylio-
logie 23(4):297-298.
Springsteen, F. J. and F. M. Leobrera. 1986. Shells of the
Philippines. Carfel Seashell Museum, Manila, 377 p.
Vokes, E. H. 1966. The genus Va,si(ni (Mollusca: Gastropoda)
in the New World Tulane Studies in Geology 5(l):l-36.
Waller, T. R. 1986. A new genus and species of scallop (Bi-
valvia: Pectinidae) from off Somalia and the definition of
a new tribe Decatopectinini. The Nautilus 100(2):39-46.
THE NAUTILUS 102(1 ):40-45, 1988
Page 40
Density, Spatial Distribution, Activity Patterns, and
Biomass of the Land Snail, Geophorus bothropoma Moellendorff
(Prosobranchia: Helicinidae)
Kurt Auffenberg
Troy Auffenberg
Malacology Division
Florida State Museum
University of Florida
Gainesville, FL 32611
ABSTRACT
The results of preliminary work done in the Republic of the
Philippines during July, 1981 on the density, spatial distribu-
tion, activity patterns, and biomass of the Philippine Island
land snail Geophorus bothropoma (Prosobranchia, Helicinidae)
are presented.
Density was estimated at 0.55/m^ utilizing the quadrat meth-
od. Using the parameters of mean crowding and patchiness,
the species was shown to be uniformly distributed spatially,
possibly due to mutual repulsion. Individual activity distance
per 24 hour period varied greatly (0.0-275.0 cm) and was
possibly correlated with rainfall Utilizing the convex polygon
and ellipsoid methods, minimum individual activity ranges for
the duration of the study were estimated to be 2.4-4.2 m^.
Biomass (live weight) of adults was estimated at 0.13 g/m'^.
The species is diurnal and does not home.
Key words: Gastropoda; Prosobranchia; Helicinidae; Geopho-
rus; density; spatial distribution; activity; biomass; Philippine
Islands.
INTRODUCTION
There are relatively few published studies dealing with
the density, spatial distribution, activity patterns, or bio-
mass of land snails (see Discussion section for citations).
Of those that do exist, almost all have dealt with pul-
monates of temperate regions, while tropical terrestrial
prosobranchs remain virtually unstudied. The present
paper reports on the results of a preliminary study con-
ducted July 7-17, 1981 on Geophorus bothropoma Moel-
lendorff, 1895 (Quadras & Moellendorff, 1895:148; Wag-
ner, 1908:152, pi. 29, figs. 20-23), a large (13.0-16.0 mm
in diameter), limestone rock-dwelling helicinid endemic
to the Caramoan Peninsula, Luzon, This represents one
of the very few times quantitative ecological methods
have been utilized for the study of terrestrial mollusks
in all of Southeast Asia. In addition, the present paper
introduces to ecological malacology several analyses used
by zoologists working in groups other than mollusks (i.e.,
birds, Odum & Kuenzler, 1955; frogs, Turner. 1960; in-
sects, Alexander, 1961, Iwao, 1970; mammals, Stumpf &
Mohr, 1962). While the database with which we worked
was rather small, the ease ot anaKses and the applica-
bility of the results suggest that such approaches to sim-
ilar questions in terrestrial snail ecolog) would be highly
beneficial.
STUDY AREA
The study area is located in the Republic of the Phil-
ippines, Luzon Island, Camarines Sur Province. Cara-
moan Municipality, 1.0 km south of Barrio Ilawod. The
work was conducted 10 m off the trail between Ilawod
and Gota Beach on a large (8 x 9 x 3 m high) limestone
boulder with almost vertical sides.
Much of the Caramoan Peninsula consists of well-
developed limestone karst of Miocene reef origin. Be-
cause the area has been relativeK recently uplifted, the
mountains are rather steep. Hills and ridges often consist
of masses of large limestone boulders with chasms tens
of meters deep. Runoff is rapid and though rainfall may
be high seasonally, the surface environment is often rel-
atively dry. Surface water collects in solution pits on
limestone and in hollows of trees.
Due to the rugged topography, the area remains blan-
keted in a multi-canopied, tropical evergreen forest.
Vegetational communities are extremely diverse. The
forest is dominated by tall dipterocarp trees in both upper
(30-40 m) and lower (5-20 m) canopies; a dense her-
baceous layer of mainK ferns exists near the ground
surface.
The Caramoan Peninsula has no definite dry season.
While it tends to rain throughout the \ear (ca. 2,900
mm), most rainfall occurs from September through Jan-
uary, with a minor peak in July associated with the onset
of the typhoon season.
As in other tropical rainforests, the Caramoan Pen-
insula shows little seasonal temperature variation. Mean
K. Auffeiiberg and T. Auffenberg, 1988
Page 41
iluil\ temperature fluctuations are greater tPian mean
seasonal variation. However, there is a marked vertical
tliermocline within the forest, in which temperatures
from the surface to 1 m are most stable; daily variation
increases at greater heights above the ground. Due to
the dense vegetation at the study site, the effects of solar
railiation and wind are minimal. Therefore, Geophorus
butliropoma lives in the most stable part of the local
warm, moist tropical forest environment. The above eco-
logical data is from Auffenberg (in press).
METHODS
An 8 m- (2 m tall, 4 m wide) vertical grid of 1 m- quadrats
was constructed against the east side of the boulder to
measure density and activity. All adult Geophorus both-
ropoma within this grid (N = 14) were numbered with
small dots of fingernail polish on the dorsal surface of
the shell. This increased shell visibilit\ , but is not believed
to have increased predation due to the nocturnal habits
of most potential predators. No additional individuals
were located during subsequent visits. The site was vis-
ited nine times during the next 11 days at 10 a.m. for
appro.ximateK 1 hour observations. In addition, the site
was visited sporadically throughout the daylight hours
and seven times at night (8-10 p.m.). This provided a set
of successive censuses, not a random quadrat sampling.
Densit\ estimates are based on the ciuadrat sampling
method. This method simply involves counting individ-
uals occurring within a quadrat of fixed size.
Analyses of spatial distribution are based on m* (mean
crowding) and m* m (patchiness) of Lloyd (1967). Mean
crowding is given by the formula:
q
- 1
(1)
where x, = the number of individuals in the jth quadrat
and q = the total number of quadrats.
Mean crowding is defined as the mean number of other
individuals per quadrat per individual. It expresses the
degree of spatial crowding experienced by an individual
because of others of the same species (or different species;
see Lloyd, 1967). A particularly important feature of this
statistic is that it is relatively independent of spatial dis-
tribution t\ pe, number of samples, and size of the means.
The most complete discussion concerning the utility of
this statistic is by Iwao (1977). It should be stressed that
this statistic does not measure crowding in an ecological
sense, as man\' variables (i.e., food availabilit\', territo-
rialit) ) must be examined before this can be determined
(Lloyd, 1967).
Patchiness (m* /m) is the ratio of mean crowding to
mean densit\ (Iwao, 1968). This index provides a relative
measure of aggregation. It equals unity (1) in random
distribution, is greater than unity in contagious distri-
butions and is less than unity in regular (uniform) dis-
tributions (Iwao, 1977).
Anal) sis of the activity of Geophorus buthropoma was
made by subdi\idiiig the I m- quadrats, plotting the
exact location of individuals and measuring straight line
distances (nearest cm) from the last observation with a
flexible tape. This provided a crude estimate of the min-
imum distance tra\eled since the last ob.servation. Cal-
culations are based only on data provided by two or more
consecutive daily observations (N = 27).
Estimations of activity ranges (N = 5) are based on
the convex polygon method and the ellipsoid method,
which is based on a covariance matrix of capture loci
(Jeunrich & Turner, 1969). The convex polygon method
involves plotting all the recapture points, drawing the
smallest convex polygon containing these points and then
determining the area. This method, defined by the equa-
tion:
A =
x.y^+i - Xi+iYi
(2)
is relatively simple but is biased b\ sample size. The
values tend to increase as the number of capture points
increases.
The ellipsoid method is not biased by sample size and
assumes that individuals are active outside the observed
range of activity (see Jennrich & Turner, 1969, for dis-
cussion). This method is defined by;
67r|S|
(3)
where |S| is the determinant of the capture point co-
variance matrix;
S =
defined by the equations;
S S
s s
(4)
1
x;^
S„
n - 2
o 2 (y. - y)^
S. = S,. = — !— 2 (X, - x)(y,
n — 2 "
X - - 7v X, ,
y = - 2j y.
(5)
Live adult snails (N = 90) were collected near the
study site and their weights (in groups of 10 individuals)
were taken to the nearest milligram on a single beam,
double pan scale. Fifty specimens were then macerated;
the shells and opercula were rinsed, dried for several
days and reweighed.
Voucher specimens of Geophorous bothropoma are
housed in the mollusk collection of the Florida State
Museum (UF 56667).
Page 42
THE NAUTILUS, Vol. 102, No. 1
Table 1. Densit\ and spatial distribution parameters based on quadrat method, n = individuals recaptured per visit, m/m-
(lciisil\, m* = mean crowdiiij;, m* m = patchiness. Visit 6 is (irnitted from densit\ and spatial distribution analyses
Visits
X
SE
1
2
3
4
.5
6
7
8
9
SD
n
9
6
5
5
2
1
3
4
3.89
m/m"
1.13
0.75
0.63
0.63
0.25
—
0.13
0.38
0.50
0.55
0.11
0.32
m*
1.11
1.00
40
1 20
00
—
00
67
0.00
0.55
018
0.52
m*/m
0.98
1.33
6-1
1 90
(100
—
1 ) 1 )( 1
1.76
0,00
83
0.28
0.79
RESUl/rS
Density and spatial distribution: Froiumnct'd \ariatiun
in population density was observed during this study
(table 1). Densities were calculated in eight quadrats for
nine visits (N = 72 quadrats). \'isit 6 yielded no recap-
tures and is omitted from density and spatial distribution
analyses. Depending on the individual visit analyzed,
mean crowding and patchiness reveal either random,
contagious, or uniform spatial distributions. However,
the means of these statistics suggest that individuals are
uniformly distributed, for the averages of both mean
crowding and patchiness are less than unity.
The regression of mean crowding on mean density
(figure 1 ) also suggests a uniform spatial distribution (« =
y intercept = —0.14, /3 = slope = 1.25) with possible
"mutual repulsion" (see Iwao, 1968, for discussion of
regression analysis).
Activity: Marked individuals were observed to be active
during all da\ light hours. None were active during visits
alter dark. .\ typical activit\' sequence consisted of an
individual moving along a meandering path for several
minutes, sometimes feeding on algal growth, stopping
for several minutes, then continuing. Some individuals
1.8 2.0
remained at the same exact location tor 1 or 2 da\s, while
for no apparent reason others remained active.
Fifty-seven recaptures were recorded (50.9^ of pos-
sible recaptures). Distances traveled per day varied greatly
between individuals (0.0-275.0 cm). Because distances
traveled rarely conform to normal distributions only the
range of distances is given.
Population activity variation was possiblv correlated
with rainfall. Light rain did not initiate activity in in-
active individuals, nor did it seem to affect individuals
already active. However, snails ceased moving about if
the rainfall became intense. Above average activity levels
were observed between July 8 and 11 (50.0-275.0 cm,
N = 11 recaptures, three visits) following intense rainfall
associated with two small typhoons in the area. This in
turn was followed by 5 days (July 12-16) of relatively
little rainfall and reduced snail activity (0.0-65.0 cm,
N = 10 recaptures, four visits). A heavy rain fell during
the night of July 16, resulting in a marked increase in
snail activity on July 17 (65.0-203.0 cm, N = 6 recap-
tures, one visit).
Perhaps the most interesting observation concerning
the activity of this species is that it does not home. In-
dividuals simply wander from one resting site to another,
usually shallow depressions or crevices in the rock. They
avoid resting sites in leaf-filled solution pits, which are,
however, usually inhabited bv Cijclophorus ceratodes
Moellendorff, 1895, Japonia ciliata (Sowerby, 1843), ].
stephanophora (Moellendorff, 1895), and several small
pulmonate species (personal observation).
2-
,^
j
%^
1 ' ^
\
\
1-
4=-
■:::y^cT^^---
_...^.....,;»
r^ a
1 .--
Fipurp I . Rejiression of mean density (ni, nr) on mean crowd-
ing (ni*) of Gi'(>i)li(inis hothropoma in study area. Curved lines
represent 95% confidence limits.
12 3 4
Figure 2. Five representative activity ranges of Geophorus
bothropojna plotted on 8 m- grid HIack dots represent recap-
ture sites. Each polygon depicts an activity range of an indi-
vidual snail ov er an 1 1 day period.
K. Auffenberg and T. Auffenberg, 1988
Page 43
Estimates of individual activity ranges (N = 5) (figure
2) are quite consistent. The convex polygon method (cor-
rected for sample size bias; Jennricli & Turner, 1969)
yielded activity ranges of 2.3-3.8 m- (x = 3.1 ± 0.3,
SD = 0.5 m-). the ellipsoid method yielded slightly (but
insignificantly), larger area estimates of 2.7-4.2 m- (x =
3.3 ± 0.3, SD = 0.6 m-).
Biomass: Mean total live weight per adult individual
in the study area was 0.59 g (N = 90). Mean shell weight
(including operculum) per individual was 0.35 g (N =
52), yielding a mean live biomass per individual of 0.24
g. The mean density estimated by the quadrat method
(table 1) of this species in this area is 0.55 individuals/
m-. Thus the estimated biomass is 0.13 g/m-.
DISCUSSION
Density and spatial distribution: Previous studies on
species comparable in size to Geophorus bothropoma
have been few (i.e.. Mason, 1970; Richardson, 1975;
Cameron, 1982), and none concerns terrestrial proso-
branchs. The only prosobranchs with available data are
the small hydrocenid Georissa monterosatiana (Godwin-
.•\usten & Nevill, 1879) and the diplommatinid Opis-
thostoma retrovertens Tomlin, 1938 of Malaysia. Berry
(1966) found these species occurring in higher densities
on moss covered rocks than on moss-free rocks, but did
not address spatial distribution. In the Caramoan area
we found Geori.'isa rufescens (Moellendorff, 1887) in ex-
tremely localized and highly aggregated populations on
vertical rock faces. This is very different from the large
Geophorus bothropoma, which is found on virtually every
vertical rock face and is uniformly distributed spatially.
In this stud\-, intersample variation in estimated mean
densities is probabK largely due to the limited size of
the study area. The choice of study area size and number
of samples must have some biological basis to estimate
population density and spatial distribution adequately.
This is usually done by preliminary sampling or guessing
(Ivvao, 1977). However, we do feel confident that the
proper quadrat size (1 m^) was chosen because the ob-
served dail\ activity was approximately 1 m (x = 93.7 ±
13.31 cm). Density estimates were possibly also biased
by the rugged microhabitat. The limestone karst provides
man> cracks, crevices, and solution pits in which a snail
could remain unseen, despite our thorough searches.
Utilizing Iwao's (1968) interpretation of m*/m regres-
sions, we determined that individuals of Geophorus both-
ropoma were uniformK distributed and may have ex-
hibited "mutual repulsion. Infra-specific competition
has never been properly substantiated in studies on ter-
restrial mollusks and we cannot substantiate it here.
Nevertheless, density was relatively low, despite the
seemingly adequate food and resting site resources that
could have been expected to sustain substantialK' higher
snail densities.
.Activity: Previous studies have shown terrestrial mol-
lusks to be very cyclic in their activity patterns (i.e..
Barnes & Weil, 1944, 1945; Wells, 1944; Dainton, 1954a,b;
Blinn, 1963; Henne, 1963; Cameron, 1970; Crawford-
Sidebotham, 1972; Baker, 1973; Bailey, 1975; Shachak
et al., 1975; Heatwole & Heatwole, 1978; Deisler, 1987).
It is apparent from these studies that the initiation and
continuation of activity is a variable response to complex,
simultaneous climatic changes (some minute, perhaps
immeasurable) in the microenvironment. More field and
laboratory studies must be undertaken before this will
be understood. In previous studies snail movements were
most numerous shortly before or after nightfall and short-
ly before dawn. However, Geophorus bothropoma is
diurnal rather than crepuscular, and the congener G.
trochiformis (Sowerby, 1842) was also observed active
throughout the day. We believe this behavioral pattern
is not a general trend in tropical terrestrial prosobranchs.
Cyclophorus ceratodes Moellendorff, 1895 and Japonia
ciliata (Sowerby, 1843) and J. stephaiiophora (Moellen-
dorff, 1895) living in the same study area are decidedly
crepuscular in their activity patterns (personal observa-
tion).
Few data are available on activity and home site ranges
of terrestrial mollusks. Heatwole and Heatwole (1978)
found individual home site ranges of the camaenid Car-
acolus caracoUus (Linne, 1758) to vary greatly (0.08-
59.0 m-). Since Geophorus bothropoma does not home
we prefer to call the area enclosed by the recapture sites
(figure 2) the activity range. The term homesite range
implies that the individual possibly returns to a certain
area after foraging outside that area. We assume that
the activity range of Caracolus caracoUus is much larger
than the enclosed polygon determined by recapture sites.
Long-term study is needed to provide a meaningful
estimate of the activity range of Geophorus bothropoma.
The data presented here only provide an estimate of the
minimum size of the activity range expected during a
wet part of the year. Activity ranges are probably sig-
nificantly smaller during the dry season.
Homing is apparently based on chemoreception (see
Cook, 1979, for review). The advantages and/or disad-
vantages of homing have not been properly addressed.
A permanent resting site should provide relative safety
from predators and fulfill the physiological requirements
of the individual at rest. The physiological and/or be-
havioral basis for Geophorus bothropoma being unable
to (or not having to) home is not known. Although the
operculum provides protection from small predators and
desiccation, this does not explain this species' lack of
homing behavior. Geophorus bothropoma most com-
monly rests on vertical rock faces. Even when inactive,
it does not retract fully into the shell, leaving the oper-
culum non-functional. Because Cyclophorus ceratodes
Moellendorff, 1895 and Japonia ciliata (Sowerby, 1843)
and J. stephanophoru (Moellendorf. 1895) were found
to return to the same solution pits after foraging (personal
observation), non-homing is not a general trend in trop-
ical terrestrial prosobranchs.
Biomass: Few data on the biomass of terrestrial mol-
lusks are available (Strandine, 1941; Jennings & Bark-
Page 44
THE NAUTILUS, Vol. 102, No. 1
ham, 1975; Richardson, 1975; Cameron, 1982) and none
concern prosobranchs, nor do they pro\ ide eas\ or direct
comparisons. Man\ more species must be studied before
general trends can be demonstrated.
Fragments of one marked specimen were found in a
solution pit, possibly preyed on by the nocturnal shrew,
Suncus marinus (Linne, 1766), an introduced species.
This species was seen on two occasions at the study site.
Many shells of Geophorus bothropoma gnawed open at
the periphery were found in solution pits nearbs . The
diurnal, snail-eating monitor lizard Varanti.s grayi Bou-
lenger, 1885 rarely preys on this species (Auffenberg, in
press).
Although short-term data such as those presented here
add much to our knowledge of this species, onl\' with
long-term life-histor\ studies will ([uestions be answered
with confidence. Combined with morphological infor-
mation, sound behavioral and ecological data uill un-
doubtedly contribute greatK to a better understanding
ot the systematics and e\olution of Philippine helicinids.
ACKNOWLEDGEMENTS
We would like to thank the U.S. Fish and Wildlife Service
and the New York Zoological Society for herpetological
grants to Walter Auffenberg that allowed us the oppor-
tunit) for the present stud>. The Philippine Parks and
Wildlife office was very helpful in issuing appropriate
permits. The illustrations were drawn by Armando Gar-
cia and Marcos Guinaldo. We appreciate the patience,
time, and insight provided bv Irvy Quitmyer, Mark
Brenner, and Michael W. Binford of the Florida State
Museum during the computer assisted regression anal-
ysis. Glenn A. Goodfriend provided suggestions during
early data analysis. Robert .\. (>ameron provided many
constructive suggestions for analyses and format. We also
thank Arthur J. Cain, Robert Hershler, and Fred G.
Thompson for reviewing the manuscript. Special thanks
are extended to our father, Walter Auffenberg, for his
tireless assistance in the field and for his time and guid-
ance during the writing of the manuscript.
LITERATURE CITED
Alexander, R D. 1961, .Aggressiveness, territoriality and sex-
ual behaviour in field crickets (Orthoptera: (iryllidae). Be-
haviour 17:130-223.
Auffenberg, W. In press. The behavioral ecology of Gray's
Monitor Lizard. University Presses of Florida
Bailey, S. R. 1975. The seasonal and daiU patterns of loco-
motor activity in the snail Helix asperse Muller, and their
relation to environmental variables. Proceedings of the
Malacological Society of London 41:415-428.
Baker, A. N. 1973. Factors contributing towards the initiation
of slug activity in the field Proceedings of the Malaco-
logical Society of London 40:329-333.
Barnes, H. F. and J. W. Weil 1944 Slugs in gardens: their
numbers, activities, and distribution Part 1 Journal of
Animal Ecology 13:140-175
Barnes, H. F. and j VV Weil 1945. Slugs in gardens: their
numbers, activities, and distribution. Part 2. Journal of
.■\nimal Ecology 1471-105
Berry, .A J 1966 Population structure and Quctuations in the
snail fauna of a Malayan limestone hill. Journal of Zoology,
London 150:1 1-27.
Blinn, W. C. 1963. Ecolog\ of the land snails Mesodon thy-
roidus and Allogona profunda. Ecology 44(3)498-505.
Cameron, R. A. D. 1970. The effect of temperature on the
activity of three species of helicid snail (Mollusca: Gastrop-
oda). Journal of Zoolog\ . London 162:303-315.
Cameron, R A D 1982 Life histories, densit\ and biomass
in a woodland snail communit\ . Journal of Molluscan Stud-
ies 48:159-166
Cook, A. 1979. Homing in gastropods. Malacologia 18:315-
318.
Crawford-Sidebotham, T. J. 1972. The influence of weather
upon the activity of slugs, Oecologia 9:141-154.
Dainton, B H. 1954a. The activit\ of slugs. I The induction
ot activity b) changing temperatures. Journal of Experi-
mental Biology 31:165-187.
Dainton. B H. 1954b The activity of slugs. II, The effect of
light and air currents. Journal of Experimental Biology 31:
187-197.
Deisler, J. E. 1987. The ecology of the Stock Island Tree Snail.
Orthalicus reses reses (Say). Bulletin of the Florida State
Museum, Biological Sciences 31(3):107-145.
Heatwole, H. and A. Heatwole. 1978. Ecology of the Puerto
Rican camaenid tree-snails Malacologia 17(21:241-315.
Henne, F. C, 1963. The effect of light and temperature on
the locomotorv activit\' of Polygyra albolahris (Sa\ ), Bios
34:129-133,
Iwao, S. 1968, A new regression method for the aggregation
pattern of animal populations. Research in Population
Ecology 10:1-20.
Iwao, S. 1970. Analysis of contagiousness in the action of
mortality factors on the Western Tent Caterpillar popu-
lation b\ using the m*-m relationship. Research in Pop-
ulation Ecology 12:100-110,
Iwao, S, 1977, The m*-m statistics as a comprehensive method
for analyzing spatial patterns of biological populations and
its application to sampling problems. In: Morisita. M (ed).
Studies on methods of estimating population density, bio-
mass and productivity in terrestrial animals. JIBP Synthe-
sis, University of Tokyo Press 17:21-46.
Jennings. T. J and J P Barkham. 1975, Slug populations in
mixed deciduous woodland. Oecologia 10:279-286.
Jeiuirich, R. I, and F B. Turner. 1969 Measurement of non-
circular home range. Journal of Theoretical Biology 22:
227-237,
Lloyd, M, 1967, 'Mean Crowding Journal of ,\niinal Ecolog)
'36(l):l-30,
Mason, C. F. 1970. Snail populations, beech litter production,
and the role of snails in litter decomposition. Oecologia 5:
215-239.
Odum, E, P, and E. J Kuenzler, 1955, Measurement of ter-
ritory and home range size in birds. .\uk 72:128-137,
(Quadras, J, F, and O, F, von Moellendorff 1895, Diagnoses
specierum novarum ex insults Philippinis auctoribus,
Nachrichtsblatt der deutschen Malakozoologischen Ge-
sellschaft 1895(9-10):137-149.
Richardson, A. M. M. 1975, Energy Dux in a natural popu-
lation of the land snail, Cepaea nemoralis Linne. Oeco-
logia 19(2):141-164.
Shachak, M., Y. Orr. and Y. Steinberger. 1975, Field obser-
vations on the natural history of Sphincterochda (S.) zona-
K. Auffenberg and T. Auffenberg, 1988
Page 45
ta (Bourguignat, 1853) (=S. boissieri Charpentier, 1847).
Argamon, Israel Journal of Malacology 5(l-4):20-46.
Strandine, E J. 1941. Quantitative study of a snail population.
Ecology 22:86-91.
Stumpf, W. A. and C. O. Mofir. 1962. Linearit\ of home
ranges of California mice and other animals Journal of
Wildlife Management 26:149-154.
Turner, F. B. 1960. Population structure and dynamics of a
Western Spotted Frog, Rana p. pretiosa Baird and Girard,
in Yellowstone Park, Wyoming. Ecological Monographs
30:251-278.
Wagner, A. 1908, Die Familie der Jlelicinidae. In: Syste-
malisches Conchylien-Cabinet von Martini und Chemnitz
ia8):113-184.
Wells, G. P. 1944. The water relations of snails and slugs. III.
Factors determining activity in Helix pomatia L. Journal
of Experimental Biology 20:79-87.
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T H E t7N AU T I L U S
CONTENTS
Volume 102, Number 2
April 29, 1988
ISSN 0028-1344
Joseph R. Pawlik
John B. O'Sullivaii
M. G. Harasewych
The egg capsules, embryos, and larvae of Cancellaria
cooperi (Gastropoda: Cancellariidae)
Martin Avery Snyder
Latirus nuirtini (Gastropoda: Fasciolariidae), a new species
from Honduras
54
Robert Hershler
Lee-Ann C. Havek
Shell variation of springsnail populations in the Cuatro
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Limnocrene Fauna
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Eva Pip
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Fred G. Thompson
Harry G. Lee
Hypselostoma holimanae new species, a pupillid land snail
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Manuel llaimovici
Eledone gaucha, a new species of eledonid octopoti
(Cephalopoda: Octopodidae) from southern Brazil
82
88
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THE NAUTILUS 102(2):47-53, 1988
Page 47
The Egg Capsules, Embryos, and Larvae of Cancellaria cooperi
(Gastropoda: Cancellariidae)
Joseph R. Pawlik
Scripps Institution of Oceanography
UniversitN of California, San Diego
La Jolla, CA 92093, USA
John B. O'Sullivan
Monterey Bay Aquarium
886 Cannery Row
Monterey, CA 93940
M. G. Harasewych'
Department of Invertebrate Zoology
National Museum of Natural History
Smitfisonian Institution
Washington, DC 20560, USA
ABSTRACT
The egg capsules, and the embr\onic and larval development
of the cancellariid gastropod Cancellaria cooperi Gabb are
described. Egg capsules are spatulate in form, having long,
narrow stalks that support the eggs above the surrounding sand.
Egg capsules contain 4,000-5,000 eggs (165 ^ni in diameter),
which undergo typical prosobranch development to hatch as
planktotrophic veligers after 27 da> s at 15 °C Larvae in culture
grew from 305 ^m to 890 ^m in shell length over 30 days, but
died before metamorphosis. Limited comparative data suggest
that long stalked egg capsules are known only in members of
the Cancellariinae, that opercula, absent in all adult Cancel-
lariidae, are present or prominent in the late larval stages of
at least some species, and that developmental type cannot be
inferred from protoconch morphology using the criteria of Shu-
to (1974) in a majority of cancellariid species.
Key words: Reproduction; development; larvae; eggs; egg cap-
sules; Cancellariidae; Cancellaria.
INTRODUCTION
Little is known about the reproductive biology and early
development of most of the approximately 200 species
that constitute the neogastropod family Cancellariidae.
The few published reports (Morch, 1869; Thorson, 1935,
1944; Knudsen, 1950; MacGinitie, 1955; Kilburn & Rip-
pey, 1982; Bouchet & Waren, 1985) are limited to de-
scriptions of egg capsules and, in some cases, ova or larval
shells attributed to cancellariids, usually on the basis of
the proximit\' of living snails.
During studies of the diet and feeding behavior of
Cancellaria cooperi Gabb, 1865, a species that is attract-
ed to and speciBcall) parasitizes the Pacific electric ray
Torpedo calif arnica .\>res, 1855 (O'Sullivan e/ ai. 1987),
a number of these snails were observed producing egg
capsules. The present study supplements our know ledge
of the natural history of this cancellariid with descrip-
Author for correspondence.
tions of its egg capsules and embryonic and larval de-
velopment, and reviews the available data on the
reproductive biology and larval development of the Can-
cellariidae.
MATERIALS AND METHODS
Twenty-three specimens of Cancellaria cooperi were col-
lected on the artificial reef "Torrev Pines #1", off San
Diego, CA (32°53'12"N, 117°50'50"'w) at depths of 20-
22 m using SCUBA. The animals were maintained in an
aquarium containing sufficient sand for complete snail
burial (4-8 cm depth) and supplied with a continuous
flow of seawater (12-16 °C). Snails were allowed to feed
on electric rays on a bimonthly basis, and had been
maintained in this manner for at least 6 months prior to
the onset of oviposition.
Individual egg capsules were freed from the aquarium
bottom and maintained in beakers containing continu-
ously aerated, 1 ixm filtered seawater at 15 °C. Developing
embryos were removed with a glass pipette through an
incision cut along the narrow edge of the egg capsule.
Hatched larvae of Cancellaria cooperi were cultured
following procedures described by Paige (1986), except
that 1 /um filtered natural seawater containing 40 mg/
liter each of the antibiotics streptomycin sulfate and so-
dium penicillin G was used instead of artificial seawater.
Larvae were fed a mixture of the green flagellates Iso-
chrysis galhana (Park, 1949) and Pavlova lutheri (Droop)
Green, 1975 at 10^ cells/ml. Prior to photography, larvae
were narcotized in a 1:3 mixture of a saturated aqueous
solution of chlorobutanol in seawater.
Eggs and larvae for SEM examination were fixed in
2% formalin in seawater, and stored in 70% ethanol.
Specimens were critical-point dried and photographed
using a Hitachi S-570 Scanning Electron Microscope.
RESULTS
Beginning April 21 and continuing through Ma\ 19,
1986, a total of 15 egg capsules were laid by at least
three snails (mean shell length = 62.3 mm), with nine
capsules being produced sequentially b>- a single female.
Page 48
THE NAUTILUS, Vol. 102, No. 2
Figure 1. Cancellaria cooperi Gabb. Apertural and right side
view of female specimen collected by SCUBA in 20-22 m at
artificial reef "Torrev Pines #1", off San Diego, CA {32°53'12"N,
!17°50'50"W), USNM 846054. 1.0 x.
Oviposition generally occurred at night or in the morn-
ing. Snails emerged partialK from the sand, with their
raised foot spread anteriorK and enfolded posteriorly,
the emerging stalk protruding from the folded portion
of the foot. Over the course of several hours, the egg case
emerged from the ventral pedal gland and was released,
whereupon the stalk straightened and the egg case was
supported well above the surface of the sand.
The spatulate capsules (figure 2; table 1) consisted of
lenticular egg cases supported on long, narrow (250 fim
diameter) stalks. Egg capsules were roughly rectangular
in transverse section, with narrow keels running along
the outer edge of each corner. A pre-formed hatching
aperture, containing a membranous plug (figure 2, mp),
was situated at the uppermost end of each egg case.
Capsules were affixed to the bottom ot the arjuarium by
holdfasts that spread from the base of the stalk.
Each egg case contained an estimated 4,000-5,000
spherical eggs (figures 3, 4; diameter = 164.5 ± 6.7 ixm,
N = 10), suspended in a clear, gelatinous matrix, all of
mp
Figure 2. Egg capsule of Cancellaria cooperi Gabb. Frontal
and side views 1.0 x. detail of hatching aperture 6.0 x. mp,
mucous plug; li, membranous sack.
which was enveloped in a membranous, transparent sack
(figure 2, li). The lenticular walls of the egg case were
slightly concave, creating a constriction along the mid-
line of the case that displaced the eggs to either side.
Spiral cleavage commenced within 12 hours of ovi-
position, and the 8 cell stage (figure 5) was reached by
the second day. Thereafter, the embryos became in-
creasingly irregular in form (figures 6-8). All the embryos
within an egg case underwent de\elopment; there were
no nurse eggs or unfertilized eggs in the cases examined.
By 10 da\s after deposition, the stomodeal in\agination
was e\ident (figure 9). .\fter 12 da\s, the protoconch and
operculum, both of conchiolin, were clearly discernable
(figures 10, 11), and torsion was complete after 14 days
(figures 12-14). The velar lobes were formed and in-
creasing in size by the 16th day (figures 15, 16). On the
Figures 3-14. Embryonic and larval development of Cancellaria cooperi Gabb at 15 °C. 3. Living etnbr\os, 1 day old 4. One-
(l.i\-(]ld embryo, critical-point dried. Scale bar = 25 ^m. ,5. lJ\iiig embrxos, 2 days old. 6. Living embr\o, 4 days old. 7. Four-
da\-old embryo, critical-point dried Scale bar = .50 fim. 8. Living embr\os, 8 days old. 9. Ten-dav-old embr\o. critical-point
dried. Scale bar = 50 /nm 10. Li\ing embryos, 12 days old 1 L Hight side and ventral views of same 12-da\-old embr\o, critical-
point dried. Scale bar = 50 ^m. 12. Living embryos, 14 days old \'.i. 14. X'cntral (13) and dorsal (14) views of 14-da\-old embrsos,
critical-point dried. Scale bar = 50 ^m
asr, apical sensory region; dml, dorsal maiille lip; Ik, larval kidney; o, operculum; pb, polar body; pc, protoconch; pt, prototroch;
s, stomodeum.
J. R. Pawlik et ai, 1988
Page 49
Page 50
THE NAUTILUS, Vol. 102, No. 2
Figures l,'j-22. Embryonic and lar\al development of Cancellaria cooperi Gabb at 15 °C. 15. Living embryo, 16 days old 16.
Sixteen-da\-old t'nibr\o, critical-point dried. Scale bar = .50 ;um. 17. Li\ing. newK -hatched larva with carmine particles in the
gut. 18. Shell ultrastrticture of newK-hatched larva, plane of fracture parallel to growing edge. Scale bar = 2 nm 19. .^pertural,
dorsal and apical views of shells of newly-hatched larvae. Scale bar = 100 ^in. 20. Living larva, 30 days after hatching 21. Shell
and operculum of larva, 30 days after hatching. Scale bar = 250 fim. 22. .-Vpical portion of adult shell (LACM 40-95.1), arrow-
indicating transition from protoconch to teleoconch, scale bar = 500 fim.
mo, month; o, ()j)crculuni; po, posterior ciliary band; pr. preoral ciliar\ band; sto, storiiach; vl, velar lobe.
20tli day after capsule depobitiou, eiiibr\ os began to move
more freely within the egg case. The gelatinous matrix
in which the developing embrvds had been suspended
became less viscous, and the swimming enil)ryos aggre-
gated randondy w illiin the case. ,\ftor an average of 27
da\s at 13 °(.' the nicnibrancjus plug occluding the hatch-
ing aperture dissolved, and swimming larvae escaped.
Shells ol the new K hatched veligers (figures 18, 19) had
a mean diameter of 304,6 /um (N = 10, SD = 3.5), and
were 3.9 ^m thick. The veligers possessed an operculum
J. R. Pawlik et a/., 1988
Page 51
Table 1 . Measurements of egg capsules of Cancellaria cooperi
Gabb. All measurements in mm (N = 10)
Character
Mean
Range
SU
Total length
79.6
70.5-S3.6
4.8
Capsule length
30.1
27.8-32.8
1.9
Capsule width
11.4
10.7-12.4
0.6
Capsule thickness
1.8
1.6-2.0
0.2
_i
Stalk length
49.5
37.7-54.0
6.0
O
>
Maximum diameter
of holdfast
9.1
7.4-lO.N
1.2
(figure 19, o), and a fully functional alimentary system.
When added to a small dish ot seawater containing sus-
pended carmine particles, the veliger larvae rapidly filled
their guts with these particles (figure 17, sto).
Attempts to rear these veliger larvae through meta-
morphosis were unsuccessful. At 20 °C, larvae grew rap-
idly, reaching a mean shell diameter of 644 ^m (figure
20) 22 days after hatching. Thereafter, the larvae began
to die, with only a single larva surviving 30 days after
hatching (shell diameter = 890 nm. figure 21). Larvae
that had survived more than 22 da\s after hatching de-
veloped propodia, but did not metamorphose in the pres-
ence of fresh or frozen Torpedo mucus, or in the presence
of sand from an aquarium containing adult snails. No
crawling or search behavior associated with the onset of
metamorphic competence was observed. Protoconchs of
adult shells of Cancellaria cooperi indicate that the larval
shell reaches 3.3 whorls (shell diameter = l,160;um) prior
to metamorphosis (figure 22; table 2).
DISCUSSION
Egg capsules of Cancellaria cooperi resemble those of
man\ neogastropods (e.g.. D'Asaro, 1970; Radwin &
Chamberlin, 1973; Bandel, 1976), with notable modifi-
cation in the length of the supporting stalk. This elon-
gation of the stalk appears to be an adaptation for un-
stable sediments in which capsules are deposited. Under
natural conditions, egg capsules are most likely attached
to buried stones or shells, with the long stalk supporting
the egg case well above the sand surface, preventing its
burial, and possibly protecting the eggs from small, bot-
tom-dwelling predators. The compressed, spatulate form
of the egg case may serve to increase the surface area
available for diffusion of gases and waste products be-
5.0
4.0
3.0
2.0
1.0
PLANKTOTROPHIC
8
• 1
4o05
7o
o9
o10
o6
-^
LECITHOTROPHIC
0.0
0.1
0.2
0.3
0.4
0.5
0.6
D / VOL
Figure 23. Relationship of the number of whorls (Vol) and
the ratio of maximum diameter to number of whorls (D/V'ol)
of cancellariid protoconchs. Solid circles denote species for which
type of development is known or inferred on the basis of num-
ber of ova per capsule. Open circles denote species for which
type of development is unknown. 1, Cancellaria cooperi, 2,
Trigonostoma foveolata. 3, .\clnicte viridula: 4, Cancellaria
reticulata: 5, Cancellaria spengleriana. 6, Trigonostoma sca-
lare: 7, Olssonella smithii. 8, \arona ntitraeformis: 9, Scalptia
obliquata; 10, Cancellaria similis.
tween the egg case and the surrounding seawater. Cap-
sules were most frequently laid with the broad face of
the case oriented into the stream of flow ing seawater.
The egg cases of Cancellaria cooperi are most similar
in morpholog} to those reported by Knudsen (1950: fig.
18j for Cancellaria sp., and to his account of the egg
cases of Cancellaria spengleriana (Deshayes, 1830). All
have the characteristic long stalk, but the latter two are
described or figured as being hemi-elliptical rather than
lenticular in profile, and triangular rather than rectan-
gular in transverse section. Egg cases of Trigonostoma
joveolata (Sowerb> , 1848), a species occurring "in sand
or gravel among rocks in low-tide pools , are similar to
those of Cancellaria cooperi in shape, but are smaller in
size, and have a proportionally shorter stalk (Kilburn &
Rippey, 1982:115). Capsules of Admete viridula (Fabri-
cius, 1780), a subtidal boreal species, are attached directly
to the substrate and lack a stalk [Thorson, 1935; fig. 71
Table 2. Protoconch measurements of the three cancellariid species for which the mode of development is known or inferred
based on number of ova per capsule. Measurements are presented in the format mean/standard deviation D = diameter in mm:
Vol = number of whorls or volutions, measured according to Jablonski and Lutz (1980:332).
Species
D
\nl
D Vol
Cancellaria cooperi. LACM 40-95.1; LACM 39-
94.1; LACM 39-116.2 (\ = 4)
Trigonostoma foveolata: XM C8393; NM C8394;
\M B7651 (N = 5)
.\dmetc viridula: USNM 189720 (N = lOt
1.16/0.02
1.33/0.21
0.78/0.05
3.29/0.02
2.24/0.21
2. 12 '0.10
0.35, 0.01
0.59 0.08
0.37 0.01
Page 52
THE NAUTILUS, Vol. 102, No. 2
Table 3. Protoconch measurements for cancellariid species for which mode of development is not known Data are presented in
the format of mean/standard deviation. D = diameter in mm; \'(il = number of w horls or volutions, measured according to Jablonski
and Lutz (1980:332).
Species
1)
V„l
D \ ol
Cancellaria reticulata: L'SNM 619108 (N = 10)
C-ancellaria spengleriana. USNM 664965; USNM
.344417 (N = 6)
Trigonnstoma scalare: USNM 846304 (N = 1)
Olssonella smithiU USNM 806986; USNM 450577;
USNM 667720 (N = 5)
Sarona mitraeformis: Petit collection (N = 3)
Scalptia ohiiquala: L'SNM 629063 (N = 4)
Cancellaria similis. USNM 664967 (N = 5)
0.99/0.11
2.90/0.06
0.34/0.01
1.08/0.07
0.96
3.02/0.09
2.0
0.36/0.02
0.48
0.85/0.04
1.06/0.03
0.89/0.03
1.26 0.06
2.62/0.16
3.60/0.15
2.45/0.09
2.75 0,00
0.33/0.02
29 0.00
n )(i 0.01
(14(1 0,02
(as Velutina iindala Brown, see Thorsoii, 1944); Bouchet
& Waren, 1985: fig. 687]. The egg capsules of all can-
cellariids studied to date have roughly elliptical, parallel
sides w ith strongly to weakly keeled margins, and a me-
dial, dorsal hatching aperture. Capsules with very long
stalks appear to be restricted to the subfamily Cancel-
lariinae.
De\elopment oi Cancellaria cooperi is similar to that
ot Thais hacniaatuma jiuridana as described by D'Asaro
(1966), although C. cooperi takes about 80% longer to
reach comparable developmental stages, and does not
produce a noticeable sinusigeral ridge. It is interesting
to note the presence of an operculum, particularly prom-
inent in the planktotrophic larval stage (figure 21), in a
family noted for the absence of opercula in adults.
The present study comprises the first direct observation
of oviposition and development of any species of can-
cellariid, although the mode of development can be de-
duced in several cases from previousK' published data
on capsule contents. Thus, Cancellaria sp. (30-40 ova/
capsule, 500 /jm in diameter; Kiiud.sen, 1950:109), Tri-
gonostovia foveolata (16 larvae/ capsule; Petit & Hara-
sewych, in preparation), Admete viridula (6-7 larvae/
capsule; Thorson, 1935:67) and Admete sp. (6 larvae/
capsule, capsule referred to by MacGinitie, 1955:51,
USNM 664468) all likely undergo lecithotrophic devel-
opment, as indicated by the low number of large larvae
or ova per capsule. To date, Cancellaria cooperi is the
only cancellariid known to undergo planktotrophic de-
velopment.
In the absence of direct information, gastropod larval
development may be inferred from the morphology of
the protoconch at the apex of the adult shell. Thorson 's
"apex theory" (Thorson, 1950; Jablonski & Lutz, 1980,
1983) asserts that a large, rounded, paucispiral proto-
conch indicates non-planktotrophic larval development,
while a narrow, polygyrate, sculptured protoconch sug-
gests planktotrophic de\elo[)meiit. In more tiuantitative
studies of this relationship, Shuto (1974) found that the
ratio ol the maximum protoconch diameter (D) to the
number of protoconch whorls (Vol) was a reasonable
indicator of developmental type. \'alues greater than 1.0
were indicative ol lecithotrophic larvae, while values
below 0.3 were more characteristic of planktotrophic
larvae, especialK it the protoconch consisted of three or
more whorls. Species with D \'ol ratios between 0.3 and
1.0 usually have lecithotrophic larvae if protoconchs con-
sist of less than 2'/i whorls. \ plot of \'ol r,s. D, \'ol values
for 10 species of cancellariids (figure 23) indicates that
although protoconch morphologv appears to be an ac-
curate indicator of the mode ol larval development for
those species of Cancellariidae that fall within the di-
agnostic regions proposed b\ Shuto (1974), develop-
mental type cannot be inferred for the majority of can-
cellariid taxa using these criteria.
As more information on the life histories and diets of
additional species of cancellariids becomes available, the
relationship between the mode of larval development
and the mobility and patchiness of prey species distri-
bution may prove fruitful ground for investigation.
ACKNOWLEDGEMENTS
We thank R. R. McConnaughey for assistance in col-
lecting and maintaining animals, and R. E. Petit for
providing information on the Cancellariidae, and for
critical review of the manuscript. Susann Braden took
the Scanning Electron Micrographs.
LITERATURE CITED
Bandel, K. 1976. Morphologie der Gelege und okologische
Beobachtungen an Buccinaceen (Gastropoda) aus der siid-
lichen Karibischen See. Bonner zoologische Beitrage 27:
98-133.
Bouchet, P. and A. Waren. 1985 Revision of the northeast
Atlantic bathyal and abyssal Neogastropoda excluding
Turridae (Mollusca, Gastropoda). Bollettino Malacologico
KSuppl.): 123-296.
D'Asaro, C. N. 1966. The egg capsules, embryogenesis. and
early organogenesis of a common oyster predator, 7'/iais
haetnastorna floridana (Gastropoda: Prosobranchia). Bul-
letin of Marine Science 16(4):884-914.
D'Asaro, C. N. 1970. Egg capsules of prosobranch mollusks
from south Florida and the Bahamas and notes on spawn-
ing in the Jaborator), Bulletin of Marine Science 20:415-
440
Jablonski, D. and R A. Lutz. 1980. Molluscan larval shell
morphology: ecological and paleontological applications.
J. R. Pawlik et al, 1988
Page 53
In: Rhoads, D. C. and R. A. Lutz (eds.). Skeletal growth
of aquatic organisms: biological records of environmental
change. Plenum Press, New York, p 323-377
Jabloiiski, D. and R. A, Lutz. 1983. Larval ecolog\ ot marine
benthic invertebrates: paleobiological implications. Bio-
logical Reviews 58:21-89.
Kiiburn, R and E Rippey. 1982. Sea shells of southern Africa
McMillan South Africa, Ltd., Johannesburg. 249 p
Knudsen, J. 1950. Egg capsules and development of some
marine prosobranchs from tropical west Africa. Atlantide
Report 1:85-130.
MacGinitie, G. E. 1955. Distribution and ecology of the ma-
rine invertebrates of Point Barrow. Alaska. Smithsonian
Miscellaneous Collections 128(9):1-201.
Morch. O. \ L. 1869. Catalogue des Mollusques du Spitz-
bergen recuillis par H. Kroyer. Memoire de la Societe
malacologique de la Belgique, Tome 4, Bru.xelles.
O'Sullivan, J. B., R. R. McConnaughey. and M. E. Huber. 1987.
A blood-sucking snail: the Cooper's nutmeg, Cancellaria
cooperi Gabb, parasitizes the California electric ray. Tor-
pedo californica .\yres. Biological Bulletin 172(3):362-366.
Paige, J. A. 1986. The laboratory culture of two aplysiids,
Aplysia brasiliana Rang, 1828, and Bursatella leachii plei
(Rang, 1828) (Gastropoda: Opisthobranchia) in artificial
sea water. The Veliger 29(l):64-69.
Radwin, G E. and J. L. Chamberlin. 1973. Patterns of larval
development in stenoglossan gastropods. Transactions of
the San Diego Society of Natural History 17(9):107-117.
Shuto, T. 1974. Larval ecology of prosobranch gastropods and
its bearing on biogeography and paleontology. Lethaia 7:
239-256.
Thorson, G. 1935. Studies on the egg-capsules and develop-
ment of Arctic marine prosobranchs. Meddelelser om
Gronland 100(5):1-71,
Thorson, G, 1944. The zoology of east Greenland: marine
gastropoda prosobranchiata. Meddelelser om Gronland
121(13):1-181.
Thorson, G. 1950. Reproductive and larval ecology of marine
bottom invertebrates. Biological Reviews 25:1-45.
THE NAUTILUS 102(2):54-55, 1988
Page 54
Latirus martini (Gastropoda: Fasciolariidae),
a New Species from Honduras
Martin Avery Snyder
7-4.5 Newtown Road
\'illanova, PA 19085
ABSTRACT
Latirus martini new species is described from shallow water
off KoalaM Island. approximateK 40 miles north ot the Hon-
duras mainland. This ne« ta.xoii differs from its nearest Carib-
bean relative, Latirus angulatus (Roding, 1798), by its sculp-
ture, coloration, shape, and size. The 13-16 axial ribs on the
body whorl constitute roughly twice the number found in other
Caribbean members of this genus
INTRODUCTION
In the summer of 1985 10 specimens of a new Latirus
were taken under coral rubble b\ divers off the north
coast of Roatan Island, Honduras. These shells were ob-
tained by Mr. Thomas Honker of Florida, who kindly
passed them on to the author. The specimens were all
collected alive, the soft parts discarded, and the opercula
glued back in place on cotton. For this reason no soft
parts were examined, although the opercidum is de-
scribed. Since that time, some additional specimens have
been taken from the same locality.
Specimens of the new species are deposited in the
collections of the Academx of Natural Sciences of Phila-
delphia (ANSP), the Delaware Museum of Natural His-
tory (DMNH), and the National Museum of Natural
History, Smithsonian Institution (USNM). .additional
specimens are retained in the collection of the author.
Family Fasciolariidae Gray, 1853
Subfamily Perislerniinae Trvon, 1880
Genus Latirus Montfort. 1810
Type species: Latirus auraiitiacus Montfort, 1810, In
nionut) p> [= L. gibbulus (Gmelin, 1791 j].
Latirus martini new species
(figures 1, 2)
Description: Shell heavy, small to medium in size (,21.7
mm to 38.4 mm in length), somewhat squat, with spire
iiearK ' s length of shell; profile inflated, w ith 8-9 whorls;
protoconch (figure 2) of 1 '/2 whorl, smooth, bulbous,
translucent pale orange-brown in color; aperture oval to
squarish, greater in length than siphonal canal; anal canal
weakly developed; axial sculpture of 13-16 prominent
ribs, crossed by numerous, pronounced spiral cords with
grooves between; cords alternateK larger and smaller,
first t\\ o on body whorl about ecjual in thickness, forming
ridge and deep suture; approximateK 45 cortls on body
whorl running onto anterior end of siphonal canal, 25
cords on the penultimate whorl, with every fifth cord
somew hat stronger than others; spiral cords darkish or-
ange-brow n turning to dark brow n between axial ribs;
grooves between cords light caramel brown w ith yellow
cast; shell appears dark orange-brown, with lighter axial
ribs; columella with 3-4 distinct plicae (figure 1); teeth
translucent, shiny, white; outer lip crenulated, with in-
dentations corresponding to spiral sculpture; smooth por-
tion of inner lip extending approximately to opposing
side of the first axial rib, followed anteriorly by 14-16
irregular w hite lirae on inside of bod\ w horl; operculum
\ellow -brow II in color, chitinous, nearK filling aperture.
Type locality: 15-20 feet, under coral rubble, north
coast, Roatan Island, Honduras, summer, 1985.
Type material: Holotv pe, ANSP 361064 (27.5 mm);
paratype 1, DMNH 169442 (23.8 mm); paratype2, USNM
859070 (26.9 mm); paratypes 3-5 in the author's collec-
tion (34.0 mm. 24.8 mm, 23.6 mm). .\11 paratv pes from
the t\ pe locality.
Etymology: The species is named for the author's eldest
son.
Discussion: This species appears to be most closeK re-
lated to the highly variable Latirus angulatus (Roding,
1798). Several forms of the latter were illustrated by
Bullock (19(58, 1974). Specimens of /.. angulatus from
the Swan Islands, approximateK 120 miles north ot Hon-
duras (Bullock, 1968: fig. 5, plate 2), and a specimen
from the northern coast of South .\merica (Bullock, 1974:
fig. 21) are superficialK similar to L. martini. The col-
oration is similar but the shape and sculpture are com-
pletel) typical of L. angulatus and thus readiK distin-
guishable from L. martini. Latirus angulatus has 7-9
axial ribs, about half the iiund)er of /.. martini. The spiral
cords on L. angulatus luc weaker than those of L. mar-
tini. The tvpical coloration is also quite different. In a
M. A. Snyder, 1988
Page 55
Figures 1. 2. Lalinis martini new species. 1. Holotype, ANSP
361064. 2. EarK « horls of parat\ pe, DMNH 169442, both from
north coast of Roatan Island, Honduras, under coral rubble in
5-7 m.
dark brown specimen of L. angulatits, the raised cords
and that portion of the whorl near the suture are dark
brown, in areas between cords where there is fine or-
namentation, the shell is light cocoa-tan to orange-brown.
Generally, L. angulatus is a more elongated shell with
the spire usualK- more than half the length of the shell.
Possilile confusion could also occur with Leucozonia
nas.sa (Gnieliii, 1791 ) which is somewhat similar in pro-
tile and coloration. This shell lacks the strong a.xial sculp-
ture of L. martini, and has a thick, black-brown oper-
culum. A characteristic narrow \\ hite band at the base,
commonly terminating in a small spine on the outer lip,
serves to distinguish this species from L. martini.
Finally, confusion might arise with the recently-de-
scribed species Latirus vermeiji (Petuch, 1986). Latirus
vermeiji has less pronounced surface sculpture, is dis-
tinctlv orangish in color, and has a caramel-orange col-
ored inner lip, whereas the inner lip in L. martini is
yellowish brown. The roughly even spiral cords in L.
vermeiji are more pronounced with every other cord
being white in color.
LITERATURE CITED
Bullock, R. C. 1968. The fasciolariid genera Latirus, Doli-
cholatirus. and Teralatinis (Mollusca: Gastropoda) in the
Western .Atlantic. Master s thesis, Universit\ of Maine,
Orono, 109 p., 8 pis.
Bullock, R. C. 1974. A contribution of the systematics of some
West Indian Latirus (Gastropoda, Fasciolariidae). The
Nautilus 88(3):69-79, 26 figs.
Gmelin,J. 1791. Systema naturae, 13th ed. Part 6:3021-4120.
Montfort, P. D. de. 1810, Conchyliologie systematique et
classification methodique des Coquilles ... 2. Paris, 676 p.
Petuch, Edward J. 1986. New South American gastropods in
the genera Conus (Conidae) and Latirus (Fasciolariidae).
Proceedings of the Biological Society of Washington 99( 1 ):
8-14.
Roding, P. 1798, Museum Boltenianum, Hamburg, 199 p.
Tryon, G. W. 1881. Manual of conchology 3. Tryon, Phila-
delphia, 310 p., 87 pis.
THE NAUTILUS 102(2):56-64, 1988
Page 56
Shell Variation of Springsnail Populations in the
Cuatro Cienegas Basin, Mexico: Preliminary Analysis of
Limnocrene Fauna
Koberl Hershler
Department of Invertebrate Zoology
National Museum of Natural History
Smithsonian Institution
Washiniitiin, DC 20560, USA
Lee-Ann C. Ilayek
Mathematics and Statistics
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560, USA
ABSTRACT
Geographic variation in shell morphometry is analyzed for
three locally endemic springsnail (Gastropoda: Hydrobiidae)
species occurring sympatrically in nine limnocrenesof the Cua-
tro Cienegas Basin, Coahuila, Mexico. Despite some correlation
of size-related variables across species, groupings of populations
based on multivariate analyses were not very similar among
species, nor were they strongly concordant with current drain-
age configurations in the basin. Groups of populations of Mexi-
pyrgu.s churinceanus Ta\ lor having different patterns of shell
sculpture and color banding (and once considered separate
species) were not separated similarK on basis of shell size and
shape. Inter-population differentiation of these snails was ap-
proximately equivalent to that of Symphuphilus minckleyi
Taylor, whereas Mexithaunia quadripaludium Taylor was less
variable
INTROULCniON
One of the more remarkable aciuatic faunas of the New
World occurs in the small (1,200 km-), intermontane
valley of Cuatro Cienegas, Coahuila, Mexico, which har-
bors at least 26 locall) endemic forms (Cole, 1984; McCoy,
1984; Minckley, 1984; Hershler, 1985). Local aquatic
taxa show a great diversity in extent of differentiation
relative to adjacent biota, ranging from slightK differ-
entiated pojMilations to highly divergent genera, sug-
gesting both long-term persistence of aquatic habitat and
multiple invasions of the valley over a broad time scale
(Minckley, 1969). .\quatic organisms are deployed among
diverse, spring-fed acjuatic habitats that comprise h\'e to
seven local drainage systems (Minckle\ , 1969; LaBounty,
1974), providing what has been termed a "matchless
natural laborator> " (Taylor & Minckley 1966:22) for eco-
logical and cvolutionars- study.
Spriiigsnails (C»astropoda: Hydrobiidae) of the basin
are diverse [nine genera (five endemic), 13 species (nine
endemic); Hershler, 1985], and occur abundantly in a
large number of easiK' accessible sites, providing an ex-
cellent (>pportunit\ to study geographic variation of pop-
ulations, the analysis of w hich is considered crucial to
understanding the speciation process (Gould & Johnston,
1972; Endler, 1977). To date, researcii on this snail fauna
has largeK been taxonomic (Ta\ lor, 1966; Hershler, 1985),
although geographic variation of one endemic species
was partly analyzed (Hershler, 1985; Hershler & Minck-
ley, 1986). Taylor (1966) and Taylor and Minckley (1966)
noted apparent diversity in extent of differentiation
among local species: Mexipyrgus Taylor, an endemic
restricted to large (> 25 m-) springpools (limnocrenes)
and stream outflows, is variable enough to have been
origiiialK considered as six nominal species (Ta\lor, 1966;
synonymized to monotypy by Hershler, 1985), whereas
other snails appear morphologically uniform, at least in
the portion oi the basin that has been well studied (i.e..
all but the southeastern lobe; Hershler, 1985). It was
suggested that heightened differentiation of Mexipyrgus
resulted from marked discontinuity of its habitat:
. . it seems that habitat of this genus is more likely to
be discontinuous than that of other snails in the area.
Mexipyrgus lives in soft flocculent ooze or mud in the
lagunas, thus not in the shallows where wave action re-
moves the fine particles Extensive marsh\ areas with small
streams connecting larger water bodies provide no suitable
widespread habitat. (Taylor, 1966:188)
Variation within Mexipyrgus has largeK- been dis-
cussed in terms of shell sculpture and color banding
(Taylor, 1966), characters absent from or poorl) devel-
oped in other local snails, and there has been no attempt
to contrast intraspecific variation among members of the
snail launa using a set of common characters, such as
shell morphometric variables. In this paper we provide
such a comparison between Mexipyrgus churinceanus
Taylor and the two other species (both local endemics)
common in basin limnocrenes. Mexithaunia quadripa-
ludium Ta\ lor and S'yniphophilus ininckU'yi Ta\ lor (see
figure 2). Specifically, we seek to answer the following
questions: 1) Do these distantly related snails (Hershler,
1985) show commonalitN of pattern of geographic vari-
ation? 2) Does shell morphometric variation among pop-
R. Hershler and L.-A. C. Havek, 1988
Page 57
Figure 1. Map of the central portion of the Cuatro Cienegas
Basin, show ing major drainage features and collecting localities
(numbered as in table 1 ), The inset (lower right) shows locations
of the state of Coahuila (dashed line) and Cuatro Cienegas (dot)
within Mexico.
ulations of Mcxipyrgus churinceanus exceed that of the
tv\o other limnocrene species (above) that occur s\ m-
patrically with this species? 3) Are Mexipyrgus popu-
lations that are well differentiated in terms of shell sculp-
ture and color banding patterns similarl\ separated by
shell size and shape variation?
MATERIALS AND METHODS
Nine limnocrenes were sampled (shown in figure 1 and
described in the Appendix), representing the majority of
springpools in the study area (all of basin excluding south-
eastern lobe) where all three forms are common. Only
isolated sources (Sites 1-4, 9) or, in cases where springs
were connected by stream, upflow pools (Sites 5-8) were
considered, in order to reduce possible effects of gene
flow from contiguous populations (Hershler & Minckley,
1986). These springs belong to four separate drainage
systems of the basin and harbor forms of Mexipyrgus
referable to four nominal species (see belowj.
Mexipyrgus churinceanus was collected by sieving soft
substrates, while the other two species, associated with
hard substrates, were gathered by w ashing travertine and
macrophytes in a bucket. Material was fixed in dilute
formalin, and preserved in 70!^ ethanol.
Figure 2. Scanning electron micrographs (printed at same
enlargement) of cleaned shells ol springsnails from Laguna Tio
Candido, Cuatro Cienegas, Mexico: a, Mexipyrgus churin-
ceanus (shell height, 6.34 mm); b, Nymphophilus minckleyi
(6.45 mm); c, Mexithauma quadripaludium (6.22 mm).
For each species, about 15 live-collected, fully mature
adults, recognizable by their complete and thickened
inner shell lips, were randomly selected from collections
from each site and dried for morphometric analysis. Af-
ter whorls were counted (WH), shells were imbedded in
clay in standard apertural aspect (figure 2) and shell
outlines were draw n using a camera lucida mounted on
a WILD M-5 dissecting microscope (12 x or 25 x ). Points
on these drawings were digitized and values for the fol-
lowing "standard" shell parameters (1-4) and shell shape
descriptors (5-8) (from Raup, 1966, and elsewhere) were
generated:
1) Shell height (SH)
2) Shell width (S\V)
3) Length of bod\ whorl (LEW)
4) Width of body whorl (WBW)
5) Translation rate (T)
6) Whorl expansion rate (W)
7) Distance of generating curve from coiling axis (D)
8) Aperture shape (SA)
A calculated variable (S), consisting of the addition of
SH and SW, was generated to serve as a more realistic
measure of size than either shell length or width. Cal-
culation of shape parameters largely followed methods
of Kohn and Riggs (1975), with the exception being W,
which was measured as the mean of a series of squared
ratios of perpendicular distances from coiling axis to
sutures (shell in apertural and not apical aspect) at half
whorl intervals. The apertural suture was not used, due
to frequent loosening of coiling during last half whorl of
growth, nor were sutures used from eroded adapical sec-
tions of the spire. Digitizing was done using the CONCH
software program (Chapman et ai, 1988; methodology
fulK described therein) and a GTCO Micro-Digi Pad
12x12 linked to a KAYPRO 2000 microcomputer.
Descriptive statistics for all morphological variables
were obtained for each species and locality. The hy-
potheses of homogeneity of mean differences and vari-
ances across localities were tested for each species. An
ANO\"A model was selected for each variable of each
Page 58
THE NAUTILUS, Vol. 102, No. 2
Table I. Descriptive statistics for each species at each locahty. Data given are mean, standard deviation, and sample size (in
parentheses). L = locality; variable abbreviations are civen in te\t
\ ariable
L
SH
sw
S
LBW
WBW
WH
W
D
T
S.\
Mexipyrgus churinceanun
1
5.14
2.62
7.76
3.37
2.39
6.18
1.90
076
6.17
1.32
(15)
0.39
0.20
0.57
0.21
0.21
0.37
0.16
O07
0.49
O06
2
6.13
3.21
9.34
3.85
3.00
7.03
1.81
0.67
5.87
1.36
(15)
0.30
0.11
0.35
0.19
012
057
0.19
O04
0.53
O06
3
4.76
2.48
7.24
294
2.32
6.57
1.75
068
6.13
1.35
(15)
0.32
0.21
49
017
019
048
0.14
0.05
0.79
0.06
4
5.02
2.72
7.74
3.44
2.47
6.23
1.97
073
5.99
1.35
(14)
0.23
0.12
029
0.16
012
027
0.16
0.06
027
O07
5
3.99
2.17
6.16
2.60
2.02
6.28
1.98
068
6.19
l.'jl
(15)
0.22
0.17
0.38
0.12
016
0.38
036
0.07
064
0.07
6
7.30
3.98
11.3
4.68
3.55
6.55
1.86
0.61
5.77
1.36
(14)
0.56
0.33
084
024
0.36
041
021
O04
067
0.04
7
6.48
3.17
9.66
4.13
2.88
7.28
2.05
O70
6,67
1.36
(15)
0.30
0.18
041
0.24
0.24
044
0.40
0.07
0,65
006
8
5.49
2.54
8.03
3.. 57
2.24
7.03
1.92
0.73
7,13
1.41
(15)
0.18
0.14
025
014
0.09
061
0.28
0.06
087
06
9
7.16
3.62
lO.S
4.39
3.45
6.58
1.76
O70
6,44
1.41
(15)
0.46
0.22
0.66
019
026
0.32
0.10
O07
0.73
O09
Mcxithaiima quadri
\i(iludiuiu
1
7.39
6.44
13.8
6.61
4.72
4.57
2,20
0.62
3,81
1.11
(15)
0.55
0.41
0.90
047
O30
0.24
022
0.05
044
0.06
2
5.82
5.07
109
5.09
3.64
4.47
2.26
O60
3.14
1.13
(15)
0.41
0.28
0.66
042
0.24
0.23
0.32
0.04
0.40
0,06
3
5.73
4.83
106
5.04
3.54
4.63
2,28
065
3.50
1.13
(15)
0.30
0.33
0.59
031
019
0.25
0.33
0.05
O60
0.06
4
6.20
5.45
11.7
5.45
3.99
4.37
2.05
062
.3..52
1.07
(13)
0.57
0.36
087
0.58
025
0.26
021
0.06
0.60
0.08
5
5.77
5.09
109
5.14
3.68
4.19
2.36
0.61
3..38
1,12
(9)
0.61
0.55
1.12
052
041
0.11
041
0.08
048
0,05
6
6.72
6.00
12,7
5.76
4.51
4.44
2,07
061
3,34
1,03
(9)
0.60
0.49
1,07
047
047
041
0.11
05
048
0,0.5
7
5.23
4.50
9.73
4.56
3.27
4.32
2.14
0.63
2,85
1,12
(15)
0.36
022
0.55
0.34
0.20
0.26
0.19
0.04
0.21
()()5
8
6.07
5.15
11.2
5.18
3.78
4.67
2.20
062
.3.17
! 10
(15)
0.48
0.30
0,72
042
0.25
029
0.25
0.07
0.37
0,06
9
6.40
5.51
11,9
5.47
4.15
4.63
2.16
0.64
3.51
1.08
(15)
0.44
029
067
0,35
0.27
0.30
0.19
004
056
O05
S'yinphophiltis minckleyi
1
6.66
5.30
12.0
.5.19
4.00
4.80
2.06
057
3.24
1.12
(15)
0.38
019
0.53
0.28
0.20
034
037
O03
0.29
0.05
2
5.99
4.56
10.6
4.51
3.76
5.03
1.88
059
3.82
1.09
(15)
0.47
031
073
034
0.22
O09
019
O05
0.43
0,03
3
5.12
4.09
9.21
3.96
3.15
4.54
2.01
0.57
3.35
1 10
(12)
0.43
0.26
067
0.31
0.20
0.37
026
0.03
0.36
004
4
5.88
4.89
10.8
4.50
3.79
5.39
1.89
054
3.32
1.05
(7)
0.39
015
042
0.25
0.09
0.35
012
0.02
033
004
5
5.56
4.34
9.90
4.38
3.48
5.28
2.08
0.60
3.60
1.14
(15)
0.34
0.23
0.50
0.28
0.19
0.38
0.29
0,03
048
0.03
6
7.17
5.98
1.3.1
5.29
4.84
5.68
1.86
054
3,19
1,13
(15)
0.30
0.26
0.45
0.24
018
0.26
0.16
0.04
042
0.05
R. Hershler and L.-A. C. Havek, 1988
Page 59
Tabic I. Oititiimcd
Variable
L
SH
SW
S
LBW
WBW
W'H
W
D
T
SA
t
.5.69
4.53
10.2
4.13
.3.74
5.22
1.76
0.58
3.45
1.07
til)
0.29
0.27
54
0.19
0.26
0.14
0.04
0,05
0.24
0.05
8
7.03
5.55
12.6
5.32
4.35
.5.23
1.88
0.57
3.34
1.18
(10)
0.25
0.36
0.55
0.28
0.27
0.18
0.29
0.04
0.35
0.07
9
7.48
5.85
13.3
5.61
4.61
5.47
1.83
0.59
3.21
1.17
(15)
0.46
30
0-6S
0.35
0..30
0.27
0.13
06
0.32
006
species unless very significant heterogeneity of variance
existed, in which case the generahzed Welch test was
used to consider mean differences. Pearson correlations
were computed across species pairs for population means
of each variable. Principal component analysis (PCA)
was applied separately to each species data matri.x to
assess and compare groupings of specimens without a
priori assumptions. Because the units of measurement
were distinct and non-comparable, the anaKses were
performed on correlation matrices. Discriminant anal\ sis
(DA) was used to determine assignment of specimens to
the locality groupings on basis of shell size and shape.
This anal\ sis w as computed in a stepwise manner in order
to identify measurements contributing to significance of
discrimination. Selection criterion was ma.ximization of
Mahalanobis D-squared between closest pairs of locali-
ties. The a posteriori procedure of classification analysis
yvas performed to determine possible error of specimen
assignment to locality . When SL and SW yvere replaced
by their sum (S), slightly better localit) separation re-
sulted in the multivariate analyses and these results are
reported.
Computations yvere performed using SYSTAT (Wil-
kinson, 1986) on an IBM-.XT. and SPSSX Ver. 2 on an
IBM 4381 VM/CMS system at the Smithsonian Institu-
tion.
prevalent among iV. minckleyi populations and partic-
ularly pronounced for those of Mexithaiima quadripa-
ludium.
For each of the three species, only three principal
components y\ere significant and y ielded meaningful in-
formation. Eigenstructuresshoyv that the first component
in each analysis is dominated by size and size-correlated
variables (table 3), and explains 39-46% of the total vari-
ation. For Mcxipyrguschitrinceaniis, tyvo shape variables
(T, SA) dominate the second component yvhich explains
almost 20% additional variability, and a sixth variable
(W) is the sole measure of importance on the third axis,
explaining 14%. For the other tyvo species, size (PCI)
explains 10% more variation than for Mexipyrgus chu-
rincearms, yvhereas shape (PC2 and 3) explains only
slightly less. Weights for shape parameters are spread
over both the second and third axes, making interpre-
tations of these more difficult.
Figure 4, consisting of plots of the first three PC's for
each species with locality means (centroids) indicated,
alloyvs comparison of relative locations of populations in
PC space among species. Spread of centroids is largely
along PCI, as expected. Each plot has one or ty\o tight
clusters of a feyv centroids, w ith cluster segregation more
RESULTS
Descriptive summary statistics for each species by lo-
cality are in table 1. Results of locality tests for both
mean differences and variance are in table 2. Hetero-
geneity- of variance yvas more pronounced for Mexipyr-
gus churinceanus (significant for six variables at P <
0.01 level) than for the other species. Inter-locality vari-
ation was marked for each species, and in ail but three
cases (LB\\', Mexipyrgus churinceanus; W, D, Mexi-
thauma quadripaludium ) the hy pothesis of mean equal-
ity of variables across localities was rejected at 0.05 level.
As an example, inter-locality variation in the size-indi-
cator variable, S (= SH -I- SW). is shoy\ n in figure 3, yvith
significant differences (P < 0.05) indicated by non-over-
lapped confidence intervals. Size range for each species
is considerable; significant differences (i.e., absence of
overlap in figure 3) are especially numerous among pop-
ulations of Mexipyrgus churinceanus, y\ ith overlap more
Table 2. Results of .\iialysis of \ariance, or Welch's test; and
Bartlett's tests for homogeneity of variance for the nine local-
ities (**: P < 0.01; *: 0,01 < F < 0,05; ns: P > 0,05).
\ ariable
SH SW S LBW WBWWH W D T SA
Mexipyrgus churinceanus
Mean ** ** ** ns
\ari-
ance ns * ** *
Mexilhaiima quadripaludium
Mean ** ** ** **
Vari-
ance ns ns ns ns
Symphophihis minckleyi
Mean ** ** ** **
X'ari-
ance ns ns ns ii'-
** ** ** ** ** *♦
ns ns ns
** **
ns ns ' ns ' ns
** ** ** ** ** **
* ** *
Page 60
THE NAUTILUS, Vol. 102, No. 2
12 -
10
E
E
CO
6 -
Mexipyrgus c h u rincean us
_i 1 I ; L.
1
3
5 7
9
Mexith a u m a qiiadripalu din m
14
-
E
12
♦
10
1
10
1
\
<
i
14
E
E
12
10 -
13 5 7 9
Ny m phop h ilu s ni in c k leyi
_j I { I I I
13 5 7 9
LOCALITY
obvious for Mexipyrgus churinceanus and xV. minckleyi
than for Mexithauma quadripaludium.
Results of discriniiiiant aiiaK ses on looalit\ are in table
4. Size (S) is the most hea\ il\ weighted \ariable on DFl
for Mexithauma quadripaludium and \. minckleyi, with
LBW weighting negatively; while LBW and S are ap-
[)ro.\imatel\ ec)ual!\ and positiveK weighted on this
tuiiction for Mexipyrgus churinceanus. .\s \s itli the prin-
cipal components anaKsis, the Brst discriminant axis ex-
plained almost lO^c more variation for Mexipyrgus chu-
rinceanus than for the other two species. Note that the
shape parameters W and D were not correlated \\ ith any
of the functions for any of the species.
Entry of the first variable (S) alone for Mexipyrgus
churinceanus \ielded significant (P < 0.01) separation
of mean values for all but a single locality pair (1, 4).
While entr\ of four additional \ariables significantK sep-
arated this final pair (P < O.O.j), the significance level
decreased to 0.082 after addition of all remaining vari-
ables. Entr\ of the first \ariable for Mexithauma quad-
ripaludium (WBW ) and \. minckleyi (S) resulted in
significant .separation of all but eight and fi\e pairs, re-
spectiveK; further addition of remaining \ariables left
four and one pairs still unseparated.
Classification error rates for indi\idual specimens in-
dicated considerable overlap of populations, and varied
across localities as follows: Mexipyrgus churinceanus,
53-100?c correct classification [87/c (overall)]; Mexithau-
nm quadripaludium. 40-87rc [62rc (overall)]; A', minck-
leyi. 60-100'7 [84^t (overall)]. Mexithauma quadripa-
ludium was the poorest classified overall. \\ ith less than
60% classification in five of nine localities. Only three
discriminant functions were significant for this species
{P < 0.05), compared to six for the other two. .additional
anaKses using onK shape parameters yielded consider-
ably poorer classification [ranging from 26-39'^c (overall)
for each species], and confirmed the low discriminating
power of these variables (see Hershler & Sada, 1987).
Differentiation among drainage systems was also ex-
amined. The study area encompasses tour local drainages
(as recognized b\ LaBount\. 1974, with localities allo-
cated to these as follows (following notes in Appendix):
Becerra System (Localit> 1 ); El Garabatal (2-4); Rio Mes-
quites S\stem (5-8); and Tio Candido System (9). Clus-
tering of populations in PC-space (figure 4) does not
closeK follow partitioning of localities into drainages, as
imlicated b\ considerable spread of centroids represent-
ing localities from El Garabatal i2-4), and from the inter-
connected springs at Los Remojos (6-8).
For further analysis, localities were re-grouped into
drainage systems. .\ di,scriminant anaKsis on each species
Figure 3. Piols of S (Sll + S\V) vs. locality for each species.
Filled circles represent medians; and bars denote simultaneous
confidence intervals around the median, con.structed so that if
parentheses do not overlap, population medians are different
at 95% confidence level.
R. Hershler and L.-A. C. Havek, 1988
Page 61
Table 3. Results of Principal Compoiit'iits .\iial\scs on cacli species. Only factor coefficients having weights > 0,25 are listed. I
= Mexipyrgus churinceanus, 11 = Mcxilhauina quadripalndium. Ill = Nijmphophihts inincklcyi, Kig. = eigenvalue; % V. = % of
variance explained.
I
11
III
PCI
PC2
PCS
PCI
PC2
PC3
PCI
PC2
PC3
s
0.30
0.30
0.28
LBW
0.30
0.30
0.27
WBW
0.29
0.30
0.27
r
0.55
44
-0.45
SA
0.44
0.43
0.59
0.36
0.31
w
-0.77
0.55
0.69
D
0.43
-0.64
60
VVH
-0.32
-0.37
-0.32
Eig.
3.22
1.51
1.10
3,19
1.22
1.00
3.49
1.35
1.11
% V,
.39 2
19.8
13.8
45.5
17.4
144
45 6
16.9
13,9
using ail \ariables \\ as able to separate the four drainage
groups only for Mexipyrgus churinceanus: only two
functions were signiBcant for the other species. Resuhs
of these analyses are in table 5. Note the low weights of
S on DFl for Mexithauma quadripalndium and N.
minckleyi compared to these for Mexipyrgus churin-
ceanus, and the relatively small amount (ca. 40%) of
variation explained by the first function for all three
species. Classification error rates for individual specimens
were similar for the three species: Mexipyrgus churin-
ceanus, 61-93% [68% (overall)]; Mexithauma Cjuadri-
paludium, 46-93% [60% (overall)]; N. niinckleyi, .57-
100% [77% (overall)], with highest classification for
Drainage 1 (87-100%) and lowest for Drainage 3 (46-
61%).
There is similarity of pattern of size variation for the
three species: populations having large-sized shells are
concentrated in the southeastern portion of the study
area (figure 3, Localities 6-9). For standard shell mea-
surements, population means were significantly corre-
lated for Mexipyrgus churinceanus and Nymphophilus
minckleyi (r > 0.61, P < 0.05 for S, SL, SW, WBW)
and Mexithauma quadripalndium with Nymphophilus
minckleyi (r > 0.60, P < 0.05 for S, SL, SW, WBW,
LBW). There were no significant correlations between
Mexipyrgus churinceanus and Mexithauma quadripa-
ludium. Population means for whorl number and shape
parameters were not correlated across species with one
exception (Mexithauma quadripalndium with Nympho-
philus minckleyi, r = 0.64, P < 0.05 for W).
Table 4. Results of Discriminant AnaKses on locality for each species. Standardized function coefficients (> 0.25) and pooled
within group correlation coefficients (> 0.25) of each function with each original variable are listed. 1 = Mexipyrgus churinceanus;
II = Mexithauma quadripalndium; III = Nymphophilus minckleyi; Eig. = eigenvalue; % V. = % of variance explained; C.C. =
canonical correlation
1
II
III
DFl
DF2
DF3
DFl
DF2
DF3
DFl
DF2
DF3
Standardized
coefficients
s
0.64
0.89
0,94
-0.66
1,51
-1.61
LBW
0.69
0.90
-0.95
-0.56
1.88
2.02
-1.09
-1.43
1.63
WBW
-0.35
-1.13
0.56
-0.94
-2.15
0.35
0.91
D
T
0.39
0.65
0.30
SA
0.34
WH
49
0.61
0.79
0.48
0.77
Eig.
13.3
2.19
0.61
3.13
0.56
0.45
8.95
2.61
0.56
% V.
79.2
13.0
3.6
70.9
12.7
10.1
69.3
20.2
4.3
C.C.
0.96
0.83
0.62
0.87
Correlation
0.60
coefficients
0.56
0.95
0.85
0.60
s
0.89
-0.38
0.88
0.31
0.84
-0.36
LBW
0.94
0.74
0.54
0.61
-0.53
0.38
WBW
0.64
-0.65
0.93
0.83
T
0.34
0.31
SA
0.46
0.34
WH
0.72
0,77
0.30
0.38
0,78
Page 62
THE NAUTILUS, Vol. 102, No. 2
1 J
Mexipyrgus churiiiceanus
Mexithauma quadripaludium
(5)@
3
®
Nymphophilus minckleyi
Despite these correlations, concordance of pattern
among plots of PC scores for the three species is not
impressive, although there is similarit) in order along
size-related PCI (figure 4). Note that the tightest clus-
tered centroids for Mexipyrgus churinceanus (Localities
1, 3, 4) are vvideK separated for the other two species.
SiniilarK . di\ t-rgeiit "outK iiig centroids lor gi\en sf>ecies
{Mcxithaiinia quaclripaludium, I, S. minckleyi, 3) are
not so differentiated in the other species.
As mentioned abo\e, the populations of Mexipyrgus
considered in this stud\ are referable to four nominal
species (fide Ta\ lor. 196(-)) on basis of shell sculpture and
color banding; Mexipyrgus churinceanus (Localities 1,
2, 4); Mexipyrgus rnojarralis (5); Mexipyrgus lugoi (6-
S); and Mexipyrgus carranzae (9) (the form present at
Localit) 3 is distiiicti\e and not clearK referable to any
nominal species). These nominal species are poorly seg-
regated on the PC axes: extent of separation of rnojarralis
(5) and carranzae (9) from other centroids, for instance,
is exceeded by that seen among three populations refer-
able to churinceanus in Los Remojos spring complex (6-
8).
DISCUSSION
For all three species, most inter-population variation in-
volved shell size and size-correlated variables. Common-
ality of geographic variation patterns was indicated by
significant correlation of population means tor some of
these variables across two of three species pairs. The
extent of this commonality was not, however, impressive
when groupings of populations based on multivariate
anal) ses were examined.
A strong correlation between geographic variation pat-
terns and current drainage configurations was not ap-
parent for any of the species, suggesting that in this
example development of intraspecific di\ersit\ of shell
inorphometrv ma\ be related to ecological as well as
historical factors (see Chernoff, 19S2, for general dis-
cussion of this subject), although we acknowledge the
possibilitv that the poor correlation w ith current drainage
configuration ma\ be obscured b\ historicalK complex
basin hydrograpln (Miiickle\, 19(i9; Hershler \ Minck-
ley, 1985). Springpools concentrated around the northern
tip of Sierra de San Marcos (where the study area is
located) are higliK unilorm in water quality (Miiickley
in Cole, 1968): single measurements taken b\ us during
the study indicated, for instance, that temperature and
conductivity ranged among the nine springpools from
25.5-34.5 °C (seven localities diftering b\- < 4 °C) and
1,825-3,500 micromhos cm (seven localities differing by
< 430 micromhos, cm), respectively. Pools do dilter con-
Figure 4. 'rliree-dimeii.sioiial plots ot PC cenlroids for the
three species t\ axis. PCI; Y, PC2; Z. PC:.'3). standardized and
viewed in perspective. Sizes of balls indicating centroids are
scaled to heighten perspective.
R. Hershler and L.-A. C. Hayek, 1988
Page 63
Table 5. Results of four-group Discriminant Analyses on drainage system for each species. Standardized function coefficients (>
0.25) and pooled within group correlation coefficients ( > 0.25) of each function with each original variable are listed. I = Mexipyrgus
churinceanus: II = Mexithauma quadripaludiutn; III = \'ymphophilus minckleiji; Eig. = eigenvalue; % V. = % of variance
explained; ('.C. = canonical correlation
I
11
III
DFl
DF2
DF3
DFl
DF2
DF3
DFl
DF2
DF3
Standardized coefficients
s
2.28
0.72
2.39
-0.49
2.11
0.84
0.87
1.76
-2.78
LBW
-2.5.3
0.71
1.43
-3.12
-0.46
1.41
-1.49
1.35
WBW
1.05
-0.98
-1.87
1.23
-0.91
-1.50
1.71
W
0.61
D
0.52
-0.46
0.66
0..57
-0.67
T
0.55
0.87
SA
0.38
0.42
0.67
WH
-0.36
-0.63
0.51
0.53
Eig.
3.21
1.51
1.10
3.24
1.26
1.03
3.49
1.35
1.11
% V.
40.2
18.8
13.8
40.4
15.8
12.9
43.6
16.9
1.3.9
c.c.
0.642
0.499
0.431
0,710
Correlation
0.426
coefficients
0.262
0744
0.717
0.217
s
0.58
0..55
0.89
0.32
0.49
0.63
LBW
0.48
0.58
0.96
0.61
0.57
\VB\\'
0.65
0.36
0.80
0..50
0.83
VV
0.61
D
-0.36
0..59
-0.50
T
0.47
0.40
0.68
-0.33
SA
31
-0..32
0,46
WH
(144
-0.47
0,31
0.48
0,68
siderablv in other potentially important parameters such
as size, substrate composition, and abundance of mol-
luscivorous cichlid fishes and the relationship between
these features and shell geographic variation merits fur-
ther study .
Groupings of populations of Mexipyrgus churinceanus
on basis of shell size and shape were not strongly con-
cordant \\ith allocation of these to nominal species de-
fined b\ shell color banding pattern differences. Fur-
thermore, both univariate and multivariate analyses
showed that these populations were no more differen-
tiated in terms of shell size and shape than were those
of seemingK monomorphic Xymphophilus winckleyi.
These results suggest that evolution within Mexipyrgus
has been mosaic, with development of striking diversity
in shell color banding patterns and a few other features
(including aspects of shell sculpture, and penial lobation
pattern) coupled w ith unremarkable divergence in shell
morphology.
It is intriguing that these snails and A', minckleyi.
which differ greatly in microhabitat and presumed po-
tential for gene flow between populations at spring sources,
have similar levels of intraspecific shell morphometric
divergence, whereas Mexithauma quadripaludiutn.
which broadly- overlaps in niche with the latter (Hershler,
1984), is less variable. The possibility that these patterns
reflect differing times of origin of lineages within the
basin is not currently testable due to absence of fossil
evidence.
ACKNOWLEDGEMENTS
Fieldwork was supported by Smithsonian Research Op-
portunities Fund #1233F650. .Assistance in the field was
pro\ ided by Dr. W. L. Minckley , Dr. P. Marsh, A. Riggs,
and C. Riggs. The assistance of staff of the Scanning
Electron Microscopy Laboratory at NMNH in photo-
graphing shells is gratefully acknowledged. Ms. Molly
Kelly Ryan, staff illustrator of the Department of Inver-
tebrate Zoology (NMNH), generously prepared figures
on very short notice. Paul Greenhall (NMNH-IZ) per-
formed the laborious task of gathering shell morpho-
metric data. Drs. M. G. Harasewy ch, R. Dillon, and an
anonymous reviewer provided useful criticism of the
manuscript.
LITERATURE CITED
Chapman, R. E., M. G. Harasewych, and R. Hershler 1988.
CONCH: an interactive computer program for the analysis
of shell coiling parameters. In preparation.
Chernoff. B. 1982. Character variation among populations
and the analysis of biogeography, American Zoologist 22:
425-439.
Cole. G. A. 1984. Crustacea from the bolson of Cuatro Cie-
negas, Coahuila. Mexico. Journal of the .\rizona-Nevada
.\cademy of Science 19:3-12.
Endler, J. .\. 1977. Geographic variation, speciation, and dines.
Princeton University Press, \J, ix -t- 246 p.
Gould, S. J. and R. F, Johnston. 1972. Geographic variation.
.\nnual Review of Ecology and Systematics 3:457-498.
Page 64
THE NAUTILUS, Vol. 102, No. 2
Hershler, R. 1984. The h\(lrobiid snails (Gastropoda: Risso-
acea) of the (^uatro C^ienegas Hasin: systematic relation-
ships and ecology of a unique fauna. Journal of the .Ari-
zona-Nevada .-Keademy of Science 19:61-76
Hershler, R. 1985. Systematic revision of the Hydrobiidae
(Gastropoda: Rissoacea) of the Cuatro Cienegas Basin.
Coahuila, Mexico. Malacologia 26:31-123.
Hershler, R. and VV. L. Minckley. 1986. Microgeographic
variation in the banded spring snail (Hydrobiidae: Mexi-
pyrgits) from the (;uatro Caenegas Basin. (;()ahuila. Mex-
ico. Malacologia 27:357-374.
Hershler, R and U VV. Sada. 1987. Springsnails (.Gastropoda:
Hydrobiidae) of Ash Meadows, .•\margosa Basin, Califor-
nia-Nevada. Proceedings of the Biological Society of
Washington 100:776-843.
Kohn. A. J. and A. C. Riggs. 1975. Morphometry of the Conus
shell. Systematic Zoology 24:346-359
LaBounty. J. F. 1974. Materials for the revision of cichlids
from northern Mexico and southern Texas. LS.\ (Pcrci-
formes: (^ichlidae). Ph.D. thesis (unpublished), Arizona
State University. Tempe. 120 p.
McCoy, C. J. 1984. Ecological and zoogeographical relation-
ships of amphibians and reptiles of the Cuatro Cienegas
Basin. Journal of the .'Vrizona-Nevada Academy of Science
19:49-59
Minckley, V\' L. 1969. Environments of the bolson of Cuatro
Cienega.s. Coahuila, Mexico L'niversity of Texas at El Paso.
Science Series Number 2, p. 1-65.
Minckley, W. L. 1984. Cuatro Cienegas fishes: research re-
\ iew and a local test of diversity versus habitat size. Journal
of the Arizona-Nevada Academy of Science 19:1.3-21.
Minckley, VV. L. and G. A. Cole. 1968. Preliminary limnologic
information on waters of the Cuatro Cienegas Basin, Coa-
huila. Mexico. Southwestern Naturalist 13:421-431
Haup. D. M 1966. Geometric analx sis ot shell coiling. Journal
of Paleontology 40:1178-1190.'
Taylor. D. VV. 1966. A remarkable snail fauna Irom Coahuila,
Mexico. Veliger 9:152-228.
Taylor, D. VV. and W. L. Minckley. 1966. Neu world for
biologists. Pacific Discovery 19:18-22.
Wilkinson, L. 1986. SYSTAT:' the system for statistics. SYS-
T.-VT, Inc., Evanston, IL.
APPENDIX
Springpools sampled are numbered as in figure 1. Lo-
cality names are either those of Minckley (1969) or reflect
local usage. Locality data represent air distances troni
Cuatro Cienegas. Dates of collection are given in paren-
theses. Catalog numbers (USNM) for voucher material
(dry shell plus alcohol specimens) from each locality are
given in follow ing order: Symphophihis ininrklciji Tay-
lor, Mcxipyrgiis cliiirinrcaiuis Ta\ lor, ami Mcxithauiua
quadripaluilium Taylor.
1. Poso de la Becerra [(south pool) 3-1X-86], 13.7 km
W-SW of C^uatro C^ienegas. A once enormous spring area
(over a kilometer long) signilicantK reduced in size hy
canal development in early 19605 (see Taylor, 1966:162;
Minckley 1969: figs. 15, 16). Currently consisting of two
large springpools (each ca. 50 x 125 m-) connected by
short section ot stream. Spring orifices few and large,
occurring in tleepest (to 7 ni) portion of pools. Water liK
stands extcnsi\ c Spring and outflow i-onstituting a major
drainage of basin now feeding canals. USNM 857909,
857918, 857927.
Sites 2-4 represent three springs in the area known as
El Garabatal, located north of Poso de la Becerra and
east of Becerra s outflow . El Garabatal drains to the north
and west, and may be considered either a separate, small
drainage or a subset of the Rio Mesquites system, the
largest drainage in the basin.
2. Laguiias de Juan .Santos (5-IX-86), 12.8 km W-SW
ot Ciuatro Cienegas. Largest of El Garabatal springs con-
sisting of series of relatively shallow (depths to 2.0 m),
inter-connected lagunas fed by springs emerging along
pool margins. Pool area somewhat larger than that of
Poso de la Becerra. Extensive marshes bordering near-
entirety of spring's perimeter. Water lily stands exten-
sive. Stream outflow extending several hundred meters
before terminating in shallow marshv area. USNM
857912, 857921, 857930.
3. Unnamed small spring, know n to biologists working
in area as North Spring (3-IX-86), 12.0 km W-SW of
Cuatro Cienegas. Small springpool ca. 12 x 18 m. about
a meter deep, w ith ca. 20^ coverage b\ water lily . Out-
flow entering second pool (not a spring); di.scharge from
latter e.vtending 50 m before disappearing into hole.
USNM 857910, 857919, 857928.
4. Unnamed large spring (.5-IX-86), 11.8 km W-SW
ot Cuatro Cienegas. Roughly circular springpool, ca. 70
m across; depths not exceeding 3.5 m. Water lily dense
in center of pool. Outflow feeding small marsh. USNM
857916, 857925, 857934.
5. "West" Laguna in El Mojarral (4-I\-86). 9.0 km
SW of Cuatro Cienegas. Moderate-sized spring (26 x 59
ni; Arnold, 1972:12), with depths to 4.5 m. Orifices few
in number, cavernous. Water lily stands few, relatively
thin. Water exiting spring \ia both a shallow, surficial
stream and single, tubular, subsurface \ent. Stream out-
flow entering "East Laguna, which in turn drains into
Rio Mesquites. USNM 857908, 857917, 857926.
Sites 6-8 are inter-coiuiected springpools known lo-
cally as Los Remojos. 9.0 km S-SW ot Cuatro Cienegas.
Site 6 drains into a large pool recei\ ing discharge from
a second pool fed by outflows from Sites 7 and 8. System
draining into Rio Mesquites.
6. Northernmost of Los Remojos springs (5-IX-86).
Pool ca. IS X 32 m. Depths generalK > 1.5 m; water
lily common. USNM 857913, 857922,' 857931.
7. Intermediate Los Remojos spring (5-IX-86). Pool
ca. 28 X 47 m. shallow (< 1.0 m). Water lib uncommon.
USNM S57914. 857923. 857932,
8. Southeriunost of Los Remojos springs (5-IX-86). Pool
moderately large (22 x 42 m), with depths increasing
in southern end of pool to 4.0 m. Water lib common.
USNM 857915, 857924, 857933.
9. Laguna Tio (-andido (5-IX-86), 12.5 km S-SW of
C.uatro Cienegas. Large spring (ca. 45 x 100 m) exten-
sively vegetated by water lily and other macrophytes.
Depths generally from 2.0-4.0 m. Outflow extending
eastward, comprising part of a major system positioned
south of Rio Mesquites drainage. USNM 85791 1, 857920,
857929.
THE NAUTILUS 102(2):65-72, 1988
Page 65
Niche Congruency of Freshwater Gastropods in Central
North America with Respect to Six Water Chemistry Parameters
Eva Pip
Department of Biology
University of Winnipeg
Winnipeg, Manitoba R.3B 2E9
Canada
ABSTRACT
Occurrences of freshwater gastropods were studied at 430 sites
in central North America with respect to total alkahnity, total
dissolved solids, chloride, sulphate, phosphorus, and dissolved
organic matter. Mean niche positions for the gastropods were
calculated for the six combined water chemistr\ parameters
by comparing mean values for each species using agglomerative
hierarchical cluster analysis. Total niche relations were exam-
ined by calculatmg and summing the amount of overlap in the
observed ecological tolerance ranges for the six parameters for
each species pair, and appKing cluster and principal compo-
nent anaKses. The results showed that the species occupied a
broad spectrum of niches, with progressively increasing eco-
logical range. Gastropods with low mean inorganic values and
narrow niche widths were interpreted as specialists for waters
with low concentrations of dissolved inorganic materials. Species
which clustered together in terms of central tendencies of oc-
currence often differed from each other in terms of overall
niche similarity, and vice versa. Competition may have been
more important in waters with low inorganic concentrations.
INTRODUCTION
A number of workers have demonstrated the importance
of water chemistry in the distribution of freshwater gas-
tropods {e.g.. Boycott, 1936; Hubendick, 1947; Aho, 1966;
Harman & Berg, 1971; Dussart, 1976; Pip, 1978, 1985,
1986; Okland, 1979; Okland & Okland, 1980). However,
aside from the pioneering study of Finnish lakes by Aho
et al. (1981). relatively little is known regarding niche
relationships between freshwater gastropod species that
coexist within a given geographical area.
The ecological niche of a species can be described as
an infinite h) perspace representing ail physical and biot-
ic aspects of the environment in which the species occurs.
Nevertheless, practical considerations restrict most niche
studies to a few selected parameters that are thought to
be important for the species (Levins, 1968 in Aho et al.,
1981). .According to the latter v\orker, the niche of a
species with respect to a given parameter can be ap-
proximated by observing the species distribution over a
range of values of the parameter in a number of envi-
ronments.
The objective of the present study was to examine
niche relationships for the freshwater gastropods in cen-
tral North America in six water chemistry dimensions:
total alkalinity, total dissolved solids, chloride, sulphate,
moly bdenum reactive phosphorus, and dissolved organic
matter (DOM). These factors were chosen because a pre-
vious study (Pip, 1987) showed that these variables con-
tribute towards variation in species richness of freshwater
gastropod communities within the study area. In the
present study a niche was defined as the range between
the minimum and maximum values observed for each
species in the study area, and was thus analogous to the
concept of the "realized niche" discussed bv Hutchinson
(1957).
MATERIALS AND METHODS
A total of 430 sites was examined in central North Amer-
ica (47°-54°N and 94°-106°W) during the May-Septem-
ber, 1972-85 seasons. All sites contained water year-round.
Ponds (< 10 ha) comprised 41.0% of the sites sampled,
lakes (> 10 ha) 41.5%, rivers (> 2 m deep) 10.3%, and
creeks (< 2 m deep) 7.2%. Each site was examined for
snails by wading or canoeing; snails on macrophytes were
obtained by dredging with a rake or by using SCUBA.
Search time at each site was limited to 1 hr.
Surface water samples were collected at most sites, but
at depths of >3 m where macrophytes did not reach the
surface a van Dorn sainpler was used. The samples were
placed on ice in darkness and frozen within a maximum
of 48 hr. Samples were analyzed using methods rec-
ommended b\ the American Public Health Association
(1971).
While most of the sites were sampled only once, ap-
proximately 50 of the locations were resampled at dif-
ferent times of the growing season and in different \ ears.
For such sites, extreme low and high water chemistry
values were used for statistical anaKses. Because of en-
vironmental heterogeneity , large lakes were sampled at
different locations which were treated as separate sites.
Mean and niche comparisons were made using ag-
glomerative hierarchical cluster analysis (Sneath & Sokal,
Page 66
THE NAUTILUS, Vol. 102, No. 2
1973). Ward's method, a minimum variance technique
(Wishart, 1969) was used for clustering. Cluster fusion
distances were calculated by means oi the squared Eu-
clidean distance measure:
Distance,, = S, (Xj — y,)-
Because Ward's method is an intense!} clustering pro-
cedure, ranking of the species in the dendrograms was
checked for misclassifications by comparing with the
ranking obtained using the unweighted pair-group meth-
od (Cliltortl k Stephenson, 1975), a weakly clustering
strateg) (Sneath & Sokal, 1973). The two methods showed
similar relative ranking; thus mi.sclassifications were not
apparent. Since the relative importance of the six pa-
rameters differed with species and situation (Pip, 1987),
ail parameters were weighted etiualK in the cluster anal-
yses.
In construction of the dendrogram for mean niche
position (tigure 1), mean values for each of the six pa-
rameters were used to construct a data matrix for the
comparison of taxa with respect to the mean positions of
their niches. Because of differences in units and statistical
distributions of the six chemical parameters, values were
converteil to standardized Z scores, each parameter w ith
a mean of and a standard deviation of 1. Ward s method
was applied to the Z scores.
The first step in niche comparison was the construction
ot a combined niche congruenc\ matrix for the six chem-
ical parameters. For each parameter, the amount of o\er-
lap in the observed ecological ranges of each species pair
was di\ided by the combined range of the two species
(range between lowest minimum and highest maximum
values). Parameter \alues were not standardized. The six
overlap proportions were then summed for each species
pair (maximum possible value 6.0). The combined coef-
ficients formed a niche congruency matrix that was ana-
i\ zed using cluster anaKsis as above to gi\e the dendro-
gram in figure 2.
The niche congruency matrix was also examined using
principal component analysis (Tatsuoka, 1971), based on
the error correlation matrix. Components were f)rthog-
onalK rotated so that the lirsl principal component lay
along the direction of greatest variance.
Statistical programs used for dendrogram construction
and principal component analysis were obtained from
SPSS. Inc.. (Chicago. Illinois.
RESULTS
The ranges of concentrations of the inorganic chemical
parameters encompassed at the stud\ sites have been
presented elsewhere (Pip, 1986). The inorganic param-
eters showed a number of significant (P < 0.05) positive
intercorrelations at the stud\ sites (table 1) and could
therefore be regarded as a block with main similar trends.
However DOVl was significantK correlated onK with
moK bdenum reactive phosphorus and total dissoK ed sol-
ids.
Five or more sets of w ater chemistr\ data were avail-
able for 36 species in the present stud\ (table 2); only
these species were included in cluster and principal com-
ponent analyses. The bulk of the mean, maximum, and
minimum values on w hicli anahses were based are gi\en
by Pip (1986) for all parameters except DOM, which is
given in table 2.
The dendrogram for mean niche positions (figure 1)
with respect to the six combined parameters reflected
the lf)cations in the ecological ranges w here each species
w as most often found. C^lnster 1 consisted of species w ith
the lowest mean values lor total dissolved solids and low
values for all other inorganic parameters. On the other
hand, Armiger crista, Physa jcnncssi. and Fossaria mo-
dicella differed the most frcmi all other species; these
gastropods showed the highest mean values for total dis-
solved solids, as well as high \alues for all other param-
eters (Pip, 1986), including DOM. Other species in the
spectrum could show low or high \alues of DOM which
were largely unrelated to the concentrations ot inorganic
parameters; for example Valvata tricarinata, Fossaria
decampi. and Stagnicola catascopium show ed the lowest
mean values for DOM ol all species except Campcloma
decisum (table 2).
The relations between species in terms ot niche con-
gruency for the six parameters are represented by the
dendrogram in figure 2. The groups that emerged formed
a spectrum ol the ranges of chemical environments in
which each species was observed. Clusters 1 and 2 con-
sisted of species that showed narrow tolerance ranges for
Table 1. Correlations between water chemistrv parameters at the stud\ sites Upper diaRonal = r. lower diaijonal = p. \ = 446-
468.
\lii|\ Ixlemini
Disso!\e<l
Total
Total
reactive
organic
dissolved so
ids
alkalinity
chloride
Sulphate
phosphorus
matter
Total dissolved solids
X
0.47*
0.66*
0.61*
0.26*
0.10*
Total alkaiinitv
<0.00I
X
0.39*
0,1.5*
0.41*
0.07
Chloride
< 0.001
<0.001
X
61*
0.05
0.04
.Sniplialc
<0.001
0.001
<0.001
X
0.05
0.04
\lul\ixlcnuni reactixe
pliosplmni''
<00{)1
<onni
14
14
X
0.1.3*
Dis.solved organic matter
0017
OS
(122
22
()()()4
X
* Significant correlation.
E. Pip, 1988
Page 67
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THE NAUTILUS, Vol. 102, No. 2
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Page 69
the inorganic parameters at low corieeMitratioiis (Pip,
1986), The narrow tolerance ranges ot these species con-
tributed towards larger interspecific differences, as seen
from the larger cluster fusion coefficients; niche overlap
with species that had broad ranges was small, while the
probability that significant proportions of the niches of
any two species w ith restricted ranges would not coincide
was increased b) the narrowness of the respective ranges.
As a result, relative niche differentiation was better de-
fined in waters with low inorganic concentrations.
Species witii broader ecological tolerance ranges, on
the other iiand, could occur in waters with high inorganic
concentrations, although in most cases they could also
tolerate comparatively low values. These species formed
the clusters on the upper part of the dendrogram in figure
2, and were more numerous than species which were
restricted to a narrow range of environments. The greater
similarity between members of each of these clusters
derived from the larger degree of overlap that necessarily
arose as a result of the broad ranges. Cluster 6, composed
of Stagnicola paliistris, Fossaria modicella, and Physa
jennessi, represents species that occurred in the broadest
range of chemical environments.
Niche relations were also examined using principal
component anal\ sis applied to the niche congruency error
matrix, which extracted two major components that ac-
counted for 55.4% and 22.7% of the variance, respec-
tively. The remaining components each accounted for
5% or less of the remaining variance. The niche relations
for individual species are plotted with respect to the first
two principal components in figure 3. The distributions
that emerged reflected the groupings of figure 2, al-
though clusters 1 and 2 from figure 2 showed close af-
finities (groups A and B) in figure 3. Species in groups
A and B showed narrow niche widths and low mean
parameter values. Species in groups E and particularly
F showed wide ecological ranges; these groups also in-
cluded the most common species in the stud\ area (Pip,
1978).
The first principal component of the niche congruence-
error matrix was significantly correlated w ith mean val-
ues for the respective species of all six parameters (total
alkalinity r = 0.49, P = 0.001; total dissolved solids,
phosphorus, and chloride, all r = 0.47, P = 0.002; sul-
phate r = 0.36, P = 0.016; DOM r = 0.33, P = 0.023;
all N = 36). This large number of correlations derived
from intercorrelations of these parameters at the sites
examined (table 1). The strong correlations with the first
component are interesting, in that the values in the orig-
inal congruency matrix were not themselves correlated
with any of the chemical parameters. The second prin-
cipal component showed no significant correlations with
the mean values of the parameters examined and was
apparentl) related to an unmonitored factor.
DISCUSSION
Cluster analysis showed that the gastropods within the
study area occupied a broad spectrum ot chemical niches.
Table 2. Mean, maximum, and minimum values of dissolved
organic matter (absorbance of acidified water at 27.5 nm) ob-
served for the gastropods in the study area.
Mini-
Maxi-
Species
X
mom
mum
\
Campeloma clecisum (Say,
1816)
0.142
0.060
0.200
6
Valvata sincera Say, 1824
0.313
0.106
0.737
7
V. Iricarinala (Say, 1817)
0.204
0.018
1.366
70
Cincinnatia cincinnatiensis
^Anthony, 1840)
0.210
0,054
1.366
42
Marstonia decepta (Baker,
1928)
0.209
0.0.53
0.665
8
Probythinella laciislris (Baker,
1928)
0.299
0.180
0.387
8
Amnicola limosa (Say, 1817)
0.263
0.039
1.428
102
A. walkeri Piisbry, 1898
0.212
0054
0,815
23
Lymnaea stagnalit Linne,
1758
0.325
0.018
1.676
285
Bulimnea megasoma (Sa\-,
1824)
0.356
0054
1,383
45
Stagnicola palustris MuUer,
1774
0.314
0.060
1.638
182
S. catascopium (Say. 1817)
0.169
0.053
0.395
13
S. caperata (Sa\-, 1829)
0.349
0.252
0.438
6
S. reflexa (Say, 1821)
0.311
0.130
1.089
7
Fossaria decampi (Streng,
1896)
0.167
0.060
0.366
9
F. exigua (Lea, 1841)
0.266
0,030
0.817
15
F. modicella (Say, 1825)
0.345
0.039
1.400
53
F. parva (Lea, 1841)
0.244
0.095
0.392
6
Phijsa gyrina Say, 1821
0.307
0.009
1.638
293
P. jennessi skinneri Tavlor,
1953
0.387
0.042
1,306
33
Aplexa hypnonim (Linne,
1748)
0.442
0.0.53
1,638
41
Ferrissia parallela (Haldeman,
1841)
0.426
0,115
0,815
6
F. ricularis (Say, 1817)
0.315
0,054
1,168
53
Helisoma trivolcis (Say, 1816)
0.341
0.018
1,968
207
H. pilsbryi infracarinatum
Baker, 1932
0.254
0.050
0.684
23
H. corpulentum (Say, 1824)
0.214
0.173
0.263
5
H campanula! urn (Say, 1821)
0.237
0.009
0.817
109
H. anceps (Menke, 1830)
0.280
0.009
1.512
122
Planorbula armigera (Sa\-,
1821)
0.435
0,088
1.638
73
P. campestris (Dawson, 1875)
0.385
0,140
0,650
7
Promenetus exacuous (Sav,
1821)
0.324
0,053
1.428
84
P. umbilicatelliis (Cockerel!,
1887)
0.240
0.053
0.625
i
Armiger crista (Linne, 1758)
0.483
0.364
0.815
17
Cyrauhis parvus (Say, 1817)
0.345
0.030
1.676
193
G. circumstriatus (Trvon,
1866)
0.347
0.053
1.428
27
C. deflect us (Say. 1824)
273
0054
1 428
70
In general, the species groupings in terms of niche sim-
ilarity formed a continuous series with progressively
greater ecological range. The niches of most species over-
lapped to some extent, and only a few were mutually
Page 70
THE NAUTILUS, Vol. 102, No. 2
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•27;
-1.0
Figure 3. Distribution of species plotted « ith respect to the
first two principal components of the niche congruenc\ error
matrix, .^.ves have been orthogonally rotated. Numbers repre-
sent the following species: 1 — Campeloma decisum, 2 — Va/-
vata sincera. 3 — V. tricarinata. 4 — Cincinnatia cincinnatien-
sis, o — Marstonia decepta. 6 — Prolnjthinella lacustris. 7 —
Amnicola limom. 8 — A. walkeh. 9 — Lymnaea stagnalis, 10 —
Bulimnea megasoma, 11 — Stagnicola palustris. 12 — S. cata-
scopium, 13 — S. caperaia, 1-4 — S. reflexa, 15 — Fossaria decam-
pi, 16 — F. exigtia, 17 — F. modicella. 18 — F. parva. 19 — Physa
gyrina, 20 — P. jennessi, 21 — Aplexa hypnorum. 22 — Ferrissia
parallela. 23 — F. riciilaris, 24 — Ilclisoma trivolvis, 25 — H.
pihbryi infracarinatum, 26 — H. corpulentum, 27 — H. cani-
panulatum, 28 — H. anceps. 29 — Planorbula armigera, 30 — F.
campestris. 31 — Promenetus exaciwus, 32 — F. umbilicatellus,
33 — .Armigpr crista. 34 — Cyraulus parvus, 35 — G. circtim-
striatus. 36 — G. deflectus.
e.xclusive for some parameters. Thus, with respect to the
variables e.xamined, the majority of the species studied
could theoretically occur together in the same eri\ iron-
meiit, near the lower end of the concentration scale for
inorganic parameters. For example 25 of the species could
occur through the entire range of 100-200 mg liter for
total dissolved solids, and an additional eight could occur
in jiortions thereof (Pip, 1986).
.Mean niche positions may be regarded as the types of
environments for which each species was most charac-
teristic. Stagnicola reflexa. Helisorna corpulentum. Bu-
limnea megasoma. Campeloma decisun}. //. campan-
ulatum. Amnicola walkeri. and Marstonia decepta were
classified in clusters 1 and 2 in both figures 1 and 2. These
species showed low mean values for inorganic concen-
trations and narrow tolerance ranges; they may be re-
garded as specialists for environments with low values
of these factors. Since the latter environments in the study
area are most common on the Precambrian Shield, it is
not surprising that these species are also limited to, or
most frequent in. Shield waters.
It is interesting that some species showed marked dif-
ferences in terms of their relative positions in the den-
drograms when mean niche positions and niche con-
gruencies were compared. For e.xample Lymnaea
stagnalis. Stagnicola palustris. Cyraulus deflectus. and
Helisoma anceps showed wide niches, but their central
tendencies of occurrence were much more moderate.
Indeed G. deflectus tended to occur most often at the
low end of its ecological range, perhaps because it does
not compete w ell w ith the nimierous other species which
frequent higher inorganic concentrations.
Species with similar mean niche positions, contained
in the same clusters in figure 1, would be expected to
compete with each other more frequently than with
species in neighboring clusters, because they would be
likely to occur in the same water bodies more often.
Thus, although each species occupies a unique ecological
niche (Hutchinson, 1957), the clusters of adaptively sim-
ilar species could be viewed as functional groups (e.g.,
Stanley, 1979) which, on average, respond to certain
aspects of the environment in similar ways. However,
species w ithin the same clusters in figure 1 often showed
differences in niche congruency in figure 2. For example,
the species comprising cluster 2 in figure 1 show ed similar
mean niche positions located at lower inorganic concen-
trations, but of these species, Marstonia decepta was
included in cluster 1 in figure 2, Ferrissia rivularis and
Cincinnatia cincinnatiensis in cluster 3, Helisoma pih-
bryi infracarinatum in cluster 4, and Cyraulus deflectus
in cluster 5. Such differences in the ecological ranges of
species w ith similar central tendencies of occurrence may
ha\e been ad\antageous in that the reduced o\erlap al-
le\ iated competition (e.g.. Hughes, 1980). since the pro-
portion of cases where the tw o species would not be able
to occur together was increased.
The most common species in the study area [i.e.. Physa
gyrina (6I.9'~( of sites sampled), Lymnaea stagnalis
(60.7'( ). Helisoma trivolvis (45.3%), Stagnicola palustris
(40.5%), Cyraulus parvus (39.5%)] (Pip, 19S7) showed
broad tolerance ranges for the variables examined, and
can occupy a wide xariety of habitats. These species
would be expected to compete w ith each other frequent-
ly, since large portions of their niches were congruent.
However, they occurred most commonly at different
mean positions w ithin their niches. Ot these five species,
w hicli clustered in two adjacent groups in terms of niche
congruenc) (figure 2), onl\ L. stagnalis and H. trivolvis
occurred in the same cluster in terms of mean niche
position (figure 1).
Nonetheless, because ot the large amoimt of overlap,
species with similar niches and broad ranges still often
coexisted in the same water bodies. For example, the five
most common species above were significantly associated
w ith each other in the stuck area (Pip, 1978). Such species
with large proportions ot niche oxerlap may minimize
competition where they coexist by partitioning the hab-
itat. For example G. parvus, a small species, feeds pri-
mariK on periphy ton, w hile the larger of the common
species also ingest macrophyte tissue, and ma\ favor
E. Pip, 1988
Page 71
separate macrophyte species within the same water bod\
(Pip & Stewart, 1976). Interspecific differences in diet
have also been reported by Reavell (1980). Other types
of habitat partitioning may occur in certain situations,
for example with respect to depth (Lacoursiere ct al.,
1975), temperature (Boag, 1981), and turbulence (Calow,
1973).
The proportionally greater niche differences between
species found at low inorganic concentrations were re-
flected b\ the lack of significant association between the
species in clusters 1 and 2 in figure 2 at the same sites
within the study area [e.xcept for M. decepta and H.
campanulatum (Pip, 1978)]. Greater niche differences
in unproducti\e environments ma\' have been associated
with more intense competition (Emlen, 1973) than in
nutrient-rich situations, where niches among many species
were similar, and a number of interspecific associations
were observed. Eutrophic waters show greater produc-
tivity of algae and macroph\ tes, w hich constitute much
of the food of freshwater gastropods (Reavell, 1980).
Food has been proposed as an important factor regulating
gastropod abundance and distribution [e.g., Dillon &
Benfield, 1982). Species richness of gastropod commu-
nities in the study area is positiveK correlated with tro-
phic state (Pip, 1987). This correlation derives from the
tendencies of many species to occur more often at higher
values of phosphorus (Pip, 1978, 1986), despite the ob-
servation that most of the species can also tolerate low-
values of this factor.
Competition has been put forward as an important
factor governing gastropod distribution (e.g., Lassen,
197.5). Species which are taxonomically and morpholog-
ically similar {i.e., "sibling species") (Aho et al., 1981)
often were not grouped within the same clusters with
respect to niche similarity (e.g., the species of each Stag-
nicola, Fossaria, Helisorna) (figures 2 and 3), indicating
intrageneric ecological differentiation, w hile other con-
geners (Gyraulus circumstriatus and G. deflectus) were
grouped together. Aho et al. (1981) found a similar lack
of consistency in the distributions of gastropods in Fin-
nish lakes and suggested that competition is only one of
a number of factors that contribute towards distribution.
Conversely, most of the groupings w hich emerged in
figures 2 and 3 contained a wide representation of dif-
ferent taxonomic and morphological entities. Accord-
ingly, while the niches within each group were similar
with respect to the parameters examined, species behav-
ior and ecological requirements were likeK to be differ-
ent, decreasing competition. Furthermore, differences
with respect to other parameters ma\ also have been
important, for example t\ pe of water bod\ and bottom
substrate (Pip, 1986), t\pe of vegetation (Pip, 1978), or
other, unmonitored, chemical and physical variables.
Principal component analysis yielded a representation
of the species in a hyperspace, of w hich the first two
dimensions accounted for more than three-quarters of
the variance. While the first dimension was clearly re-
lated to the types of chemical environments in which
the species most often occurred, the groupings in figure
3 also reflected the geographical distributions and types
of habitats frequented by the respective species. Species
in groups A and B were present largely in Precambrian
Shield w aters characterized by low total dissolved solids,
and were absent or rare west of the Shield boundary.
Species in group C frequented ponds and small lakes
(Pip, 1986) with intermediate water chemistry values.
Group D occurred frequently in lakes. Group E species
occurred in a very w ide variety of water chemistry types,
and differed w ith respect to preferences for water body
type, in some cases lacking discernable preferences en-
tirely. Species in group F occurred in the most extreme
ranges of water chemistry, but were found most fre-
quently in ponds (Pip, 1986).
While the niche means and widths utilized here are
those observed for the study area as a whole, ecological
ranges may vary for a given species in different geo-
graphical regions (Pip, 198.5), and are likeh different for
individual populations. The importance of particular pa-
rameters ma\ also vary among habitats with different
chemical and physical characteristics (Pip, 1987). Thus
the exact position of the niche may be a plastic attribute
that can vary to some extent with situation as well as
time.
LITERATURE CITED
Aho, J. 1966. Ecological basis of the distribution of the littoral
freshwater molluscs in the vicinity of Tampere, South Fin-
land. Annales Zoological Fennici 3:287-322.
Aho, J, E. Ranta, and J. Vuorinen. 1981. Species composition
of freshwater snail communities in lakes of southern and
western Finland. Annales Zoological Fennici 18:2.33-241.
American Public Health Association. 1971. Standard methods
for the examination of water and wastewater. American
Public Health .Association, New York, 874 p.
Boag, D. .\. 1981. Differential depth distribution among
freshwater pulmonale snails subjected to cold tempera-
tures. Canadian Journal of Zoology 59:733-737.
Boycott, C. 1936. The habitats of the freshwater molfuscs in
Britain. Journal of Animal Ecology 5:118-186.
Calow, P. 1973. Gastropod associations w ithin Malham Tarn,
Yorkshire. Freshwater Biology 3:521-534.
Clifford, H. T. and VV. Stephenson. 1975. An introduction to
numerical classification. Academic Press, New York, 229 p.
Dillon, R. T., Jr. and E. F. Benfield. 1982. Distributions of
pulmonate snails in the New River of Virginia and North
Carolina, U.S.A.: interaction between alkalinity and stream
drainage area. Freshwater Biology 12:179-186.
Dussart, G. B. J. 1976. The ecology of freshwater molluscs in
north west England in relation to water themistry. Journal
of Molluscan Studies 42:181-198.
Emlen, J. M. 1973. Ecology: an evolutionarx approach. .\d-
dison- Wesley Co., Reading, \i.\, 493 p.
Harman, W. N. and C. O. 15erg. 1971. The freshwater snails
of central New York with illustrated ke\s to the genera
and species. Search: Cornell L niversit\ Agricultural Ex-
perimental Station 1:1-68.
Hubendick, B. 1947. Die Verbreitungsverhaltnisse der lim-
nischen Gastropoden in Sudschweden. Zoologische Bid-
rucke Uppsaliense 24:419-.559.
Hughes, R. N. 1980. Strategies for survival of aquatic organ-
Page 72
THE NAUTILUS, Vol. 102, No. 2
isms. In: Barnes, R. S. K. and K. H. Mann (eds.). Funda-
mentals of aquatic ecosystems. Blackwell Scientific Pub-
lications, Oxford, p. 162-184.
Hutchinson, G. E. 19.57. Concluding remarks. C^old Spring
Harbor Sym[X)sium. Quantitative Biology 22-41.5-427
Lacoursiere, E., G N'aiiiancourt, and R. Goiilure 1975. Re-
lation entre les plantes aquatiques et les gasteropodes (Mol-
lusca. Gastropoda) dans la region de la centrale nucleaire
Gentilly 1 (Quebec). Canadian Journal of Zoology 53:1868-
1874.
Lassen, H. H. 1975. The diversity of freshwater snails in view
of the equilibrium theory of island biogeography. Oeco-
logia 19:1-8.
Okland, J. 1979. Distribution of en\ ironmental factors and
fresh-water snails (Gastropoda) in .Norway: use of Euro-
pean invertebrate surve\ principles Malacologia 18:211-
222.
Okland, J. and k. A. Okland. 1980. Acidification threatens
trout diet. Research in Norway 1980, p. 21-27.
Pip, E. 1978. A survey of the ecolog\ and composition of
submerged aquatic snail-plant communities, Canadian
Journal of Zoology 56:2263-2279.
Pip, E. 1985. The ecolog\ of freshwater gastropods on the
southwestern edge of the Precambrian Shield. Canadian
Field-Naturalist 99:76-85.
Pip. E. 1986. The ecology of freshwater gastropods in the
central Canadian region. Nautilus 100:56-66.
Pip. E. 1987. Species richness of freshwater gastropod com-
munities in central North America Journal of MoUuscan
Studies 53:16.3-170.
Pip, E. and J M Stewart. 1976 The dynamics of two aquatic
plant-snail associations. Canadian Journal of Zoolog\ 54:
1192-1205.
Reavell, P. E. 1980. A study of the diets of some British
freshwater gastropods. Journal of Conchology 30:253-271.
Sneath, P. H. A. and R. R. Sokal. 1973. Numerical taxonomy.
\V. H. Freeman and Co., San Francisco, 573 p.
Stanley. S. M. 1979. Macroevolution, pattern and process. VV.
H. Freeman and Co., San Francisco, 332 p.
Tatsuoka. M. M. 1971. Multivariate analysis: techniques for
educational and psychological research. John \\ ile\ & Sons,
New York, 310 p.'
Wishart. D. 1969. Mode analysis: a generalization of nearest
neighbour which reduces chaining effects. In: Cole, .\. J.
(ed). Numerical taxonomy. Academic Press, London, p.
282-308.
THE NAUTILUS 102(2):73-77, 1988
Page 73
Factors Affecting the Distribution of Sphaeriid Bivalves in
Britannia Bay of the Ottawa River
B. \^ . Kilgour
G. L. Mackie
Departmt'iit of Zoology
Universit) of Guelph
Guelpli, Ontario NIG 2W1
Canada
ABSTRACT
Distribution of Sphaeriidae in Britannia Bay of the Ottawa
River was not homogeneous and was primarily affected by
sediment particle size and depth. Total sphaeriid diversity is
maximal at particle sizes near 0.18 mm. Total sphaeriid density
and abundance of the genus Pisidiurn increase with decreasing
water depth. Distribution of species within the genera Pisidiurn
and Musculium is variable. These data indicate that compar-
isons of pisidiid distribution should consider at least depth, and
particle size in the sampling design.
INTRODUCTION
The importance of sphaeriids as part of the aquatic com-
munity has lead to a number of studies dealing with the
distributional patterns of this group (Healey , 1978; Mack-
ie et ai. 1980). Most studies, liowever, ha\e dealt with
distribution patterns among a group of lakes, rivers, or
sections of a river. Usually it is assumed that variation
among sites is greater than the variation within sites.
However, data in Avolizi (1976) and Holopainen (1979)
suggest that the abundance of some species can be vari-
able within a single site, but little work has been done
to determine the extent of this variation.
Britannia Bay , a section of the Ottawa River just above
Ottawa and Hull, Canada, is relatively unpolluted, and
is known to contain a diverse sphaeriid fauna ( 15 species),
including members of Musculium. Pisidiurn, and
Sphaerium (Mackie, 1971). The purpose of this study
was to determine if the depth distribution of sphaeriids
in Britannia Bay is homogeneous, and if distribution is
a function of the sediment particle size, organic matter
content, algal biomass, and depth.
MATERIALS AND METHODS
During mid-.\ugust. 1985. benthos and sediment samples
were taken at eight depths (0.25. 0.35, 0.50, 1.0, 2.0, 3.0.
4.0, and 6.0 m) along a transect in Britannia Bay of the
Ottawa Ri\er (figure 1).
At each depth, five Ekman grab (15 x 15 cm, 0.32
mm mesh screen on top) samples were taken, except at
0.36 m where four samples were taken. A sediment sam-
ple was taken at each depth from which three subsamples
(approximately 100 ml) were taken. Two for determi-
nation of percent organic matter content and one for
geometric mean particle size.
From each benthic sample, all sphaeriids and algae
(entangled masses of Lyngbya sp. and Vaucheria sp.)
were hand picked. Sphaeriids were counted and iden-
tified to species. Algae was dried at 70 °C and weighed
to the nearest 1.0 mg. Total sphaeriid diversity (Shannon
& Weaver, 1949) and density (N/m-), and individual
species abundances were calculated for each benthic
sample.
The organic content of the sediments was estimated
by ashing two of the sediment samples from each depth
and expressing the result as percent loss on ignition. Geo-
metric mean particle size was determined with the third
sediment sample from each depth using the methods
outlined by Lotspeitch and Everest (1981).
Stepwise multiple regressions (Ostle & Mansing, 1975)
were performed to determine which environmental fac-
tors significantly affected total sphaeriid diversity and
density, and the abundances of individual genera and
species. Only those variables found to be independent
by simple correlations (table 1), were used coincidentally
in the regression models.
Box plots (McNeil, 1977) determined that all data,
except diversity values, were most normally distributed
when the depth means were transformed by logjQ. All
simple correlations and stepwise multiple regressions were
calculated using the normalized data.
RESULTS
Depth means of algal biomass, and ?? organic matter
content are presented in figures 2 and 3. Both parameters
increased with increasing water depth. Trends in sedi-
ment particle size are given in figure 4. Compared to
other depths, sediments were coarser at the 1.0, 2.0. and
6.0 m depths.
Twelve sphaeriid species were found in Britannia Bay
Page 74
THE NAUTILUS, Vol. 102, No. 2
laoa-
Figure 1. Map of the Ottawa River, and an enlarged map of
Britannia Bay above Ottawa indicating location of tfie transect.
Depth contours on enlarged map are in metres.
with P. casertanum the most abundant species. Mean
abundances at each depth for each species are presented
in table 2.
Diversit) values were highest at the shallow depths
(0.2.5-0.50 in) and at .'3.0 and 4.0 m (figure 5). Stepwise
multiple regression indicated that diversity increased with
decreasing particle size (table 3). Figure 6 indicates a
unimodal distribution oi diversity in relation to geo-
metric mean particle size. Maximum diversilN occurred
near a particle size of 0.18 mm.
Regression analysis indicated that total sphaeriid den-
sity significantly increased w ith decreasing water depth
Table I. C^nrrelatinns between some environmental variables.
Sigiiilitant rorrclaliniis [P < O.O.'j) are indicated 1)\ asterisks
(•)
.Mgal
biomass
Organic
matter
Depth
Organic matter
Depth
Particle size
0.787*
0.839*
f) Vyl
0.863*
-0,043
80D-
<00-
o.so 1.0 2.a
Depth (in)
«.0
6.0
Figure 2. Mean algal biomass at each depth along the transect
with 95^< confidence limits indicated b\ vertical lines.
(table 3). Density at 0.25 m was less than that at 0.36
and 0.5 m (figure 7).
Abundance of the genus Fisidium significantK in-
creased w ith decreasing water depth (table 3). A similar
relationship was found for P. variabile. Abundance of P.
casertanum significantK increased with finer sediments
and lower organic matter content. None of the other
seven Pisidium species were correlated with the en\i-
ronmental variables.
Abundance of the genus Musculiniu did imt correlate
with any of the variables tested, .\bundance ot A/, trans-
versum increased significantK with decreasing water
depth (table 3). Abundance of A/, secitris correlated pos-
itively with algal biomass.
No relationship was found tor the genus Sphaerium
which was represented b\ onK one species, S. striati-
nuin.
at 2-
322
0.25 0.36 0.50 1.0 2.0 3.0 t. 6.0
Depth (m)
Figure 3. Mean organic matter content of the sediments at
each depth along the transect with 95^t confidence limits in-
dicated l>\ \frtital lines.
B. W. Kilgour and G. L. Mackie, 1988
Page 75
Figure 4. Geometric mean particle size at each depth along
the transect.
2.5-
1.5-
0.50 l.D
Depth (m)
6.0
Figure 5. Mean diversity values at each depth along the tran-
sect with 95^0 confidence limits indicated bv vertical lines.
DISCUSSION
In Britannia Bay, total sphaeriid diversity significantK- in-
creases with finer sediments. The finer sediments at 3.0
and 4.0 m, which correspond with high diversity values,
contradict the positive correlation which exists between
particle size and depth in rivers (Hamill, 1975). The sand
bar between the 3.0 and 4.0 m sites (figure 1) may have
affected the distribution of sediments in the same manner
that P'olke and Ward (1957) describe for a sand bar in
the Brazos River.
The unimodal distribution of diversity in relation to
particle size (figure 6) indicates that diversity is maximal
at a particle size near 0.18 mm, and decreases with finer
or coarser material. Finer substrates enhance burrowing
(Rogers, 1976), production (Hamill, 1975), growth and
reproduction (Mackie & Qadri, 1978), and may improve
survival in young clams (Gale, 1976). Fine substrates,
however, may reduce oxygen a\ailability (Meier-Brook,
1969; Hartnoll, 1983). The particle size at which diversity
is maximized may represent the optimal grain size for
efficient burrowing and /or oxygen/ nutrient availability'.
Total sphaeriid density increases in shallow water in
Britannia Bay. From the literature, it appears that re-
source availability and utilization may be affected by
depth. Bacteria and phytoplankton (especially diatoms),
which are food resources for sphaeriids (Holopainen,
1985), are generally more abundant in shallow water (0-
2.0 m) than in deep water (> 5.0 m) (Hargrave, 1970).
The drop in total sphaeriid density at 0.25 m which
Table 2. Mean abundance (No. /m-) of individual species at each depth along the transect.
Depth (m)
Total
no.
Species
0.25
0,36
0.5
1.0
2.0
3,0
4.0
6.0
sampled
Musculium
M. securis (Prime, 18.32)
17.8
8.9
88.9
80.0
115.5
35.6
39
M. transversum (Say, 1829)
444.4
266.6
26.6
8.9
17.8
80
Pisiditim
P. casertatuim (Poli, 1795)
942.1
1,833.2
1,173.2
35.6
17.8
142.2
106.7
437
P. duhitim (Sav, 1816)
17.8
11.1
26.7
8.9
8.9
8
P. fcrrugineum (Prime, 1852)
22.2
44.4
8.9
8
P. hcnsloivanum (Sheppard, 1825)
188.9
168.9
8.9
37
P. lilljeborgi (Esmark and Hoyer, 1886)
124.4
600.0
1,262.1
8.9
8.9
35,6
88.9
35.6
230
P. nitidiim (Jenvns, 1832)
231.1
944.4
1,528.7
17,8
26.7
26.7
291
P. punctatum (Sterki, 1895)
133.3
422.2
160.0
8.9
17.8
8.9
75
P. lahabile (Prime, 1852)
106.7
100.0
195,5
8.9
44
P. ualkcri (Sterki, 1895)
11.1
1
Sphacriuni
S. striatiniim (i.amarck, 1818)
195.5
26.7
35,6
231.1
62.2
17.8
64
Total no. sampled
227
396
539
10
20
59
49
14
1,314
Page 76
THE NAUTILUS, Vol. 102, No. 2
2.5-
.'-*-,
2-
A
\
1.5-
'' \
s,.
1-
1 1 ^'**«,«„,^^
.5-
5
1
.2
1
.25
.
sooo-
Geometric Mean Particle Size (mm)
Figure 6. Relationship between mean diversity and geometric-
mean particle size with 95/( confidence limits indicated b\
vertical lines.
4500-
<ooo-
'^ 3500-
E
o 3000-
>^ 2500-
i 200O-
Of
CD
c 1500-
o
Of
^ 1000-
500-
^
0. 50 1.0 2.
Depth (m)
3.0
4.0
6.0
Figure 7. Mean densits at each depth along the transect with
95% confidence limits indicated h\ vertical lines.
is inconsistent with the regression model, ma\ result from
wave action and or summer lethal temperatures, which
can significantly reduce the numbers of sphaeriids in the
shallower areas in Britannia Ba\ (Mackie, 1971; Mackie
& Qadri, 1978).
Regression analysis suggests that the genus Pisidium
prefers shallower water in Britannia Bay. The relation-
ship between P. variabile and depth is consistent with
the distribution of the genus. In contrast, P. casertanum
prefers finer substrates with lower organic matter con-
tent. The distribution of P. casertanum suggests that
there can be variabilitv in the distributional patterns of
species within a genus.
That the distribution of the genus Musciiltum is not
related to an\ oi the variables tested may be due to
variation in the distribution patterns of the two species
of Musculiiim. Competition between M. securis and M.
transversttni in Britannia Bay, as described b\ Mackie
et al. (1978), ma\ also be responsible for these results. It
is likely that two species are not sufficient to determine
a general distribution pattern of a genus.
Table 3. Significant relations between the dependent \ ariables
(D) and the independent variables (I), where, R = correlation
coefficient between 1) and I, a = algal biomass, b = depth, c =
geometric mean particle size, and d = % organic matter content
of the sediments
Significance
D
I
fi (D.I)
level
Total sphaeriid diversity
c
-0.791
P < 0.020
Total sphaeriid density
b
-0.720
P < 0.050
Musculiiuu scrtnis
a
as 19
P < 0.025
Mttsculinni trunsrcr^iun
1,
-0 940
P < 0.010
I'isidiuni
b
-0.733
P < 0.030
Pisidium cascfidiniin
c
-0.711
d
-0614
total R- =
0.883
P < 010
Pisidium vnhahilc
1.
(19.39
P < OOIO
The positive relationship between abundance of M.
securis and algal biomass has also been described by
Mackie and (^adri (1978) who suggested that M. securis
utilizes the algae Lyngbya and Vaucheria as food re-
sources in Britannia Ba\. The \ariabilit\ in the distri-
bution of the two species of Musculium ma\ also be
related to the food forms with which the clams are as-
sociated.
That the abundance of the genus Sphaeriuin and most
Pisidium species do not correlate with the em ironmental
variables ma\ be a result of modal relationships with
these variables, or relationships with variables that were
not tested. The relationships found centre on the im-
portance of particle size and depth, w hich have been
shown b\ Green ( 1971 ) to be important factors affecting
distribution of sphaeriids among lakes. It is suggested
that in studies dealing with the distribution of sphaeriids,
particle size and depth are important factors to be con-
sidered in the sampling design.
ACKNOWLEDGEMENTS
We wish to thank Mr. Clarence Kilgour for help with
the sampling. Comments from Mr. Chris McCiall and
two anonymous reviewers improved the manuscript. The
study was funded b\' the Natural Sciences and Engi-
neering Research Council of Canada, Grant No. A-9882
awarded to C; LM.
LITERATURE CITED
.\volizi, R J. 1976. Biomass turnover in populations of vivip-
arous sphaeriid clams: comparisons of grow th. fecundity,
mortalitv and liiomass production Hvdrobiologia 51: 163-
168.
Folke, R. L and W C W ard 1957 Brazos River Bar: a study
in the significance of grain size parameters. Journal of
Sedimentary Petrology 27:3-26
Gale, W. F. 1976. \'ertical distribution and burrowing be-
B. W. Kilgour and G. L. Mackie, 1988
Page 77
haviour of the fingernail clam, Sphaeriuni transvcrsum.
Malacologia 9:121-125.
Green, R. H. 1971. .\ multivariate statistical approach to the
Hutchinsonian niche: bivalve molluscs of central Canada.
Ecologv 52:543-556.
Hamili. S. E. 1975. Production of sphaeriid clams and am-
phipod crustaceans in the Ottawa River near Ottawa-Hull,
Canada. M.Sc. thesis. University of Ottawa, 98 p.
Hargrave, B. T. 1970. Distribution, growth, and seasonal
abundance of Hyallela azteca (Amphipoda) in relation to
sediment microflora. Journal of the Fisheries Research
Board of Canada 27:685-699.
Hartnoll, R. G. 1983. Substratum. In: Earll, R. and D. G.
Erwin (eds. ). Sublittoral ecology: the ecologv of the sub-
littoral benthos. Clarendon Press, 0.\ford, p. 97-124.
Healey, M. C. 1978. Sphaeriid mollusc populations of eight
lakes near Yellowknife, Northwest Territories. Canadian
Naturalist 92:242-251.
Holopainen, I. J. 1979. Population dynamics and production
ol Pisiditim species (Bivalvia, Sphaeriidae) in the oligo-
trophic and mesohumic lake Paajarvi, Southern Finland.
Archiv fiir Hvdrobiologie 54iSuppl ):466-508.
Holopainen, I. 1985 Feeding biology of Pisidiidae (Bivalvia)
with special emphasis on functional morphologv of the
digestive tract Lammi Notes 12:5-9.
Lotspeitch, R. B and F. H Everest. 1981 A new method for
reporting and interpreting te.xtural composition of spawn-
ing gravel United States Forest Service Research Note
PN\\-369.
Mackie, G. L. 1971. Some aspects of the distributional ecology
of macrobenthos in an industrialized portion of the Ottawa
River near Ottawa and Hull, Canada M.Sc. thesis, Uni-
versity of Ottawa, 161 p
Mackie, G. L. and S. U. Qadri. 1978. Effects of substratum
on growth and reproduction of Musculium securis (Bi-
valvia: Sphaeriidae). The Nautilus 92:135-144.
Mackie, G.L.S. U.Qadri, and R. M. Reed. 1978. Significance
of litter size in Musculiuryi securis (Bivalvia: Sphaeriidae).
Ecology 59:1069-1074.
Mackie, G. L., D. S. White, T. W. Zdeba, and N. A. Thomas
1980. A guide to freshwater mollusks of the Laurentian
Great Lakes with special emphasis on the genus Pisidium.
U.S. Environmental Protection Agency, Duluth, MN, 144 p.
McNeil, D. R. 1977. Interactive data analysis: a practical
primer. John Wiley and Sons, Toronto, 186 p.
Meier-Brook, C. 1969. Substrate relations in some Pisidium
species (Eulamellibranchiata: Sphaeriidae). Malacologia 9:
121-125.
Ostle, B. and R. W. Mansing. 1975. Statistics in research: basic
concepts and techniques for research workers, 3rd ed. Iowa
State University Press. Ames, 596 p.
Rogers, G. E. 1976. Vertical burrowing and survival of sphae-
riid clams under added substrates in Pool 19, Mississippi
River. Iowa State Journal of Research 51:1-12.
Shannon, C. E. and W. Weaver. 1949, The mathematical
theory of communication. University of Illinois Press, Ur-
bana, 125 p.
THE NAUTILUS 102(2):78-81, 1988
Page 78
Hijpselostonia holimanae New Species, a Pupillid
Land Snail from Thailand
Fred G. Thompson
Florida State Museum
University of Florida
Gainesvilie, FL .32611, USA
Harry G. Lee
709 Lomax Street
Jacksonville, FL 32204,
USA
ABSTRACT
Hypselostorna holimanae new species (Gastropoda, Pulmonata,
Pupillacea, X'ertiginidae) is described from a limestone range
near Kancliaiiaburi. Thailand Its morphological characteristics
are distinct to the extent that no close relationship between this
form and other nicmbers of the genus is apparent. Hypselos-
torna tubifcrum (Benson, 1856) from Burma is it closest phy-
logenctic and geographic congener. The two species are alike
in details ol sculpture and umbilical width, but differ widely
in shell shape and aperture barriers. Similarities of reduced
apertural dentition among other species of Hypselostorna are
cnnsidered to be due to convergence.
INTRODUCTION
The land snail fauna of Thailand is \ery poorly known.
The genus Hypselostorna is wideK distributed from Bur-
ma through CJamhodia, Vietnam, Mala\a, the Loo C^hoo
Islands and the Philippine Archipelago. Yet not a single
species has hitherto been recorded from Thailand. The
species de.scribed in this paper was collected by Stephen
C. Holimaii while serving as a Peace Corps volunteer in
Thailand. During his few opportunities of leisure, Mr.
Holiman collected mollusk specimens for his mother,
Mrs. Stanley (Bonnie) Holiman of Jacksonville, Florida,
who has a.ssemhled a private collection of fair importance
becau.se of the data that accompanies the specimens. The
mollusks from Thailand were submitted to us by Mrs.
Holiman for identification. The new species of Hypse-
lostorna described in this paper shows strong similarity
to the generic t\ pc species from Burma, and differs con-
spicuously troi7i other known species. We are honored
to name this snail after Mrs. Holiman in recognition of
her bringing this species to our attention.
Hypselostorna holimanae new species
(figures 1-6)
Description: Shell small, about 2.6-2.9 mm wide and
2.3-2.6 mm high; about 0.8.5-0.9.5 times as high as w ide.
Shell turlMii-shaped with a moderateK long conical spire
(figures l-.'3, holotype). Last whorl conspicuously en-
larged, and with a distinct peripheral angle and a basal
angle. Last whorl flat below peripheral angle, and strong-
1\ shouldered above. Occasional specimens may be weak-
1\ furrowed below periphery. Last whorl ascending at
about 10° to longitudinal axis of spire (figures 2, 4). Neck
of last w horl becoming narrowed behind aperture, and
extending forward for about ',5 of minor diameter of last
whorl (figure 3); indented externally over junction of
angulo-parietal lamella and slightK so over colinnellar
lamella. Base of shell broadK umbilicate due to lateral
expansion of last whorl. Umbilicus about 0.40-0.53 times
minor diameter of last w horl as measured across the basal
angle. Whorls 4.6-4.9. Protoconch consisting of about 1.5
whorls that appear smooth under light microscopy; at
higher magnifications the whorls are sculptured with a
dense mesh of fine reticulating threads that have an
underlying spiral arrangement (figure 5). Whorls of te-
leoconch sculptured with raised spiral threads that are
nearK uniformly distributed o\er the surface of the shell
(figure 4). Threads weak liut distinct on spire; most con-
spicuously developed on last whorl (figure 6). Spiral
threads interrupted at irregular intervals b\ incremental
growth striations, which in some specimens ma\' cause
the spiral sculpture to appear cancellate or beaded on
the spire. Color dark brow n with a light brow n aperture
and white lamellae within the aperture. Face of aperture
translucent w ith fine radiating lirow n lines due to raised
spiral threads on opposite surface Peristome broadly ex-
panded and nearK uniformly wide around aperture. Ap-
erture barrier with four teeth located on inner rim of
the aperture just behind expanding peristome and ar-
ranged o|5posite each other in a cross-configuration. .An-
gulo-parietal lamella rectilinear, torming a single short
undulating blade; bifid as is typical for genus, with an-
gular segment smaller and separated from parietal por-
tion by a weak notch. Palatal and basal plicae short and
lateralK llattcned; coiiliiied to inner ritu of peristome.
Columellar lamella tubercular and located on a slight
callus.
Measurements in mm for the holot> pe and three para-
t\ pes (I'F 1 13428) selected to show \ariation follow , The
minor diameter is the transverse w idth of the bod\ w horl
posterior to the neck of the aperture. Other measure-
ments were made of standard parameters.
F. G. Thompson and H. G. Lee, 1988
Page 79
1
Figures 1-6. Hypselostoma holimanae new species. 1-3. Holotvpe (UF 113427). x 26. 4-6. Parat\pe (UF 113483). 4. x 40.
5. X 120, 6. X 160.
Page 80
THE NAITILUS, Vol. 102, No. 2
Specimen Height Maj. w. Min \v .\per w. L'mbil Whorls
Holotype
■2A
2.6
2.4
1.4
1.2
4.7
Paratvpe
2.3
2.7
2.3
1.4
0.9
4.8
Paralype
2..3
2.6
2.3
1.2
1.0
4.6
Paralype
2.6
29
2.5
1.6
1.0
49
Type locality: Thuiluiid. Kaiicliaiiaburi Pro\ ince, small
limestone range on the west border of the Kanchanaburi
Agricultural College, about 15 km w est of Kanchanaburi.
Holot\pe: IF 113427; collected March 15, 1987 b\ Ste-
phen Holiman. Paratypes: UF 113428 (12), UF 113483,
Mahidol Universit\ Malacology Collection, Bangkok (12),
and the private collections of Harry G. Lee (12) and
Bonnie Holiman (12).
The specimens constituting the t\ pe series are recently
dead shells. They were collected at the height of the dry
season from leaf-litter in a forested knoll at the top of
the limestone range.
COMPARISONS
Hijpselostoma Benson, 1 856 belongs to a group of genera
that also includes Boysidia .\n\e\. 1881, Paraboijsidia
Pilsbry, 1917, Gyliotrachela Tomlin, 1930, Aulacospira
Moellendorff, 1890, Anauchen Pilsbry, 1917, and Sys-
tenostoma Bava\ and Dautzenberg, 1909. These genera
were monographed b\ Pilsbry (1917). More recently,
Jutting (1950) reviewed the known species of the first
four genera. Members of this group have protoconch
sculpture similar to that described above for H. holi-
manae. The variation of this sculpture w ithin the group
and its ph\ logenetic significance v\ill be discussed else-
where (Thompson, in preparation). The classification of
species within these genera is sometimes problematic,
and no one who has worked with the group seems to
have been comfortable with the systematic arrangements
of previous authors (Pilsbry, 1917; Jutting, 1950, 1961;
Solem, 1981; Thompson & Dance, 1983). Much of the
problem centers on the emphasis that is placed on the
development of the parietal and angular lamella, the
reduction ot the aperture barrier, the sculpture on the
teleoconch and the degree to which the aperture is at-
tached to or free of the previous whorl. Hypselostoma
hohmanae is placed w ithin Hypselostoma because of the
fused angulo-parietal lamella, the free aperture and the
spiral sculpture ot the teleoconch. The angulo-parietal
lamella is weaker than in most other species of the genus,
and the aperture barrier in general approaches the con-
dition that characterizes Anauchen. However, all of the
species of Anauchen lack any indication of an angular
lamella and none have raised spiral sculpture on the
teleoconch.
Few species of Hypselostoma have thus far been de-
scribed from the mainland on southeast .\sia. Pilsbrv
(1917) discussed the species known at that time. More
recently. Jutting (1950) briefly reviewed the genus and
described several additional species (1950, 1961, 1962).
Hypselusluma. as used b\ both Pilsbrv and Jutting, is a
poKpIn letic assemblage that contains species belonging
to at least three genera. F"our species (H. terae Tomlin,
1939, H. megaphona Jutting, 1949, H. elaphis Jutting,
1949, // pcrigyra Jutting. 1949) were transferred to Boy-
sidia (Dasypupa} b> Thompson and Dance (1983:109).
Three others related to H. dayanum Stoliczka, 1871 be-
long in another genus (Thompson, in preparation).
The remaining species placed in Hypselostoma differ
strikingK from //. holimanac. although H. tubiferum
(Benson, 1856) from Burma appears to be the most closely
related congener. It has a similarly broad umbilicus and
similar spiral sculpture on the teleoconch, though not as
strong. Because of these similarities the two species are
considered to be more closeK related than either is to
other known species w ithin the genus. The two differ in
several conspicuous features. Hypselostoma tubiferum
is much more depressed, the aperture is turned upw ard
above the apex of the spire and the aperture barrier
consists of 6-7 lamellar teeth (see Pilsbrv, 1917 for a
description and illustrations). Hypselostoma holimanae
differs from all other mainland species b\ its strong spiral
sculpture, its reduced aperture barrier with tubercular
columella lamella, its strong circumumbilical basal keel,
and its broad umbilicus. Some Philippine species {H.
sibuyanicum Moellendorff, 1896. H. quadrasi Moellen-
dorft, 1896, H. roebeteni Moellendorff, 1894, and H.
latispira Thompson and Auffenberg, 1984) are similar
to H. holimanae in that the aperture barriers have been
reduced to denticles, although they are not as weak as
in //. holimanac. None has sculpture as strongK- devel-
oped as //. holimanae. none has a basal keel circum-
scribing the umbilicus, nor are any as widely umbilicate.
Other differences that separate H. holimanae from these
Philippine species are the shape of the shell and the
contour of the whorls. Hypselostoma roebeleni and H.
latispira are \er\ depressed species in which the up-
turned aperture reaches almost to the level of the apex
of the spire. Hypselostoma sibuyanicum and H. quadrasi
have more slender shells w ith regularh increasing whorls,
and both are narrovvlv rimate. The combination of all
of these morphological differences indicate that the sim-
ilar aperture barriers of the Philippine species and H.
holimanae have evolved independentlv through tooth
reduction from more comple.x ancestral conditions, and
that no close relationship can be inferreil on the basis of
these barriers.
ACKNOWLEDGEMENTS
We wish to thank Mrs. Stanley (Bonnie) Holiman for
generously donating the holotvpe and part of the para-
type series to the Morida State Museum and to Mahidol
University. Stephen Holiman provided us with infor-
mation about the type localitv. The illustrations com-
prising figures 1-3 were rendered bv Ms. Wendy B.
Zomlefer, Staff Illustrator, Florida State Museum.
LITERATURE CITED
Jutting, VV. S. S. van Bentliem 1950. The Malavan species
of Boysidia. Paraboysidia, Hypselostoma, and Gyliotra-
F. G. Thompson and H. G. Lee, 1988
Page 81
chela (Gastropoda. Pulnioiiata, Nertiginidae) with a cat-
alogue of all the species hitherto described Bulletin of the
Raffles Museum 21 5-47
Jutting, W S. S. van Benthem 1961. Additional new species
and new localities of the family X'ertiginidae and the gen-
era Oophana and Opisthostoma from Mala>a. Bulletin of
the Raffles Museum 26:34-48, pis. 8-14.
Jutting, W. S. S, van Benthem. 1962. Coquilles terrestres
nouvelles de quelques collines calcaires du Cambodje et
du Sud X'ietnani. Journal de Conchyliologie 102:3-15.
Pilsbrv, Henr\ .\. 1916-18. Manual of conchology, Ser. II,
24. Pupillidae, Gastrocoptinae. Philadelphia, p. i-xii, 1-
380, pis. 1-49.
Solem, A. 1981. Small land snails from Northern Australia,
1: species of Gtjiiotraclicia Tonilin, 1930 (Mollusca, Pul-
monata, Nertiginidae). Journal ot the Malacological So-
ciety of Australia 5:87-100
Thompson, F. G. and K. Auffenberg 1984. Hypselostoma
latispira, a new pupillid land snail Irom the Philippine
Islands. Proceedings, Biological Societv of Washington 97:
86-89.
Thompson, Fred G. and S. Peter Dance 1983. Non-marine
mollusks of Borneo, II. Pulmonata: Pupillidae, Clausili-
idae. III. Prosobranchia: Hydrocenidae, Helicinidae Bul-
letin Florida State Museum 29:101-152.
THE NALTILL'S 102(2):82-87, 1988
Page 82
Eledone gaucha, a New Species of Eledonid Octopod
(Cephalopoda: Octopodidae) from Southern Brazil
Manuel Haimovici
Dt-partamento de Oceaiiografia
Fuiida^ao Universidadf de
Rio Grande
Cx. Postal 47-1, Rio Crande RS 96.200
Brazil
ABSTRACT
A lieu species of Eledone is described from tlie southwestern
Atiaiitif at depths of 60 to 160 m, off Rio (Jrande do Sul, Brazil.
The characters that distinguish this species from the other species
of the genus are presented, as well as a morphometric com-
parison with the sympatric Eledone massyae Voss, 1964.
INTRODUCTION
Several cephalopods were collected during a survey of
the demersal resources of the inner shelf of Rio Grande
do Sul between Solidao (30°40'S) and Chui (34°20'S)
at depths to 100 m (figure 1) by the R/V "Atlantico Sul"
of Funda^ao Universidade de Rio Grande (FURG). Hai-
movici and Andriguetto (1986) stated that two species
of the octopod genus Eledone were found. One of them,
E. rnassijae, was described by \ oss (1964) and the second
was a new species. Both sympatric species possess the
generic character of papillae at the tips of the non-
hectocot) iized arms of the males. Morphological analysis
presented here as well as ijiochemicai dilferences fouml
by Levi et al. (1985) separate these two similar species.
MATERIALS AND METHODS
All specimens studied were killed with fresh water, fixed
in 10% formalin for 24 hr and preserved in 70% ethanol.
Measurements were taken in millimeters, and all mea-
surements and indices used are among tho.se described
by Roper and Voss (1983). The drawings of £. gaucha
are by Jose Angel Alvarez Perez. The types are deposited
in the Museu Oceanografico de Rio Grande (MORG),
Museu Nacional de Rio de Janeiro (MNRJ), Museu de
Zoologia da L ni\ersidade de Sao Paulo (MZUSP), Museo
Nacional de Historia Natural, Liruguay (MNHN), Museo
de Ciencias Naturales de La Plata, Argentina (MCNLP),
University of Miami Mollusks Laboratory (UMML) and
National Museum ot Natural History, Smithsonian In-
stitution (USNM).
This work has been partially supported by a grant of
the Conselho Nacional de Pesquisas Cientificas e Tec-
nologicas (CNPq), Proc. 403293-83 and the surve\ fi-
nanced bv the Comissao Interministerial de Recursos do
Mar (CIRMj.
Eledone gaucha new species
(figures 2-14, table 1)
Material examined: Holot\ pe; 6 32.5 mm ML, K \'
Atlantico Sul, cruise 13/83, S'ta. 37, 32°58'S, 5ri9'W, 56
m, trawl, 17 Nov. 1983, MORG 23544. Parat\pes: 7 6
27-41 mm ML and 5 9 32-34 mm ML, R \' Atlantico
Sul, cruise 13/83, Sta. 37, 32°58'S, 51°19'W, 56 m, trawl,
MORG 23544; 2 <5 32-34 mm ML, R/\' Atlantico Sul,
cruise 13/83, Sta. 39, 32°50'S, 50°45'W, 87 m, trawl, 18
Nov. 1983, MORG 23545: 1 S 21 mm ML. R \" Atlantico
Sul, cruise 13/83, Sta. 2, 31°46'W, 100 m, 9 Nov. 1983,
MORG 23846; 1 9 33 nun ML, R/\' Atlantico Sul, cruise
10/83, Sta. 51, 33°43'S, 52°13'W, 60 m, trawl. 29 Aug.
1983, MORG 23547; 4 9 28-33 mm ML, R/\' Atlantico
Sul, cruise 10/83, Sta 57, 33°13'S, 51°25'\\'. trawl, 60 m,
30 Aug. 1983, MORG 23548; 1 <J 33 mm ML and 1 9 44
mm ML, R/V Atlantico Sul, cruise 10/83, Sta. 53, 33°52'S,
5r55'W, 52 m, trawl, 29 Aug. 1983, UMML 32.2066;
1 6 34 mm ML and 1 9 42 mm ML, R \' Atlantico Sul,
cruise 10/83, Sta. 53, 33°52'S, 5r55'W, 52 m, trawl,
29 Aug. 1983, USNM 816613; 1 <5 26 mm ML and 1 9
29.5 mm ML, R/V Atlantico Sul, cruise 10/83, Sta. 51,
33°43'S, 52''13'W, 74 m, trawl, 29 Aug. 1983, MZUSP
25242; 1 S 32 mm ML, R/\' Atlantico Sul, cruise 10/83,
Sta. 55, 32°52'S, 51°43'\V, 140 m, trawl, 30 Aug. 1983,
MGNLP 4681; 1 9 29.5 mm ML, R/V Atlantico Sul,
cruise 10/83, Sta. 57, SS'IS'S, 51''25'W, 60 m, trawl, 30
Aug. 1983. MC:NLP 4682; 1 <? 30 mm ML and 1 9 38
mm ML, R/ \' Atlantico Sul, cruise 10/83, Sta. 53, 32°52'S,
51°55'W, 52 m, trawl, 29 Aug. 1983, MNHN 14762,
MNHN 14763; 1 S 35 mm ML and 1 9 42 inm ML, R/V
Atlantico Sul, cruise 3/83, Sta. 42, 33°02'S, 51°30' \V, 130
m, trawl, 17 June 1980, MNRJ 5626, MNRJ 5627.
M. Haimovici, 1988
Page 83
DESCRIPTION
Animal small, maximum observed mantle length 65 mm
(figures 2, 3). Mantle firm, not very thick, ovoid and
elongated (MVVl 3: 61.0; 2: 59.2) separated from head
by small constriction. Body surface smooth with some
papillae on the dorsal mantle and head. Head narrow er
than mantle (HWl S: 36.2; 9: 37.3); eyes slightly protu-
berant. One supraocular cirrus. Funnel long (FLI S: 44.6;
9: 45.4) with anterior half free (FFuLI 6: 21.6; 5: 21.1);
funnel organ (figure 4) W shaped.
Arms long and rather slim. Arms length order
1 >2>3>4 in most specimens \\ ith dorsal arms markedly
longer than others. All arms longer in males (except hec-
tocotylized arm) than in females (ALl I to IV S: 273.2-
239.8-134.9-208.1; 9: 250.0-213.8-200.6-191.0).
Suckers small, uniserial, well separated and deeply set
into arms. Suckers somewhat crowded near tips of the
arms on females. Two rows of minute fleshy papillae on
all non-hectocotylized arms of males (figure 10). Number
of suckers on basal half of the first right arms varies from
17 to 23 (ASC 6: 20.1; 9: 19.4), suckers slightly larger on
all arms of males (lASI S: 7.7; 9; 6.5).
Web extends over half the length of arms and de-
creases from dorsal to ventral surface; web formula most
trequentlv A:B:C:D:E. Web indices similar in both sexes;
24.4-24.1-21. 6-18.6-15.6 for males and 24.4-23.9-21.1-
18.3-14.9 for females.
Third right arm in males hectocotylized (figure 9)
(HcAI: 58.9). Ligula small (LLI: 8.8), without differen-
tiated calimus (figure 9); spermatophore grove deep.
Gill count in external hemibranch from 7 to 10, most
trequentlv 8 in males and 9 in females (Gilc S: 8.2; 9:
8.9).
Males reproductive system with no special figures (fig-
ure 11). Penis long and tubular (PLI: 23.4) with a rather
short diverticulum (PdLI: 7.8). Spermatophores undif-
ferentiated (figure 13) from 12 to 20 mm (SpLI: 45;
SpLWT: 1.56). Number of spermatophores in a sample
of 40 mature males from 7 to 92 (mean 32.1).
The proximal o\ iduct long, the oviductal glands small,
the distal oviducts shorter and somewhat stouter (figure
12). Intraovaric eggs oval (figure 14); maximum lengths
of apparently mature eggs approximately 8 mm. Num-
ber of developing eggs in a sample of 42 maturing fe-
males ranged 10-55 (mean 30.2).
Buccal mass well developed, with small anterior sali-
vary glands and larger posterior salivary glands. Esoph-
agus connects to developed crop leading to muscular
stomach and smaller spiral caecum united by two ducts
to the large digestixe gland. Intestine thin and leads to
the anus adjacent to ink sac opening. Ink sac superficialK
embedded in the digestive gland (figure 7). Radula with
a tricuspid rachidean tooth, three lateral teeth and a
marginal plate (figure 8).
Color of living animals changed from brow n to almost
white dorsally always remaining clear ventrally. Color
of specimens preserved in alcohol purplish gray dorsally
and pale \ellow ventralK'. Inner surface of the arms,
mouth, and ventral mantle with few chromatophores.
29°
\^
'vBRaSIL ,
)
M
^.\ J
^^ PORTO (N "sy
■*==5X5^«1-EGBE U J
30"
1
.320
V<^'.'/
V^
\:
k
(1
soLmAoy.
5^0
1 ")" '/ *''
31°
<^
r^
32°
>-
tot
^J/
fGRANDE
IJ
K
)^
/
'
V,
•
/
J^r^ /^
oi
•
,,-— '
> oW/
!>l
■
33°
t
/
7
' • . -e-
ARROIO CHuf^
?
'.■' o
/
•■. ■■
.■ • o
34°
y
--■■.
Ci
i
-=-1 r
— 1 — 1 — I
50t 50m
1 1 1 r
__^
.00*
54° 53° 52° 51° 50°
Figure 1. Sample localities of Eledone gaucha new species.
Type locality: 32°58'S, 51°19'W, south Rio Grande, Bra-
zil in 56 m.
Etymology: The name gaucha refers to the coastal planes
of Argentina, Uruguay, and southern Brazil and its peo-
ple.
Distribution: Eledone gaucha is knov\n onK from off
Rio Grande do Sul between Solidao (30°40'S) and Chui
(34°20'S) (figure 1).
DISCUSSION
The new species belongs in the genus Eledone because
of a single row of suckers, the heteromorphic arms in
the males, with the non-hectocot\ lized arms having the
suckers at their tips modified into fleshy papillae or lam-
inae, and the hectocot\ lus without a differentiated cal-
imus. These characters distinguish Eledone from related
genera Pareledone, Vosseledone, Graneledone and oth-
ers (Palacio, 1978).
The genus Eledone occurs on the Atlantic continental
shelves of South America. Africa, and Europe and in the
Mediterranean Sea. It includes six described species: E.
moschata (Lamarck, 1798) and £. cirrhosa (Lamarck,
Page 84
THE NAUTILUS, Vol. 102, No. 2
;-. V" :■■ r;" ■■■:1'. . ' Vj
-'T^
■r \ mm ....
!5
I,,
10
M. Haimovici, 1988
Page 85
Table 1. Ranges and means of measurements and
Eledune gaticlui new species, from southern Brazil
indites of 10 males and 10 females each of Eledone massyae V'oss, 1964 and
Eledone mass,
yae \ dss
, 1964
Eledune gaucha new species
Males
Females
Males
Females
Lower
Upper
Lower
Upper
Lower
Upper
Lower
Upper
Index
limit
Mean
limit
limit
Mean
limit
limit
Mean
limit
limit
Mean
limit
Total length (TL)
148.0
176.4
207.0
195.0
218.2
236.0
81.0
123.2
137.0
100.0
131.9
178.0
Mantle length (ML)
45.0
54.0
63.0
60.0
65.3
71.0
21.0
31.6
41.0
28.0
35.5
50.0
Mantle width index (MWI)
65.1
76.2
88.2
75.8
79.3
88.9
47.6
61.0
71.9
40.0
59.2
70.0
Head width index (HWI)
40.4
47.4
56.5
38.2
41.9
45.0
34.2
37.5
40.0
28.0
36.2
43.6
1° right arm length index
(1 ALI)
184.0
201 4
216.0
185.0
209.9
247.0
244.0
275.2
322.0
227.0
250.0
284.0
2° right arm length index
(II .ALII
195.0
203.3
235.0
190.0
217.3
248.0
193.0
239.8
281.0
162.0
213.8
242.0
3° right arm length index
(III ALI)
155.0
199.1
225.0
185.0
214.1
2.52.0
85.0
134.9
159.0
155.0
200.6
2.36.0
4° right arm length index
(IV ALI)
191.0
204.0
224.0
186.0
215.5
243.0
134.0
208.1
256.0
162.0
191.0
213.0
Arm formula (AF)
4:2:1:3
2:4:3:1
1:2:3:4
1:2:3:4
A web depth index (A WDI)
19.1
24.3
28.5
19.8
22.5
27.4
16.8
24.2
30.9
18.9
24.4
28.6
B web depth index (B WDI)
18.2
24.5
27.5
21.0
23.8
29.6
19.0
24.1
30 1
18.9
23.9
25.8
C web depth index [C WDI)
22.9
25.9
28.3
21.2
24.1
28.9
18.8
21.6
28.9
17.6
21.1
23.7
D web depth index (D WDI)
22.7
24.8
28.6
20.9
24.3
27.6
14.6
18.6
23.0
15.8
18.3
20.5
E web depth index (E WDI)
13.4
20.6
25.3
18.6
20.8
22.9
11.5
15.6
21.6
12.3
14.9
17.4
Web formula (WE)
C;D:B:A:E
D:C:B:A:E
A:B:C:D:E
A:B:C:D:E
Gill lamellae count (GiLC)
8/10
9.4/9.2
11,7
9/10
9.7/9.6
8/10
7/9
8.2/8.5
9/9
8/8
8.9/8.5
10/10
Funnel length index (ELI)
37.8
41.9
45.9
40.8
44.2
48.2
36.6
44.6
51.9
32.5
45.4
51.7
Free funnel length index
(FFuLI)
17.5
23.2
29.5
20.9
23.8
27.9
14.6
21.6
31.0
10.0
21.1
30.3
Arm sucker count (ASC)
16
18.1
20
16
17.8
20
17
19.4
22
18
20.1
23
Arm sucker index I (I ASI)
6.3
i . i
9.1
7.0
7.9
8.7
4.7
7.7
9.3
6.0
6.5
8.0
Arm sucker index II (II ASI)
6.3
8.0
9.1
7.0
8.2
9.6
4.7
7.3
9.3
5.0
6.4
8.0
Arm sucker index III
(III ASI)
5.5
7.4
8.9
7.0
7.9
8.5
4.7
7.6
9.6
5.0
6.0
7.5
Arm sucker index I\'
(IV ASI)
5.5
7.2
8.9
6.7
7.7
8.4
4.7
6.7
7.8
3.0
5.1
6.7
Penis length index (PLI)
19.3
28.1
39.3
19.5
23.4
28.1
Penis diverticulum length index
(PdLI)
6.6
15.0
22.2
4.9
7.8
10.5
Spermatophore length index
(SpLD
20.9
31.6
37.7
38.3
45.0
50.0
Spermatophore w idth index
(SpWI)
1.8
2.5
3.1
1.2
1.6
2.1
Hectocotvlized arm index
(HcAI)'
59.0
69.2
81.4
41.7
58.9
67.4
Ligula length index (LLI)
5.3
80
97
49
8,8
12.5
1798) both from the NE Atlantic and Mediterranean, £.
thijsanophora Voss, 1962 and E. caparti Adam, 1950
from the SE .\tlantic, and E. massyae Voss, 1964 and £.
gaucha trom the SW .Atlantic.
Eledone gaucha seems to be a relatively abundant
species off Rio Grande do Sul. The possible reasons why
it has not been recognized to date are its small size, its
similarity to E. massyae, and the scarcity of scientific
cephalopod collections by research vessels in this area.
Due to its small size it is not retained in the cod ends of
commercial trawlers and e\en in the R \' Atlantico Sul
surveys it most often was found entangled in the wings
of the net. Palacio (1977) reviewed several museum col-
lections of Argentina, Uruguay, and Brazil and found
Figures 2-12. Anatomical features of Eledone gaucha new species. 2. 3. Dorsal (2) and lateral (3) views of holot>pe (MORG
23544, 32 mm ML). 4. Funnel organ. 5. Upper mandible. 6. Lower mandible. 7. Digestive tract. 8. Radula. 9. Hectocot) lized
arm tip 10. \on-hectocot\ lized arm tip of male 1 1. Male reproductive organs. 12. Female reproductive organs.
Page 86
THE NAUTILUS, \ol. 102, No. 2
Figures 13, 14. Rpprocliictive products of Eledone gaucha
new species. 13. Sperinatophore. 14. Egg.
onl\ one eledoiiid. E. inaasyae. A survey of the MORG
collection by the author showed one specimen of E. gau-
cha (MORG 15'341) lornierK classified as E. massijae. It
is expected that reviews in other collections w ill expand
the range of £. gaucha.
The sympatric species E. nias.syae and E. gaucha ini-
tially look similar but many differences may be seen in
a more detailed stud\'. In ortler to compare both species
morphologicalK, the same indices were calculated for
10 males and 10 females of E. massyae {collected in the
same survey) which were fixed, preserved, and measured
in the same way as the new species (table 1). Eledone
gaucha is smaller antl has a narrower mantle and head.
The arms are thinner, longer, and decrease in size while
in £. massyae all arms are approximately the same length.
The hectocotvlized arm is shorter and the web depth
decreases troni the dorsal to the ventral surface in E.
gaucha. while in E. massyae the web is shorter only
between the ventral arms. The funnel organ is W shaped
in E. gaucha, w shaped in £. massyae. The number of
inner and outer gill lamellae is one unit lower and the
arms sucker count two units higher in E. gaucha. .Arm
sucker indices are similar in males of both species, but
in females they are smaller in E. gaucha. Perhaps the
best single diagnostic character to distinguish mature
males of both species is the spermatophore, which is
shorter and much thinner in £. gaucha. ExternalK, the
best distinctive character is the arm length pattern.
Eledunc caparti was described b\ .Adam (1950) based
on five specimens, two males and three females collected
in the equatorial west .Africa at depth ranging from 60
to 170 m. No figures or tables were included in the
original description. The decreasing arm length and web
depth of £. caparti are similar to those of the new species.
However, £. caparti does not have a supraocular cirrus,
has enlarged suckers at the base of the lateral arms of
the males, and the number ot suckers on the dorsal arms
is almost double that in E. gaucha. The radula of £.
caparti has an A2 seriation and the spermatophore is
insufficiently described for comparisons.
Eledone thysanophora was described by \'oss (1962)
based on a single male specimen collected in a tide pool
in western South Africa. The morphometric description
is short, but the number of papillae on the tips of the
non-hectocotylized arms and the structure of the sper-
matophore, w ith the inner w all ot the horn portion lined
with teeth, differentiate £. thysanophora irom the new
species.
Summary descriptions of £. cirrhosa (Lamarck, 1798)
and £. moschata (Lamarck, 1798) are presented in Roper
et al. (1984), and the species are compared b\ Rees
(,1956). Both species can be distinguished from £. gaucha
by several characters. Eledone cirrhosa has moderately
short arms, a ridge along the mantle, non-hectocot\lized
arms of males w ith a single row of compressed sucker-
like cirri, and spermatophores with spines. Eledone mos-
chata has subequal arms, 11 to 12 filaments on the outer
hemibranch of the gills, big, sausage-shaped eggs 15 mm
long, and a characteristic musk odor.
ACKNOWLEDGEMENTS
J. M. Andriguetto Filho and J. A. Alvarez Perez helped
with collection of the specimens, G. L. Voss and an anon-
ymous reviewer improved dramaticalK the final version,
and E. G. Rios stimulated m\ interest in mollusks. To all
of them m\' sincere gratituile.
LITERATI RE CITED
.Adam, W. 1950. Notes sur les cephalopodes .\.\11. Deux nou-
velles especes de la cote africaine occidentale. Bulletin
Institut Ro\ al des Sciences Naturelles de Beigique 2(i(45):
1-9.
Haimovici, M. and J. M .Andriguetto Fo. 1986. Celalopodes
costeiros capturados na pesca de arrasto do iitoral sul do
Brasil .Archives de Biologia e Tecnologia. Parana 29(3):
473-495.
Levi, J. A., M. Haimovici. and M. B. Concei^ao. 1985. Car-
acteriza9ao eletofortica de dois morfotipos do genero Ele-
done (Cephalopoda: Octopodidae). Resumes do XII Con-
gresso Brasileiro de Zooiogia. (lampinas. 27 Jan to 1 Feb.
1985:39.
Palacio, J. F. 1977, .A stud\ of the coastal ceplialopods from
Brazil with reference to Brazilian zoogeography. Ph.D.
thesis, University of Miami, 311 p.
M. Haimovici, 1988
Page 87
Palacio, J. F. 1978. Vosseledorie charrua. a new Patagonian
ceplialopod (Octopodidae) with notes on related genera,
bulletin of Marine Science 28(2):282-296.
Rees, W J. 1956. Notes on the European species ol F.lcdone
with special reference to eggs and larvae. Bulletin British
Museum of Natural Histor\ .3(6):283-292, pis. 9-10,
Roper, C. F. E., M. J. Sweene>, and C. E. Nauen. 1984. FAO
species catalogue, Vol. 3. Cephalopods of the world. FAO
Fisheries Synopsis No. 125, 3:227.
Roper, C. F. E. and G. L. Voss. 1983. GuideHnes for taxonomic
descriptions of cephalopod species. Memoirs of the Na-
tional Museum of Victoria 4449-63.
Voss, G. L. 1962. Soutli African cephalopods. Transactions of
the Royal Societ)' of South Africa 36(Part 4):245-272.
Voss, G. L. 1964. A note on .some cephalopods from Brazil
with a description of a new species of octopod: Eledone
massyae. Bulletin of Marine Science of the Gulf and Ca-
ribbean 14:511-516.
THE NAUTILUS 102(2);88, 1988
Page 88
News and Notices
.54TH ANNUAL MEETING OF THE AMERICAN
MALACOLOGICAL UNION
CHARLESTON. SOUTH CAROLINA
RADISSON FRANCIS MARION HOTEL
June 19-24, 1988
The 54th annual meeting of the American Malacological
Liiioii \\ ill be held June 19-24, 19SS in (Charleston, South
Carolina. Charleston is a historical cit\, many parts of
which have been beautifully restored, as has the Radisson
Francis Marion Hotel, which is located downtown, with-
in walking distance of many restaurants, shops and other
attractions. Charleston is easily accessible both by air and
b\ interstate highway.
Three symposia are planned: Applications of Nucleic
Acid Techniques to the Study of Molluscan Evolution,
convened b> Dr. M.G. Harasewych, Department of In-
vertebrate Zoology, National Museum of Natural His-
tory, Smithsonian Institution; S\ stematics and Evolution
of Non-marine Mollusks, convened by Dr. Robert
Hershler, Department of Invertebrate Zoology, National
Museum of Natural History, Smithsonian Institution; and
History of Malacolog\, convened by Dr. W. Backhuys,
E.J. Brill, Inc Leyden, The Netherlands.
In addition to these ssmposia, contributed papers and
poster presentation, scheduled events w ill include a tour
of historic Charleston, guided field trips to terrestrial and
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For further information, please contact:
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President, AMU
P.O. Box 30
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Telephone (803) 249-1454
16TH CONCHOLOGISTS OF AMERICA
CONVENTION
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SHERATON HARBOR HOUSE HOTEL
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THE NAUTILUS
Volume 102, Number 3
August 29, 1988
ISSN 0028-1344
A quarterly devoted
to malacology.
Marine Biological Laboratory .
LIBRARY
SEP 6 1988 i
Woods Hole, Mass.
EDITOR-IN-CHIEF
Dr. M. G. Harasawych
Division of Mollusks
National Museum of
Natural Histor\
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Washington, DC 20560
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.■\merican Malacologists, Inc.
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Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
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National Museum of
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THE €9 N An^ LJicaiXio/ i L U S
CONTENTS
j LIBRARY
SEP 6 1988
Volume 102, Number 3
August 29, 1988
ISSN 0028-1344
Woods Hole, Mass.
Timothy M. Askew
A new species of pleurotomariid gastropod from the
western Atlantic
89
IM. G. Harasewych
Shirley A. Pomponi
Timothy M. Askew
Spongivory in pleurotomariid gastropods .
92
James H. McLean
Ricardo Silva Absalao
Renato Luiz
dos Santos Cruz
A new species of Macrarene (Turbinidae: Liotiinae) from
Brazil
99
Geerat J. Vermeij
Timothy M. Collins
Nerita fortidentata, a new gastropod from the Neogene of
Panama, with comments on the fossil record of Nerita in
tropical America
102
Anthony D^Attilio
Barbara W. Myers
A new species of Favartia from the eastern Pacific
(Gastropoda: Muricidae)
106
William K. Emerson
Salter E. Sage III
Conus baccatus G. B. Sowerby III, 1877: A Panamic
faunal constituent
110
Kenneth J. Boss
References to Molluscan Taxa Introduced by Linnaeus in
the Systema Naturae (1758, 1767)
115
Mark E. Gordon
Frederick Benjamin Isely: Biographical Sketch and
malacological contributions
123
William F. Adams
Andrew G. Gerberich
Rediscovery of Planorbella magnifica (Pilsbry) in
southeastern North Carolina
125
Donald M. McKinstry
Bite by Octopus joubini: a case report 127
Oscar J. Polaco
Wolfgang Mendl
Occurrence of mites in Mexican land snails 129
Richard E. Petit
Axelella, new name for Olssonella Petit, 1970, a
preoccupied taxon (Mollusca: Cancellariacea)
130
THE NAUTILUS 102(3):89-91, 1988
Page 89
A New Species of Pleurotomariid Gastropod
from the Western Atlantic
Timothy M. Askew
Harbor Branch Oceanographic
Institution, Inc.
5600 Old Dixie Highway
Fort Pierce, FL 34946, USA
ABSTRACT
Perotrochus charlestonensis, a new species of pleurotomariid,
is described from off the coast of South Carolina. This is the
12th Recent pleurotomariid taxon to be described from the
western Atlantic. Its habitat is described, environmental data
are provided, and comparisons made with closely related con-
geners.
Key words: Gastropoda; pleurotomariid; Perotrochus; slit shells;
western Atlantic; JOHNSON-SE,\-LINK
INTRODUCTION
Since the discovery of the first living species of the pre-
dominant!) Mesozoic and Paleozoic family Pieuroto-
mariidae in the western Atlantic over a century ago
(Fischer & Bernard!, 1856), 24 Recent species and sub-
species have been described, usually on the basis of one
or a very few specimens. The habitat of these animals,
generally steep-wailed, hard substrates at depths in e.xcess
of 100 meters, accounts for their infrequent collection
by such methods as trawling, dredging, and grab sam-
pling, and, therefore, for the paucity of data on the
biology and distribution of most species. Since the pub-
lication of a review of the Recent pleurotomariids that
included six species from the West Indies (Bayer, 1966),
three species (Bayer, 1967; Rios & Mathews, 1968; Leme
& Penna, 1969) and two subspecies (Okutani & Goto,
1983, 1985) have been described from the western At-
lantic.
Another new species of pleurotomariid, described
herein, was collected while conducting fish population
studies approximately 90 nautical miles east of Charles-
ton, South Carolina, utilizing the submersible JOHN-
SON-SEA-LINK I (Harbor Branch Oceanographic In-
stitution, Inc., Fort Pierce, Florida). Bottom topography
at the stud\ area was extremely rugged, making sampling
by any other means difficult.
SYSTEMATICS
Family Pleurotomariidae Swainson, 1840
Genus Perotrochus P. Fischer, 1885
Perotrochus charlestonensis new species
(figure 1)
Description: Shell (figure 1) moderately large (maxi-
mum diameter 87.4 mm, minimum diameter 80.1 mm,
height 73.0 mm), broadly turbiniform, very thin, fragile;
spire angle 89°, spire slightly convex in profile; proto-
conch of 1.0 whorls, translucent, glassy; transition to te-
leoconch marked by axial costae, with selenizone ap-
parent by second postnuclear whorl; teleoconch of 8'/3
whorls; early whorls nearly flat-sided, becoming pro-
gressively more inflated; selenizone near suture in early
whorls, shifting to slightly below mid-whorl by fifth post-
nuclear whorl; anal slit depth at upper margin 89°, at
lower margin 57°; anal slit width 4 mm; suture adpressed;
periphery rounded; base inflated, convex, non-umbili-
cate; nacreous umbilical callus extending '/6 the distance
from axis to periphery; spiral sculpture of 21 uniformly
sized spiral cords between suture and anal slit, 20 cords
of variable thickness between anal slit and periphery, 40
cords along base; selenizone with 0-5 broad cords, num-
ber increasing with shell size; axial sculpture of weak
nodes on early whorls (88 on fourth postnuclear whorl),
forming cancellate sculpture; axial sculpture decreasing,
sculpture limited to spiral cords by sixth postnuclear whorl
above selenizone, and seventh postnuclear whorl below
selenizone; aperture broadly ovate; columellar lip slightly
thickened, weakly recurved; color creamy white with
diffuse brownish orange axial streaks and blotches; na-
creous layer visible through porcellaneous layer, creating
iridescent hue; color lighter on base than on dorsal sur-
face; selenizone margins with cream colored lines most
evident on penultimate and body whorls; aperture na-
creous, iridescent; operculum multispiral (7 whorls),
horny, brownish-yellow, translucent; soft parts unknown.
Type localitv: 90 nautical miles east of Charleston, South
Carolina (32°43'80"N, 78°05'60"W), in 213 m, R/S
JOHNSON-SEA-LINK I, dive 1250, August 6, 1982.
Holotype: USNM 859961, maximum diameter 87.4 mm.
Etymology: Named after the t\pe localit\ , commonly
referred to as the Charleston Lumps.
THE NAUTILUS, Vol. 102, No. 3
Figure 1. Perotrochus charlestunensis new species. Apertural, right lateral, apical, and basal views of the holotype (USNM 859961 ),
off Charleston, South Carolina (32°43'80"N, 78°05'60"W), in 213 m, maximum shell diameter 87,4 mm.
Ecology: This species is known only from the type lo-
cality, an area of extremely rugged terrain where the
bottom topograph}' consists of steep, large hills and val-
leys. Topographical features (figure 2) consist of a pave-
ment of relithified phosphorite and fibrous concretionary
apatite composed of calcium phosphate and other min-
erals (Manheim et ai, 1980). This pavement, which ranges
in thickness from 10 cm to almost a meter, has been
undermined in some areas, causing large pieces to break
off and fall down-slope forming rubble and boulder zones.
Hills range in height from several meters to about 30
meters. Valleys contain sand composed primarily of brown
to black phosphorite. This area was formed during the
middle Tertiary and has remained stable since the Mio-
cene (Baturin, 1982).
Marine life in the vicinity indicates an area of high
productivity resulting from warmer Gulf Stream waters.
Large snowy grouper [Epinephehts niveatus (Valen-
ciennes, 1828)] and blue-lined tile fish [Caulolatilus mi-
crops (Goode & Bean, 1878)] are abundant in the area
together with many species of small, deep-reef fish, which
generally inhabit rocky terrain. Common invertebrates
include basket and brittle stars, sea urchins, solitary and
colonial anemones, solitar\ corals, arrow, spider, and gal-
T. M. Askew, 1988
Page 91
RCXKSOULOER LECX3E
HOCK RUBBLE
'-a.-..
i^^, SAND PEBBLE
BARREN SAND
Figure 2. Cross-section of bottom topography at collection
site. Hills range in height from 3 to 30 m.
atheid crabs, barrel and encrusting sponges, and hy-
droids. A pink featherlike hydroid covers many of the
broken boulders along the ridge tops. Prevalent gastro-
pods include Perotrochus amabilis (Baser, 1963), Cal-
liostoma sayana (Dall, 1889), Stenorhytis pernobilis
(Fischer & Bernardi, 1857), Aurinia gouldiana (Dall,
1887), and Pterynotus phaneus (Dall, 1889).
Remarks: Perotrochus charlestonensis is a member of
the species complex consisting of P. midas Bayer, 1966,
P. pyramiis Bayer, 1967, P. africanus (Tomlin, 1948),
P. teremachii (Kuroda, 1955), P. tangaroana Bouchet &
Metivier, 1982, and an undescribed species from off
northwestern Australia (Group B, Bayer, 1966:745). All
are characterized by having large, thin shells with in-
flated whorls and proportionally large, broadly ovate ap-
ertures. This new species most closely resembles P. af-
ricana and Perotrochus sp. (Bayer, 1966; fig. 29) from
Japan, but differs from these taxa in having a thinner
shell w ith more inflated whorls, and a more convex profile
of the spire. Perotrochus africanus has a more stepped
spire, a more strongly recurved and thicker columella,
and a broader umbilical callus (Vi distance from axis to
periphery). Of the western Atlantic species, P. charles-
tonensis is most similar to P. pyramus. but is more than
twice the size, and is much higher-spired. Perotrochus
charlestonensis also somewhat resembles Perotrochus
midas, but lacks its characteristic flat, blunt spire and
angular periphery. Perotrochus charlestonensis occurs
in shallower water (213 m) than either P. pyramus (420-
648 m) or P. midas (600-770 m).
ACKNOWLEDGEMENTS
I thank the following persons, without whose help and
guidance this paper would not be possible: Dr. Richard
Cooper, University of Connecticut; Dr. Roger Theroux
and Joseph L zmann. National Marine Fisheries Services,
Woods Hole, Massachusetts; Dr. Walter Nelson and El-
mer Gutherz, National Marine Fisheries Service, Pas-
cagoula, Mississippi; and Tom Smoyer, Harbor Branch
Oceanographic Institution photographer, for his excel-
lent photographs. This is Harbor Branch Oceanographic
Institution Contribution No. 642.
LITERATURE CITED
Baturin, G. N. 1982. Phosphorites on the sea floor — origin,
composition and distribution. Developments in Sedimen-
tology 33:111-112.
Bayer, F. M. 1963. A new pleurotomariid gastropod trawled
in the Straits of Florida by R/V Gerda. Bulletin of Marine
Science of the Gulf and Caribbean 13(3):488-492.
Bayer, F. M. 1966. New pleurotomariid gastropods from the
western Atlantic, with a summary of the Recent species.
Bulletin of Marine Science 15(4):737-796.
Bayer, F. M. 1967. Another new western Adantic pleuroto-
marian gastropod. Bulletin of Marine Science 17(2):389-
397.
Bouchet, P. and B. Metivier. 1982. Living Pleurotomariidae
(Mollusca: Gastropoda) from the South Pacific. New Zea-
land Journal of Zoology 9:309-318.
Dall, VV. H. 1887. [A letter containing notes on Antillean
mollusks.] Conchologist's Exchange 2(1):9-10.
Dall, W. H. 1889. Reports on the results of dredging ... in
the Gulf of Mexico (1877-1878) and in the Caribbean Sea
(1879-1880), by the U.S. Coast Survey steamer "Blake"
. . . XXIX. Report on the Mollusca. Part II. — Gastropoda
and Scaphopoda. Bulletin of the Museum of Comparative
Zoology, Harvard 18:1-492, pis. 10-40.
Fischer, P. 1885. Manuel de conchyliologie et de paleonto-
logie conchyliologique. Histoire naturelle des mollusques
vivants et fossiles. Fascicule 9:785-896. Libraire F. Savy,
Paris, xxiv -I- 1369 p., 23 pis. (1887).
Fischer, P. and A. C. Bernardi. 1856. Description dun pleu-
rotomaire vivant. Journal de Conchvliologie 5:160-166,
pi. 5.
Fischer, P. and A. C. Bernardi. 1857. Descriptions d'especes
nouvelles. Journal de Conchyliologie 5:292-300, pis, 8, 9.
Kuroda, T. 1955. A new Pleurotomaria from Japan with a
note on a specimen of P. rumphii Schepman collected
from Taiwan. Venus 18(4):21 1-221, pis. 8, 9,
Leme, J. P. L. and L. Penna. 1969. Ocorrencia de Mikado-
trochus no Brasil com descri^ao de uma nova especie.
Papeis Avulsos de Zoologia (Sao Paulo) 22(21 ):225-230.
Manheim, F. T., R. M. Pratt, and P. F. McFarlin. 1980. Com-
position and origin of phosphorite deposits of the Blake
Plateau. The Society of Economic Paleontologists and Min-
eralogists, Special Publication No. 29:117-137.
Okutani, T. and Y. Goto. 1983. A new subspecies of Adanson's
slit shell from Bermuda. Venus 42(4):305-311.
Okutani, T. and Y. Goto. 1985. A new subspecies of Pero-
trochus quoyanus from Bermuda. Venus 44(1):27-31.
Rios, E. C. and H. R. Mathews. 1968. Nova especie de Pleu-
rotomariidae do Brasil (Mollusca: Gastropoda). Arquivos
Estaceo de Biologia Marinha da Universidade Federal do
Ceara 8(l):65-68.
Tomlin, J. R. le B. 1948. A new species of Pleurotomaria.
Journal of Conchology 23:2, pi. 1.
THE NAUTILUS 102(3):92-9S, 1988
Page 92
Spongivory in Pleurotomariid Gastropods
M. G. Harasewych
Department of Invertebrate Zoology
National Museum of Natural History-
Smithsonian Institution
Washington, DC 20560, USA
Shirley A. Pomponi
Timothy M. Askew
Harbor Branch Oceanographic
Institution, Inc.
5600 Old Dixie Highway
Ft. Pierce, FL 34946, USA
ABSTRACT
Direct in situ observations of feeding together with analyses
of the gut contents of Perutruchus ruidas and P. amabilis in-
dicate that these species feed predominantK and selectively on
sponge tissue. Foraminiferal and diatom tests previously re-
ported in the gut contents of pleurotomariids are derived from
planktonic sediment coating the surfaces of sponges and do not
significantK contribute to the nutrition of these gastropods. The
family Pleurotomariidae represents an adapti\e radiation to
spongivory that most likely occurred after the divergence of
the herbivorous Scissurellidae and Haliotidae from the pleu-
rotomariid progenitor in the late Paleozoic, but prior to the
appearance of umbilicate pleurotomariids with deep anal ca-
nals in the Upper Jurassic.
Key words: Pleurotomariidae; diet; sponges; spongivory; Pero-
trochus; Mikadotrochus, Entemnutrochus.
INTRODUCTION
Since the discovery of the first living species of the family
Pleurotomariidae in the mid-nineteenth century, nu-
merous papers have been published on various aspects
of the biology and anatomy of these, the most primitive
living gastropods (e.g., Bouvier & Fischer, 1899, 1902;
Woodward, 1901; Fretter, 1964, 1966; Yonge, 1973;
Hickman, 1984a, b). As pleurotomariids are predomi-
nantly restricted to hard substrates and bathyal depths
in the Recent fauna, sampling has pro\ed difficult and
most published observations are based on limited ma-
terial, often poorly preserved.
The common occurrence of sponge spicules, forami-
niferal tests, and diatoms in the alimentary systems and
fecal pellets of pleurotomariids has been variously in-
terpreted as being indicative of a spongivorous diet con-
sisting of a single species of sponge (Woodward, 1901:
252); a diet consisting principally of sponges (Thiele,
1935:1129; Hyman, 1967:.360; Yonge k Thompson, 1976:
52); a diet of encrusting invertebrates, predominantly
sponges (Hickman, 1984a:29); or detrital feeding and
vegetarian diet (Fretter & Graham, 1976:1). The pres-
ence of sponge spicules in the gut contents, however, is
not necessarily evidence of spongivory. Although mem-
bers of the pleurotomariacean family Haliotidae are
known herbivores (Leighton, 1961; Leighton & Boo-
lootian, 1963; Shepherd. 1973; Fretter & Graham, 1976:
6), Leighton and Boolootian (1963:229) reported sponge
spicules, comprising 1-2% of the gut content \olume, in
25-50/c of the specimens of Haliotis cracherodii they
examined. Similarly, Herbert (1987:289) identified sponge
spicules, foraminiferans, annelid setae, and crustacean
remains in the alimentary systems of members of the
Solariellinae (Trochidae), a group adapted to feeding on
superficial and interstitial detritus. Three species of pleu-
rotomariids maintained in aquaria have been reported
to feed on a wide variety of foods, including starfish,
bivalve meat (Arakawa et al., 1978) and sliced, raw fish
(Matsumoto et al, 1972; Sekido et al, 1976).
The increasing use of research submersibles in deep
sea investigations has made possible in situ observations
on the feeding of two species of pleurotomariids. These
observations, supported by gut content analyses of col-
lected voucher material, form the basis of this report.
MATERIALS AND METHODS
During the course of numerous di\es aboard the research
submersible JOHNSON-SEA-LINK-II throughout the
northern and central Bahamas, li\ ing Perotrochus midas
Bayer, 1966, were observed on 16 occasions at depths
ranging 670 to 853 meters. Observations were noted and
specimens photographed when practical, using a BEN-
THOS 35 mm camera, with 85 mm lens. Two of the
Figures 1, 2. Perotrochus midas Bayer, 1966, and Strongylophora hartmani \'an Soest, 1981, in situ. 1. Perotrochus midas feed-
ing on Strongylophora hartmani. JSL-II dive 1501, #2, west end of Island at Ciouldings Cay, New Providence Island, Bahamas,
25°00'00"N, 77°34'06"\V. in 766 m. October 20, 1987. Specimen not collected Note area of sponge consumed by snail. 2. Perotrochus
midas and Strongylophora hartmani along steep wall, JSL-II dive 1505, Chub Cay, Berry Islands, Bahamas, 25°22'22"N, 77°50'25"W,
in 777 m. October 23, 1987. Specimen not collected. Note layer of planktonic sediment.
M. G. Harasewvch et al, 1988
Page 93
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THE NAUTILUS, Vol. 102, No. 3
specimens were collected, one together with its prey,
using the hydraulic arm and clamshell scoop. These spec-
imens were fixed in formalin and preserved in 70% eth-
anol.
Fifty-six specimens of Perotrochus arnabilis (Bayer,
1963) were observed and collected during four dives
aboard the research submersible Nekton Delta on the
"Charleston Lumps", an area of rough bottom topog-
raphy approximately 90 miles east of Charleston, South
Carolina (32°43'78"-32°44'90"N, 78°05'68"-78°06'00"\V)
(Askew, 1988:91), at depths ranging from 200 to 230 m.
Samples were collected by suction through a 2 inch (51
mm) diameter hose. Specimens were frozen on dry ice
and maintained at — 80°C.
After identification of the intact sponge on which a
specimen of P. midas was feeding, a spicule sample for
SEM examination was prepared by digesting a portion
of the sponge in 70% nitric acid (HNO5) until only sili-
ceous material remained. Two specimens of P. midas
and three specimens of P. amalnlis were dissected, and
sections of the esophagus between the esophageal valve,
situated behind the buccal mass (Fretter, 1964:181, fig.
5, ov; 1966:609, fig. 2, v), and the long sphincter at the
opening of the stomach (Fretter, 1966:609) were excised.
Also removed were portions of the rectum (Fretter, 1966:
fig. 1, r; 1964: fig. 2, r). These sections were teased apart
and examined under a dissecting microscope, then treat-
ed with warm diluted bleach (1-2% sodium hypochlorite,
NaOCl) to dissolve organic material while leaving the
siliceous and calcified remains. The preparations were
rinsed in distilled water, filtered through 0.4 ^m Nucle-
pore membrane filters, and the filters mounted directly
onto SEM stubs. Samples were coated with carbon and
gold, and photographed using a Hitachi S-570 scanning
electron microscope. A transverse section of the intact
sponge was critical point dried prior to SEM examination.
The following voucher material is deposited at the
National Museum of Natural History:
Perotrochus midas: Specimen 2. USNM 857097, JSL-II
dive 1501, #2, West end of Island at Gouldings Cay,
New Providence Island, Bahamas, 25°00'00"N,
77°34'06"W, in 766 m. October 20, 1987.
Perotrochus amabilis: Specimens 1, 2, and 3. USNM
846900, DELTA Dive 560, 129 km due east of Charles-
ton, South Carolina, USA, 32°43'95"N, 78°05'72"W, in
198-210 m. May 3, 1987.
RESULTS
Of the 16 sightings of Perotrochus midas. this species
was observed in close proximity to, or actively feeding
upon, the sponge Strongylophora hartmani Van Soest,
1980 (class Demospongiae, order Haplosclerida) on six
occasions (figures 1, 2). Two specimens of P. midas were
collected, one (specimen 1) together with the sponge on
which it was feeding. The esophagus of both snails was
distended and full of sponge tissue. Comparisons of spic-
ules taken from the esophagus of both specimens of Per-
otrochus midas (specimen 1, figure 5) with those of Stron-
gylophora hartmani (figures 3, 4) confirmed the identity
of the prey species. The rectum of specimen 1 contained
spicules of S. hartmani (figures 6, 7) as well as spicules
tentativeK attributed to a species of PachastreUa (figure
7). The rectal contents of specimen 2 included spicules
of S. hartmani, but spicules of an unidentified sponge
belonging to the order Hadromerida (figure 8) comprised
an estimated 80% of the \olume. The contents of the
esophagus and rectum from both specimens of P. midas
consisted almost entirely (estimated >95% by volume)
of sponge spicules, with foraminiferal and ostracode tests
accounting for most of the other identifiable remains.
Of the 53 specimens of Perotrochus amabilis observed
and collected, the majority were on phosphorite blocks
ranging in size from several centimeters to over a meter
in length. Only three specimens were on or near a sponge.
In all three instances, the sponge, which was small and
roughly spherical (<6 cm diameter), could not be col-
lected. Examination of the contents of the esophagus and
rectum of these three specimens revealed a more het-
erogeneous assemblage of sponge spicules (about 80%),
diatoms, and foraminiferal tests (about 20%) (figures 9,
10). In no instance could more than an estimated 50%
of the contents of either the esophagus or the rectum be
attributed to a single species of sponge.
DISCUSSION
Direct feeding observations as well as gut content anal-
yses indicate that Perotrochus midas feeds on the sponge
Strongylophora hartmani b> rasping large, deep depres-
sions (>2 cm diameter, >1 cm depth) in its surface
(figure 1). In each of the two specimens dissected, the
voluminous esophagus was distended, and contained a
corresponding volume (2-3 cm') of sponge tissue. Rectal
contents also consisted almost exclusively of sponge spic-
ules, though from species belonging to the orders Cho-
ristida and Hadromerida. The most abundant groups of
deep water sponges (in terms of biomass) in the tropical
western Atlantic and Caribbean are the orders Haplo-
sclerida and Choristida, with the most common species
often being highly silicified (Pomponi, unpublished ob-
servations). Although six of the 16 sightings of P. midas
were on or near a haplosclerid sponge, it is unclear
Figures 3, 4. Strongylophora fiartmani Van Soest, 1981. 3. Transverse section, outer surface at top (SEM), scale bar = 300 ^lm.
4. Nitric acid spicule preparation (SEM), scale bar = 200 urn. Figure 5. Perotrochus midas, contents of mid-esophagus of spetimen
1, showing high concentration of Strongylophora hartmani spicules (SEM), scale bar = 200 ^m. Figures 6, 7. Perotrochus midas,
rectal contents of specimen 1. 6. Strongylophora hartmani spicules (SEM), scale bar = 200 ^m. 7. Spicules of S. hartmani as well
as of ?Pachastrella sp. (arrows) (SEM), scale bar = 200 ^m. Figure 8. Perotrochus midas, rectal contents of specimen 2, with
spicules of unidentified sponge, order Hadromerida (arrows) (SEM), scale bar = 200 ^m.
M. G. Harasewych et al, 1988
Page 95
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THE NAUTILUS, Vol. 102, No. 3
Figures 9, 10. Perotrochus amabilis (Bayer, 1963), contents of mid-esophagus of specimen collected 90 miles due east of Charleston,
SC, in 210 m. 9. Spicule of an unidentified species, order Poecilosclerida (SEM), scale bar = 20 ^m. 10. Spicules of Strongylophora
sp. (small arrow) and an unidentified species, order Choristida or Spirophorida (large arrows) (SEM), scale bar = 100 ^m.
whether prey species are selected, or mereK reflect the
relative abundance of sponges in a nutrient-poor envi-
ronment. In either case, available evidence indicates that
sponges comprise the major component of the diet of P.
midas, and that the small amounts (<5%) of foraminif-
eral and ostracode tests as well as other planktonic sed-
iment found in the esophagus and rectum of this species
were present on the surface of sponges (figures 1, 2), and
probably do not contribute significantly to its nutrition.
Although far more specimens of Perotrochus amabilis
were observed and collected, fewer were in the proximity
of a sponge. Due to the smaller size of this species and
the inability to collect the sponges, no direct observations
of feeding can be documented. E.xamination of the con-
tents of the esophagus and rectum of three specimens
indicate that sponge spicules comprise approximately
80% of the bleach insoluble mass, the remaining fraction
consisting mostly of foraminiferal tests and diatom frus-
tules. Fretter (1964:182) reported similar contents in the
stomach of this species and concluded that the animal
was a microphagous scavenger. We suggest that P. ama-
bilis, like P. midas, feed exclusively on sponges, but, due
to their smaller size and correspondingly shorter snout,
these animals are not able to penetrate as deeply into the
tissues of the sponge, and feed on surface tissues covered
with deposits of planktonic sediment, thus accounting
for the higher proportion of diatoms and foraminifera
in the gut.
With a single exception (Barnard, 1963), all published
reports on the gut contents of pleurotomariids (table 1)
list sponge spicules as a major component. Woodward
(1901:252) was the first to speculate that the distinctive
radula, shared by all pleurotomariids, was adapted for
spongivory, with the hooked teeth "tearing away great
pieces of the sponge , and the brush teeth used to "rasp
away some of the flesh from the spicules". The occur-
rence of brush or filament-tipped teeth, long considered
unique to Pleurotomariidae, in unrelated, sponge-feed-
ing mesogastropods of the genus Seila. has led Hickman
(1984a:35) to conclude that this tooth morpholog\ is a
functional adaptation to sponge predation and not a phy-
logenetically constrained, conservative, morphological
feature.
Although exceptions have been documented (e.g., Gra-
ham, 1939; Perron, 1975), archeogastropods are gener-
all\ considered to be herbivores (Yonge & Thompson.
1976). This has led several authors (Yonge, 1973; Hick-
man, 1984a) to imply that carnivory was not the original
mode of feeding of pleurotomariids. Yonge (1973) fur-
ther suggested that the change to a carnivorous diet may
have been associated with the ecological shift of this
family from shallow water reefs, which it inhabited dur-
ing the Paleozoic and Mesozoic, to deeper water (>200
m), hard substrates by the end of the Eocene.
As members of all three Recent pleurotomariid genera
have radulae with filament-tipped teeth, it would appear
that spongivory in Pleurotomariidae predates the diver-
gence of the genus Entemnotrochus Fischer, 1885, char-
acterized by a deep, broad umbilicus and an anal slit
extending nearK 180° back from the aperture, from the
genera Perotrochus Fischer, 1885, and Mikadotrochus
Lindholm, 1927, both with shallow (<90°) anal slits and
without umbilici. Inclusion of the genus Conotomaria
Cox, 1959, which is also characterized by having shells
w ith deep anal slits and umbilici, in the same clade as
Entemnotrochus would date this divergence, and there-
fore the adaptation to spongivory, prior to the Late Ju-
rassic (Knight et al., 1960). The restriction of the family
M. G. Harasewych et ai, 1988
Page 97
Table 1. Published reports on the contents of the alimentary system and fecal pellets of pleurotomariid gastropods.
Species Food Reference
Mikadotrochxis heyrichi
Mikadotrochus hirasei
Perotrochus africanus
Perotrochus amahilis
Perotrochus amahilis
Perotrochus midas
Sponge spicules, order Poecilosclerida
Sponge spicules, orders Haplosclerida,
Poecilosclerida, and Hadromerida
Amorphous mass with few tiny foraminiferans
Sponge spicules, foraminiferans, diatoms, and
algal fragments
Sponge spicules, foraminiferans, and diatoms
Sponge spicules, foraminiferans, and diatoms
Woodward, 1901:252
Arakawa ei ai. 1978
(table 1)
Barnard, 1963:156
Fretter, 1964:182
Present study
Present study
to bathyal depths at the time of formation of the psy-
chrosphere, the lower, cooler layer of a two-layer ocean
(Bruun, 1957; Benson, 1975) was accompanied by a con-
siderable reduction in diversity, compared to Paleozoic
and Mesozoic fauna (Woodward, 1885).
Many sponges contain secondary metabolites that are
toxic (Green, 1977; Bakus & Thun, 1979). Nevertheless,
mollusks, echinoderms, fishes, and marine turtles are nat-
ural predators of sponges, with nudibranchs being among
the most species-specific spongivores (Sara & Vacelet,
1973). Halichondria okadai (Kadota, 1922), the preferred
pre\ of several species of nudibranchs, for e.xample, con-
tains potent cytotoxic, antifungal, and tumor promoting
compounds (Tachibana et ai, 1981; Fujiki et ai, 1987).
Strongylophora hartnmni, the sponge species eaten by
Perotrochus midas, contains puupehenone, a cytotoxic
compound (Komoto et al.. 1987). It is possible that in-
corporation of diet-derived toxic metabolites may confer
some protection from predators upon pleurotomariids,
but this hypothesis remains to be tested.
ACKNOWLEDGEMENTS
We thank Richard Cooper and Peter Auster of the
NOAA National Undersea Research Program, University
of Connecticut, Avery Point, for making submersible
time available at the Charleston Lumps site. The assis-
tance of M. Cristina Diaz with the identification of the
intact sponge, and Susann Braden with the scanning elec-
tron microscopy is gratefully acknowledged. This is Har-
bor Branch Oceanographic Institution Contribution No.
643.
LITERATURE CITED
Arakawa, K. Y., D. Nakano, O. Tsukuba, and T. Hoshino. 1978.
On faecal pellets and food habits of emperor's slit shell,
Mikadotrochus hirasei (Pilsbry). Venus 37(3):116-120.
Askew, T. M. 1988. A new species of pleurotomariid gastro-
pod from the western Atlantic. The Nautilus 102(3):89-
92.
Bakus, G. J. and M. Thun. 1979. Bioassays on the toxicity of
Caribbean sponges. In: C. Levi and \. Esnault {eds. ).
CoUoques Internationaux du C.N.R.S., No. 291, Biologie
des Spongiaires, C.N.R.S., Paris, p. 417-422.
Barnard, K. H. 1963. Notes on the animals Cyrina gigantea
(Lam.) and Pleurotomaria africana Tomlin. Proceedings
of the Malacological Society of London 35:155-158.
Bayer, F. M. 1963. A new pleurotomariid gastropod trawled
in the Straits of Florida by R/V Gerda. Bulletin of Marine
Science of the Gulf and Caribbean 13(3):488-492.
Bayer, F. M. 1966. New pleurotomariid gastropods from the
western Atlantic, with a summary of the Recent species.
Bulletin of Marine Science 15(4):737-796.
Benson, R. H. 1975. The origin of the psychrosphere as re-
corded in changes of deep-sea ostracode assemblages. Le-
thaia 8:69-83.
Bouvier, E. L. and H. Fischer, 1899. Reports on the results
of dredging ... in the Gulf of Mexico and the Caribbean
Sea, and on the east coast of the United States, 1877-1880,
by the U.S. Coast Survey Steamer "Blake," . . . XXXVIII.
Etude monographique des Pleurotomaires actuels. Bulletin
of the Museum of Comparative Zoology, Harvard 32:193-
249. Also Published in Journal de Conchyliologie 47(2):
77-151, pis. 4-7.
Bouvier, E. L. and H. Fischer. 1902. L organisation et les
affinites des gasteropodes primitifs d'apres I'etude anato-
mique du Pleurotomaria heyrichi. Journal de Conchy-
liologie 50(2): 117-272, pis. 2-6.
Bruun, A. F. 1957. Deep sea and abyssal depths. In:
Hedgepeth, J. W. (ed). Treatise on marine ecology and
paleontology. Geological Society of .\merica. Memoirs 67:
641-671.
Cox, L. R. 1959. Thoughts on the classification of the Gas-
tropoda. Proceedings of the Malacological Societv of Lon-
don 33:190-202.
Fischer, P. 1885. Manuel de conchyliologie et de paleonto-
logie conchyliologique. Histoire naturelle des moUusques
vivants et fossiles. Fascicule 9:785-896. Libraire F. Savy,
Paris, xxiv -I- 1369 p., 23 pis. (1887).
Fretter, V. 1964. Observations on the anatomy of Mikado-
trochus amahilis Bayer. Bulletin of Marine Science of the
Gulf and Caribbean 14(1):172-184.
Fretter, V. 1966. Biological investigations of the deep sea. 16.
Observations on the anatomy of Perotrochus. Bulletin of
Marine Science 16(3):603-6i4.
Fretter, V. and A. Graham. 1976. The prosobranch molluscs
of Britain and Denmark. Part 1 — Pleurotomariacea, Fis-
surellacea and Patellacea. The Journal of Molluscan Stud-
ies Supplement l:iv -I- 37 p.
Fujiki, H., M. Suganuma, H. Suguri, S. Yoshizawa, M. Ojika,
K. Wakamatsu, K. Yamada, and T. Sugimura. 1987. In-
duction of ornithine decarboxylase activit)- in mouse skin
by a possible tumor promoter, okadaic acid. Proceedings
of the Japanese Academy, Series B, 63(2):51-53.
Graham, A. 1939. On the structure and function of the ali-
Page 98
THE NAUTILUS, Vol. 102, No. 3
mentary canal of the style-bearing prosobranchs. Pro-
ceedings of the Zoological Society of London 109:75-112.
Green, G. 1977. Ecologvof toxicity in marine sponges. Marine
Biology 40:207-215.'
Herbert, D. G. 1987. Revision of the Solariellinae (Mollusca:
Prosobranchia: Trochidae) in southern Africa. Annals of
the Natal Museum 28t2):283-382.
Hickman, C. S. 198-4a. Form and function of the radulae of
pleurotomariid gastropods. The X'eliger 27(l):29-36.
Hickman, C. S. 1984b. Pleurotomaria. pedigreed persever-
ance? In: Eldridge, N. and S. M. Stanley (eds. ). Living
fossils. Springer-V'erlag, New York, NY, p. 225-231.
H\nian, L. H. 1967. Mollusca I. The invertebrates. Vol. 6.
McGraw-Hill Book Co., New York, NY, vii -I- 792 p.
Knight, J B., L. R Cox, A. M. Keen, R. L. Batten. E. L.
Yochelson, and R. Robertson. 1960. Systematic descrip-
tions [Archeogastropoda] In: R. C. Moore (ed.). Treatise
on invertebrate paleontology, Part I, Mollusca 1. Geolog-
ical Society of .America and University of Kansas Press,
Lawrence,' KS, p. n60-I331.
Komoto, S., O. McConnell, .\. Wright, F. Koehn, W. Thomp.son,
M Lui, and K. Snader. 1987. Puupehenone, a cytotoxic
metabolite from a deep water marine sponge. Journal of
Natural Products 50:336.
Leighton, D L 1961. Observations of the effect of diet on
shell coloration in the red abalone, Haliotis rufescens
Svvainson. Veliger 4:29-32, pi. 6.
Leighton, D. L. and R. A. Boolootian. 1963. Diet and growth
in the black abalone, Haliotis cracherodii. Ecology 44(2):
227-238.
Lindholm, VV . .\. 1927. O Pleuro!.oniariabeyrichilii\geudor{
(Gastropoda) v kolleklsiakh zoologicheskogo muzeia AN 5
zametkoi o rode Pleurotomaria s. lat. Dokladv Akademii
Nauk USSR 24:409-414.
Matsumoto, Y., T. Kataoka, and M. Sekido. 1972. On the
behavior of Beyrich's slit shell, Mikadotrochus beyrichi
(Hilgendorf) kept in the aquarium. Venus 30(4): 147-152.
Perron, F. 1975. Carnivorous Calliostoma (Prosobranchia:
Trochidae) from the northeastern Pacific. The Veliger 18(1):
52-54, fig. 1.
Sara, M. and J. V'acelet. 1973. Ecologie des Desmosponges.
In: P.-P. Grasse (ed.), Traite de zoologie. Tome III. Spon-
giaires. Masson et Cie, Paris, p. 462-576.
Sekido, M., O. Tsukada, Y. Matsumoto. and T. Kataoka. 1976.
On the keeping of Teremachi's slit shell, Perotrochus te-
remachii. Journal of the Japanese .Association of Zoological
Gardens and .Aquariums 18(2):49-52.
Shepherd, S \. 1973. Studies on southern .Australian abalone
(genus Haliotis) I. Ecology of five sympatric species. .Aus-
tralian Journal of Marine and Freshwater Research 24:
217-257.
Soest, R. W. M. Van 1980. Marine sponges from Curacao
and other Caribbean localities Part II Haplosclerida.
Studies of the Fauna of Curacao and the Caribbean Islands
62(191):1-173.
Tachibana, K., P. J. Scheuer, Y. Tsukitani, H. Kikuchi, D. Van
Engen, J. Clardy, Y. Gopichand, and F. J. Schmitz. 1981.
Okadoic acid, a cytotoxic polyether from two marine
sponges of the genus Halichondria. Journal of the Amer-
ican Chemical Society 103:2469-2471
Thiele, J. 1935. Handbuch der Systematischen VVeichtier-
kunde. Part 4:1129.
Woodward, H. 1885. On Recent and fossil Pleurotomariae.
Geological Magazine, Decade 3, 2(101:433-439, pi. 11.
Woodward, M. F. 1901. The anatomy of Pleurotomaria bey-
richii, Hilg The Quarterly Journal of Microscopical Sci-
ence 44(2):215-268, pis. 13-16.
Yonge, C M. 1973. Observation of the pleurotomarid En-
temnotrochus adansoniana in its natural habitat Nature
241(5384):66-68.
Yonge, C. M. and T. E. Thompson 1976. Living marine
mollusks. William Collins Sons & Co. Ltd., London, 288
p., 16 pis.
THE NAUTILUS 102(3):99-101, 1988
Page 99
A New Species of Macrarene (Turbinidae: Liotiinae)
from Brazil
James H. McLean
Los Angeles County Museum of
Natural Histor)-
900 Exposition Boulevard
Los Angeles, California 90007, USA
Ricardo Silva Absalao
Renato Luiz dos Santos Cruz
Departamento de Zoologia,
Instituto de Biologia, CCS.
Universidade Federal do
Rio de Janeiro
Ilha do Fundao, Rio de Janeiro,
Brasil 21940
ABSTRACT
The new species Macrarene digitate from the northeast Bra-
zilian coast represents the first record of this principally eastern
Pacific genus in the western Atlantic. The species had previ-
ously been known from juvenile specimens reported as Liotia
admirabilis E. A. Smith, the holotype of which is not a member
of the subfamily Liotiinae.
INTRODUCTION
The species here described was first recognized as a mem-
ber of the Brazilian fauna by Rios (1975, 1985), who
referred it to Liotia admirabilis E. A. Smith, 1890. That
species was described from the oceanic island of Saint
Helena. According to the original description. Smith's
species has a maximum dimension of I'A mm. The 15
syntypes were examined at the British Museum (Natural
History) (catalogue numbers 1889.10.1.1554-68) by the
senior author in 1984 and found to be similar (although
not clearly referable) to the skeneiform genus Parviturbo
Pilsbry & McGinty, 1945, which is not a member of the
Liotiinae.
The Brazilian species was clearly undescribed, but its
true generic affinity was not readily apparent because
the specimens available to Rios, which have been ex-
amined by McLean (figure 2), were not mature and the
expression of the mature lip was impossible to determine.
More recently, two larger specimens have come to light
and it can now be maintained that the species has the
characters of the genus Macrarene Hertlein & Strong,
1951.
Abbreviations for institutions are as follows: LACM,
Los Angeles County Museum of Natural History; MORG,
Museu Oceanografico, Universidade do Rio Grande, R.S.,
IBUFRJ, Instituto de Biologia, Universidade Federal do
Rio de Janeiro.
SYSTEMATICS
Family Turbinidae Rafinesque, 1815
Subfamily Liotiinae H. & A. Adams, 1854
Shells of the subfamily are characterized by turbiniform
to discoidal profiles, nacreous interiors, fine lamellar
sculpture, intritacalx (calcified periostracum) in most
genera, circular apertures, and multispiral opercula with
calcareous beads. Radula like that of members of other
turbinid subfamilies.
Although previously treated by most authors as a full
family, the Liotiinae have recently been ranked as a
subfamily of Turbinidae by McLean (1987).
Genus Macrarene Hertlein & Strong, 1951
Type species (original designation): Liotia californica
Dall, 1908. Recent, off Baja California, Mexico.
Macrarene species are characterized by turbinate white
shells, broad umbilici, and presence of axial ribs and
spiral cords that form spines at their intersections. Spac-
ing of the axial ribs increases in the final whorl. In some
species the ribs then become more closely spaced in the
final quarter whorl. The final lip is not thickened at
maturity.
Some Macrarene species reach relatively large sizes.
The genus differs from Arene in lacking shell pigmen-
tation and in having the spacing of the axial sculpture
increasingly separated as the shell matures. The white-
shelled genus Liotia is smaller and retains tight spacing
of the axial sculpture.
The white-shelled, new world Liotiine genera Mac-
rarene, Liotia, and Cyclostrema differ as a group from
those of the Indo-Pacific and Australasian regions in lack-
ing the thickened mature lips that characterize the gen-
era Bathtjiiotina Habe, 1961, Liotina Fischer, 1885,
Dentarene Iredale, 1929, and Austroliotia Cotton, 1948.
Page 100
THE NAUTILUS, Vol. 102, No. 3
Figures 1, 2. Macrarene digitata new species. 1. Holotype, IBUFRJ 1562. x 7.1. 2. Immature specimen, MORG 18.359. x 10.0.
For further remarks on the Indo-Pacific group see McLean
(1988).
There are si.\ previously described species of Macra-
rene: M. californica (Dall, 1908), M. cookeana (Dall,
1908), M. diegensis McLean, 1964 (Pliocene); M. far-
allonensis (A. G. Smith, 1952), M. lepidotera McLean,
1970, and M. spectahilospina Shasky, 1970. All occur
offshore in the tropical to temperate eastern Pacific.
Macrarene digitata new species
(figures 1, 2)
Description: Shell small for genus, turbinate, white,
interior weakly nacreous, ma.ximum diameter 6.7 mm,
whorls 3.5, aperture only slightK ohliciue, final lip not
thickened. Whorls circular in outline; final w horl in con-
tact with previous whorl at tips of axial ribs. Shell surface
marked by microscopic lamellar growth increments; in-
tritacalx present. Protoconch diameter about 200 ^m.
Suture deeply impressetl, first and second whorls rising
above protoconch, third whorl descending, resulting in
flat-topped profile for earlv whorls. F"irst teleoconch whorl
nearly smooth (except for fine lamellar sculpture), second
with about 20 strong axial ribs and nine, nearly equal
spiral cords defining deep, rectangular pits. Inter.sections
of axial and spiral sculpture produce sharpK- projecting
spines that are slightly upturned adapicalK . Spiral cords
of final whorl increasing to 12; axial ribs decreasing to
15. Spines produced on final whorl by interaction of spiral
and axial sculpture; spines with w eblike interconnections.
Axial ribs narrow across umbilical wall, forming single
descending row of sharp-pointed projections along in-
nermost spiral cord. Operculum and radula unknown.
Type locality: Off northeast coast of Brazil (03°59'N,
49°35'W), 100 m, Brazilian Naval Research Vessel Al-
mirante Saldaiilia. station 1913, May 6, 1968, 1 speci-
men.
Type material: Holotype (figure 1), IBUFRJ 1562.
Height 5.0 mm, diameter 6.7 mm. The holot\pe is a
dead collected specimen in good condition. Parat\pe,
LACM 2377, off Cabo San Roque, Rio Grande do Norte,
Brazil (04°30'S, 50°03'\\'), 146 m, Brazilian Naval Re-
search Vessel Almirante Saldanha. station 1921, May 8,
1968 (height 4.3 mm, diameter 6.6 mm). The paratope
agrees with the holot)pe in size and sculptural details
but is in subfossil condition with attached sedimentary
deposits on the base.
Referred material: MORG 18.359, 2 immature speci-
mens [height 2.7, diameter 4.3 mm (figure 2); height 1.5,
diameter 2.6] from Paripueira, Alagoas, Brazil, in beach
drift collected b\ P. S. Cardoso, December, 1964; MORG
20.620, 2 immature specimens (height 2.1, diameter 3.7
mm; height 2.4, diameter 3.6 mm), Fernando de No-
ronha Island, 6 m, collected b\ L. Barcellos, January,
1979.
Etymology: From the Latin digitatus. having fingers.
J. H. McLean et at., 1988
Page 101
Remarks: Macrarene digitata is the smallest species of
the genus described to date. However, it is not certain
that the holot\pe is mature. Although this genus does
not form a thickened lip, maturit) in other members of
the genus is indicated by closer spacing of the axial ele-
ments in the final quarter whorl of growth, as indicated
in the original illustration of M. spectabilospina of Shasky
(1970: fig. 2). The absence of such closer spacing of the
axial element suggests that a quarter whorl of additional
growth (and a substantial increase in diameter) is possible
for M. digitata.
Macrarene digitata is unique in the genus in having
all elements of the spiral sculpture of similar strength,
rather than having a strongly projecting peripheral
carination. Such a sculptural distinction is not regarded
as a generic level character because generic characters
in the Liotiinae are more reliably based on apertural
morpholog\', particularly the structure of the final lip.
The most characteristic feature of this species is the
fingerlike aspect of the projecting spines. It cannot easily
be confused with any other member of the Liotiinae.
ACKNOWLEDGEMENTS
We are grateful to Prof. E. C. Rios for the loan of the
referred material of this species.
LITERATURE CITED
Dall, W. H. 1908. Reports on the dredging operations off the
west coast of Central America to the Galapagos, to the
west coast of Mexico, and in the Gulf of California . . .
,\1V. The Mollusca and Brachiopoda Bulletin of the Mu-
seum of Comparative Zoology, Harvard University 43:
205-487, pis. 1-22.
Hertlein, L. G. and A. M. Strong. 1951. Eastern Pacific ex-
peditions of the New York Zoological Society, XLIII. Mol-
lusks from the west coast ol Mexico and Central America.
Zoologica, Scientific Contributions of the New York Zoo-
logical Society 36:67-120, pis. 1-11.
McLean, J. H. 1964. New species of Recent and fossil west
American aspidobranch gastropods. The Veliger 7:129-
133.
McLean, J. H. 1970. New species of tropical eastern Pacific
Gastropoda. Malacological Review 2:115-130.
McLean, J. H. 1987. Angariinae and Liotiinae — the primitive
living trochaceans. .Annual Report of the Western Society
of Malacologists 19:16.
McLean, J. H. 1988. Two new species of Liotiinae (Gastropo-
da: Turbinidae) from the Philippine Islands. The Veliger
30:408-411.
Rios, E. C. 1975. Brazilian marine mollusks iconography.
Museu Oceanografico, Fundagao Universidade do Rio
Grande, 331 p., 91 pis.
Rios, E. C. 1985. Seashells of Brazil. Museu Oceanografico,
Funda^ao Universidade do Rio Grande, 328 p., 102 pis.
Shasky, D. R. 1970. New gastropod taxa from tropical western
America. The Veliger 13:188-195.
Smith, A. G. 1952. Shells from the bird guano of southeast
Farallon Island, California, with description of a new species
of Liotia. Proceedings of the California Academy of Sci-
ences, Series 4, 27:383-387.
Smith, E. A. 1890. Report on the marine Mollusca of the
island of Sta. Helena. Proceedings of the Zoological Society
of London 1890:247-322, pis. 21-24.
THE NAUTILUS 102(3): 102-105, 1988
Page 102
Nerita fortidentata, a New Gastropod from the Neogene of
Panama, with Comments on the Fossil Record of
Nerita in Tropical America
Geerat J. Vermeij
Department of Zoology
University of Maryland
College Park, MD 20742, USA
Timothy M. Collins
Department of Geology and
Geophysics
Yale University
New Haven, CT 06520, USA
ABSTRACT
Nerita fortidentata new species is described from the Neogene
of Bocas del Toro, Panama It is most closely related to the
Recent Caribbean \. fulgurans Gmelin, 1791. The fossil record
and biogeography of tropical American Nerita are reviewed.
At least two lineages of Nerita present in tropical America
during the Tertiary have become restricted in the Recent fauna
tn the Infln-Wf^t-Pacific region
INTRODUCTION
Neritid gastropods are prominent members of tropical
rocky intertidai communities. Because ihe fossilization
potential of tfiese animals is poor, little is known about
tlie historical development of the genus Serita Linnaeus,
1758. It was, therefore, of considerable interest to find
an excellently preserved specimen of an apparently hith-
erto unrecognized species from the Neogene of tropical
America. Here we describe the new species and review
briefly some biogeographically interesting aspects of the
history of the genus Nerita in tropical America.
METHODS
Species of Nerita are distinguished conchologically by
characters of shape, apertural dentition, external sculp-
ture, form and sculpture of the parietal callus, and the
shape and sculpture of the calcareous operculum. Two
ratios are especially helpful in describing the overall shell
form of Nerita (X'ermeij, 1973). The first is globosity, G,
defined as the distance H, between the dorsal surface of
the body whorl and the center of the parietal callus
divided by the geometric mean betw een the shell's major
diameter Dj and minor diameter D2: G = H,/(D,D2)'''.
The second ratio is the degree of basal excavation. The
plane of the parietal callus typically lies at an angle to
the horizontal w hen the shell lies aperture-down on a
flat surface. The greater the angle, the greater is the
degree of basal excavation. An approximation of the de-
gree of basal excavation is given by the ratio E = H2/
H,, where Ho is the distance from the horizontal plane
on w hich the shell rests to the dorsal surface of the body
whorl.
SYSTEMATIC DESCRIPTION
Class Gastropoda
Subclass Prosobranchia
Order Neritacea
Family Neritidae
Genus Nerita Linnaeus, 1758
Type species: Nerita peloronta 1758, Recent, tropical
Western Atlantic.
Subgenus Theliostyla Morch, 1852
Type species: Nerita albicilla Linnaeus, 1758, Recent,
Indo- West-Pacific.
Nerita (Theliostyla) fortidentata new species
(figures 1, 2)
Diagnosis: Shell thick, moderately globose (G = 0.59),
base little excavated (E = 1.10), apex of spire barely
raised above rest of shell. Outer lip very thick, inner edge
with 12 strong teeth; two teeth nearest spire \er\ large
and protruding, as is third tooth from abapical end of
lip; columellar lip with tw^o strong centrally placed teeth;
adapical portion of parietal region with a fold of about
the same size and strength as adapical tooth, w hich curves
into aperture; sculpture consisting of 21 regularK -spaced
flat-topped smooth spiral cords, w hich Dare slightly from
base to barely overhang incised interspaces about one-
third the width of ribs; parietal callus small, its surface
sculptured with about 10 strong ridges that bear up to
3 large granules each; holotype shows faint radial color
pattern of alternating continuous and discontinuous pro-
socline bands of off white and grey-black; operculum
unknown.
G. J. Vermeij and T. M. Collins, 1988
Page 103
Figure 1. Nerita fortidentata new species, from Bocas del
Toro, Panama, Holotype, USNM 423644, height 19.7 mm,
Apertural view, showing thickened outer lip and enlarged teeth
on adapical portion of outer lip.
Holotype: United States National Museum number
423644. Major diameter 21.3 mm, minor diameter 16.9
mm, Hi 11.2 mm, H^ 12.3 mm, standard shell height
19.7 mm (apex abraded), standard shell diameter 19.9
mm, shell thickness at midpoint of outer lip 3.5 mm.
Type locality: Panama, Province of Bocas del Toro,
Archipelago of Bocas del Toro, Punta Robalo quadrangle.
Island of Cayo Agua, eastern side about 400 meters south
of Punta Nispero on the shoreline in clayey, tuffaceous,
quartzose, blue-grey siltstones with dense shelly horizons.
We have followed Woodring (1982) in referring to the
Late Miocene-Pliocene deposits of the Bocas del Toro
area as the Limones Formation. The true relationships
between the Miocene-Pliocene of Bocas del Toro with
respect to the Costan Rican Limones Formation and the
Gatun Formation of the Canal Zone have not been elu-
cidated. Laurel B>bell of the U.S.G.S. (personal com-
munication) has assigned a preliminary age of Late Mio-
cene to Early Pliocene to the locality from which the
holotype of N. fortidentata was collected. This age de-
termination is based on the presence of the calcareous
nannofossils Sphenolithiis abies Deflandre, in Deflandre
and Fert, 1954 (last occurrence middle Pliocene), and
Discoaster brouweri Tan, 1927 (first occurrence middle
Miocene). Thomas M. Cronin, U.S.G.S. (personal com-
munication), examined the ostracode faunas from several
adjacent localities of the same formation on Cayo Agua.
He noted a remarkable similarity between the ostracodes
from these samples and the ostracode fauna described
by van den Bold (1967) from the type Gatun Formation
of the Canal Zone. On the basis of these similarities he
suggests a preliminary age of Late Miocene for this fau-
na. Harry Dowsett, also of the U.S.G.S. (personal com-
munication), examined the planktic foraminifera from
an adjacent localit\ of the same formation on Cayo Agua
and found an assemblage indicative of planktic zone
N17-NT8 (Late Miocene-Early Pliocene). The consensus
at this stage, therefore, is that the beds from which N.
fortidentata was collected are Late Miocene to Early
Pliocene in age.
Figure 2. Nerita fortidentata new species, from Bocas del
Toro, Panama. Holotype, UNSM 423644, height 19.7 mm. Ab-
apertural view.
Remarks: The new species clearly belongs to the sub-
genus Theliostyla Morch, 1852 (type N. albicilla Lin-
naeus, 1758), which is characterized by granulate sculp-
ture on the parietal region, a barely protruding spire,
and w ell-developed external spiral sculpture. Among the
four Recent species of this subgenus in tropical America,
N.fulgurans Gmelin, 1791, bears the closest resemblance
to A', fortidentata. Measurements of 17 specimens of N.
fulgurans in the Vermeij collection from the Atlantic
coasts of Panama, Costa Rica, Venezuela, and Jamaica
show that this species is less globose (G = 0.54 ± 0.020,
range 0.50-0.57) and basally much more excavated
(E = 1.27 ± 0.05, range 1.21-1.39) than is the new
species. Nerita funiculata Menke, 1851, the Recent east-
ern Pacific cognate of N. fulgurans, is also less globose
(G = 0.51 ± 0.04, range 0.42-0.58, based on 15 speci-
mens in the Vermeij collection from Costa Rica, Panama,
and Ecuador) and more excavated (E = 1.28 ± 0.15,
range 1.13-1.60) than is N. fortidentata. Both N. ful-
gurans and N. funiculata have weaker and more nu-
merous denticles on the outer lip, weaker and more finely
granulated ridges on the parietal region, and spiral cords
that are more numerous and more variable in size on the
body whorl (18-35 in N. fulgurans, usually more than
30 in N. funiculata). The spiral cords of N. fulgurans
show a tendency to bifurcate on the body whorl, whereas
no such tendencN is seen in iV. fortidentata. The three
subspecies of N. ascensionensis Gmelin, 1791, from is-
lands in the tropical south Atlantic (Vermeij, 1970) differ
from iV. fortidentata by having a nearly smooth parietal
region and by the very weak dentition on the outer lip.
The West Indian A', tessellata Gmelin, 1791. differs from
A', fortidentata b\' having low rounded spiral cords bro-
ken irregularl) bv high and low areas correlating to the
characteristic black and white checkered pattern found
in this species, a finely granulated parietal region of
relatively large extent, and a weakK denticulated outer
lip (Russell, 1941).
Jung (1965) recorded Serita fulgurans from the Mid-
dle Miocene and Upper Pliocene of Venezuela, but he
pointed out that his specimens differed from Recent shells
by having stronger apertural dentition and a less concave
Page 104
THE NAUTILUS, Vol. 102, No. 3
(that is, less excavated) parietal region. Later Jung (1969)
found a similar shell in the Late Miocene Melajo Clay
Member of the Springvale Formation of Trinidad. Like
the Venezuelan material, the shell from Trinidad has
only 16 ribs, but instead of having 2 centrali\ placed
columcllar teeth, as in the Venezuelan material, the Trin-
idad specimen has one upper tooth which curves into
the aperture (this upper tooth is probably equivalent to
the parietal fold found on the tvpe of \. fortidentata]
and 2 somewhat lower denticles. Jung tentati\ely re-
ferred both lots to N. exuvioides Trechmann, 1935, a
species described on the basis of one incomplete speci-
men from the Pliocene of Carriacou in the Grenadines.
Jung (1971) redescribed this latter specimen as having
only 12 ribs whose edges overhang the adjacent inter-
spaces. Vokes (1983) clarified the status of N. exuvioides
when she described a very strongly ribbed shell with 1 1
ribs from the Gatun Formation of Panama. This speci-
men closely resembles the one from Carriacou and is
clearly referable to S. exuvioides. We believe that Jung's
(1965, 1969) specimens from Venezuela and Trinidad
belong neither to N. fulgurans nor to N. exuvioides, but
instead to our new species, iV. fortidentata. This species
appears therefore to be intermediate in sculpture be-
tween N. exuvioides with only 1 1-12 ribs and the Recent
.v. fulgurans, usually with more than 21 ribs. The re-
lationship between N. fortidentata and the Late Oli-
gocene or Early Miocene N. tampaensis Dall, 1892, from
the Tampa Formation of Florida is unclear. Merita tam-
paensis, whose granulated parietal region suggests place-
ment in Theliostyla, is a small species with weakly de-
veloped apertural dentition and highly \ariable external
sculpture, some shells being nearly smooth whereas oth-
ers have fine spiral cords of varying sizes.
The shape of xV. fortidentata suggests that this species
inhabited the upper zones of rocky shores. Merita ful-
gurans, its most similar living relative, is usually found
in areas of reduced salinity , such as the mouths of harbors
(Russell, 1941) or protected embayments. Vermeij (1973)
showed that shells with low globosity, high basal exca-
vation, weak sculpture, and relatively broad apertures
with weak dentition are found in middle to lower inter-
tidal species of Merita; whereas species with a globose,
little excavated, strongly sculptured shell and a small
aperture bordered by strong teeth are found at higher
shore levels. The latter shell form is especially charac-
teristic of the subgenera Cyrnostyla von Martens, 1887,
and Ritena Gray, 1858. Of the living and fo.ssil members
of the subgenus Theliostyla, most of which live in the
middle zones of the intertidal, M . fortidentata most closely
approaches species of Ritena. Other fossils collected with
N. fortidentata, including Oliva, Olivella, Conus, Na-
tica, Polinices, Strombus, Phalium, Dentalinm. and
Corlmla, suggest a variety of different environments,
implying post-mortem transport and mixing of assem-
blages.
BIOGEOGRAPHY OF AMERICAN MERITA
Among the fossil species of Merita that have been de-
scribed from late Eocene and younger strata in tropical
.\merica, at least two have close affinities with li\ing
Indo-\^'est-Paci£ic species. Merita listrota Woodring,
1973, from the late Eocene (?) Gatuncillo Formation of
Panama has a finely ribbed shell w ith a peripheral keel,
fine teeth on the outer lip, 7 teeth on the columellar lip,
and a sparseK papillate parietal region. Woodring (1973)
noted the striking similarity' between A', listrota and the
recent mangrove-associated M. planospira Anton, 1839
(the type and only known species of the subgenus Ily-
nerita von Martens, 1887) from the tropical Indo-Pacific.
He doubted that the tw^o species were closely related, in
part because M. planospira has 5 rather than 7 columellar
denticles. We consider the similarities to be so numerous
that an inference of close relationship seems warranted.
If A', listrota belongs to the subgenus Ilynerita, as we
believe it does, that subgenus may be added to the grow-
ing list of taxa whose distributions became restricted to
the Indo- West-Pacific during the Tertiary (N'ermeij,
1986). As Vokes (1983) points out, \. (Theliostyla) exu-
vioides may represent a second lineage that has become
restricted (as N. exuvia Linnaeus, 1758) to the Indo-
West-Pacific. Merita exuvioides differs from the Western
Pacific M. exuvia chiefly by having 1 1 instead of 14 strong
overhanging spiral cords on the body whorl.
The other fossil species of Merita that have been de-
scribed from late Eocene and younger deposits in tropical
America do not easih fit with any living members of the
genus. These are M. hadra Woodring, 1973, from the
late Eocene (?) Gatuncillo Formation of Panama and N.
oligopleura Dall and Ochsner, 1928, from the Pleistocene
of the Galapagos. Merita hadra has very fine spiral threads
on the bod\' whorl and a smooth parietal region, whereas
M. oligopleura has 3 broad spiral ribs on the body whorl.
Although the record of Merita in tropical America is
very meager, the history of the genus points to multiple
instances of extinction and geographical restriction. It is
too earl\ to assess the scope of these changes, but the
record of Merita suggests that intertidal species have been
no less affected by events leading to extinction and re-
striction than have species from the better-sampled fossil
environments of subtidal bottoms.
ACKNOWLEDGEMENTS
T.M.C. gratefully acknowledges a Smithsonian Tropical
Research Institute short-term fellow ship, directed by Jer-
emy Jackson, during w hicli large collections of Late Ter-
tiary fossil mollusks were made. Collecting was made
more fruitful and pleasant by the expertise and company
of the other members of the field part\-, A. G. and J. M.
Coates. We thank Laurel B\ bell, Thomas M. Cronin, and
Harry Dowsett of the U.S.G.S. for their age determi-
nations.
LITERATURE CITED
Anton, H. E. 1839. Verzeichniss der Conchylien welche sich
in der Sammlung von Hermann Eduard .^nton befinded.
Halle, xvi -I- 110 p
Bold, W. A. van den. 1967. Ostracoda of the Gatun For-
mation, Panama. Micropaleontology 13(3):306-318.
G. J. Vermeij and T. M. Collins, 1988
Page 105
Dall, W. H. 1892. Contributions to the Tertiary fauna of
Florida; with especial reference to to the Miocene silex
beds of Tampa and the Pliocene beds of the Caloosahatchie
River. Transactions of the Wagner Free Institute of Sci-
ence 3(2):201-473, pis. 13-22.
Dall, W. H. and W. H. Ochsner. 1928. Tertiary and Pleis-
tocene Mollusca from the Galapagos Islands. Proceedings
of the California Academy of Science 17{4):89-139, pis.
2-7.
Gray, J. E. 1858. Observations on the genus Nerita and its
operculum. Proceedings of the Zoological Society of Lon-
don 26:92-94.
Gmelin, J. F. 1791. In: Linnaeus' Systema naturae, Tom. I,
Pare VI, p. 3021-3909.
Jung, P. 1965. Miocene Mollusca from the Paraguana Pen-
insula, Venezuela. Bulletins of American Paleontology
49(223):389-652.
Jung, P. 1969. Miocene and Pliocene mollusks from Trinidad.
Bulletins of American Paleontology 55(247):293-657.
Jung, P. 1971. Fossil molluscs from Carriacou, West Indies.
Bulletins of American Paleontology 61(269):147-262.
Linnaeus, C. 1758. Systema naturae, 10th ed. Stockholm,
824 p.
Martens, E. von. 1887-89. Die Gattungen Nerita und Neri-
topsis. Svstematisches Conchylien-Cabinet von Martini und
Chemnitz 2(11):1-147.
Menke, K. T. 1851. Conchylien von Mazatlan, mit kritischen
Bemerkungen. Zeitschrift fiir Malakozoologie, yr. 8.
Morch, O. A. L. 1852 Catalogus conchyliorum quae reliquit
D. Alphonso d Aguirra & Gadea Comes de Yoldi, Copen-
hagen, 170 p.
Russel, H. D. 1941. The Recent molluscs of the family Ne-
ritidae of the western .Atlantic. Bulletin of the Museum of
Comparative Zoology at Harvard College 88(4):347-404.
Trechmann, C. T. 1935. The geology and fossils of Carriacou,
West Indies. The Geological Magazine 72(858):528-555,
pis. 20-22.
Vermeij, G. J. 1970. The Nerita ascensionensis species com-
plex (Gastropoda: Prosobranchia) in the South Atlantic.
The Veliger 13(2):135-138.
Vermeij, G. J. 1973. Morphological patterns in high intertidal
gastropods: adaptive strategies and their limitations. Ma-
rine Biology 20:319-346.
Vermeij, G. J. 1984. The systematic position of the neritid
prosobranch gastropod Nerita polita and related species.
Proceedings of the Biological Society of Washington 97(4):
688-692.
Vermeij, G. J. 1986. Survival during biotic crises: the prop-
erties and evolutionary significance of refuges. In: D. M.
Elliott (ed.). Dynamics of extinction. Wiley, New York, p.
234-246.
Vokes, E. H. 1983. Nerita exuvioides Trechmann (Mollusca:
Gastropoda) from the Gatun Formation of Panama. Tu-
lane Studies in Geology and Paleontology 17(4): 131-134.
Woodring, W. P. 1973. Geology and paleontology of Canal
Zone and adjoining parts of Panama: description of Ter-
tiary Mollusks (additions to gastropods, scaphopods, pe-
lecypods: Nuculidae to Malleidae). U.S. Geological Survey
Professional Paper 306-E:453-539.
Woodring, W, P. 1982. Geology and paleontology of Canal
Zone and adjoining parts of Panama: description of Ter-
tiary Mollusks (Pelecypods: Propeamussidae to Cuspida-
riidae; additions to families covered in P.306-E; additions
to gastropods; cephalopods). U.S. Geological Survey
Professional Paper 306-F:541-759.
THE NAUTILUS 102{3):106-1{)9, 1988
Page 106
A New Species of Favartia from the Eastern Pacific
(Gastropoda: Muricidae)
Anthony D'Altilio
Barbara ^ . Myers
Department of Marine Invertebrates
San Diego Natural History Museum
San Diego, CA 92101, USA
ABSTRACT
Favartia (Murexiella) shasktji is described from Isla del Coco,
Costa Rica, and compared with related species from the eastern
Pacific, This species is known onl\ from this isolated oceanic
island
INTRODUCTION
Isla del Coco (also known as Cocos Island), one of the
National Parks of Costa Rica, is a small, uninhabited
island situated approximately 600 kilometers south south-
west of Puntarenas, Costa Rica, at 5°33'N latitude and
87°03'W longitude. Isla Cascara and Roca Sucia are two
of the many islets surrounding Cocos Island. In May,
1985, Donald R. Shasky of Redlands, California, and
Kirstie L. Kaiser of La Canada, California, collected five
specimens of the new species described herein, at depths
of 50-80 feet (15.2-24.4 m), under dead coral off these
two islets. In a previous paper (D'Attilio, M>ers & Shasky,
et al., 1987), a new species of Phyllonotus Swainson,
1833, was described from the same area. Montoya (1983,
1984) published a bibliography of Cocos Island molluscan
faunal studies.
SYSTEMATICS
Family Muricidae Rafinesque, 1815
SubfamiK Muricopsinae Radvvin & D',\ttilio, 1971
Genus Favartia Jousseaume, 1880
Type species: Miirex breviculus Sowerby , 1 834, by orig-
inal designation.
Subgenus Murexiella Clench & Perez Farfante, 1945
Type species: Mtirex hidalgoi Crosse, 1869, b>- original
designation.
Favartia (Murexiella i shaskyi new species
(figures 1-10)
Description: Shell (figures 1, 2) broadly fusiform, spire
'/3 shell length; protoconch of holot\pe eroded, proto-
conch of paratype 1 (figure 3) of 3'/2 smooth, lustrous,
convex whorls, with axial buttresses arising from the te-
leoconch; teleoconch with 5-6 subangulate whorls; suture
weakly impressed; whorls gentK sloping from suture to
shoulder; body whorl % shell length; aperture ovate; in-
ner lip erect posteriorly; outer lip thin, erect, crenulate,
reflecting the spiral cords; siphonal canal '3 shell length,
broad proximalK', tapering distalK', with narrow \entral
opening, terminally recurved, tube-like; siphonal fascicle
of 3 fine distal portions of previous canals; body whorl
with 6 broad varices, penultimate whorl with 8 varices;
varices broader than intervarical regions; spiral sculpture
of 6 cords on body whorl, all nearK equal in width,
packed closely together; 2 additional cords between
body whorl and siphonal canal, 2 major and 1 minor
cord on canal; 2 cords per whorl on spire; shoulder of
body whorl without spiral cords; all cords terminate as
spines on varices (figures 4, 5), posterior spines recurved,
anterior spines long, projecting ventrally; spiral cords
(figures 6, 7) strongly scabrous; scales prominent, fine,
white, erect, closely packed, disguising the contour of
cords; cords and scales microscopicalK grooved, incre-
mentally incised; adapertural sides of varices, especially
last varix, completeK- scaled; scales arranged in 3 tiers
(figure 6), with tube-like spines on tier farthest from
aperture; radula (figure 8) t>picall\ muricopsine, with
broad, U-shaped basal plate, and strongK projecting cusps.
Paratypes 1 and 2 with only 4 postnuclear whorls; suture
more strongly impressed, spire angulate.
Figures 1, 2. Favartia (Murexiella) shaskyi new species. 1 . Holot> pe, L'SNM 860012, Isla Cascara, Cocos Island, Costa Rica, under
dead coral slab, in 24.4 m, 23.0 mm long. 2. Paratype 1, SDNHM 91873, Roca Sucia, Cocos Island, Costa Rica, under dead coral
slab, in 24.4 m, 14.0 mm long.
A. D'Attilio and B. W. Myers, 1988
Page 107
Page 108
THE NAUTILUS. Vol. 102, \o. 3
Figures 3-7. Favartia (Murexiella) shaskyi new species, details of shell sculpture 3. Protoconch of parat\pe 1. 4, 5. Adapertural
(4) and abapertural (5) views of last varix of holotype. 6, 7. Microscuipture along adapertural (6) and abapertural (7) surfaces of
a spiral cord of the holotype. Figure 8. Rachidian and right lateral teeth from the holotype.
Color: Base color buff to tan, with diffuse darker band
encircling body whorl; scales white, aperture white, si-
phonal canal white.
Type locality: Isla Cascara, Cocos Island, Costa Rica,
under dead coral slab, in 24.4 m.
Holotype: USNM 860012 (figure 1 ), 23.0 mm long, 14.2
mm wide.
Paratypes: Paratype 1: SDNHM 91873 (figure 2), 14.0
mm long. Paratype 2: Collection of D. R. Shasky, 11.8
mm long, both from Roca Sucia, Cocos Island, Costa
Rica, under dead coral, in 24.4 m.
Additional material examined: Tw o specimens, collec-
tion of K. Kaiser, 19.2 mm long, and 14.7 mm long, Isla
Cascara, Cocos Island, Costa Rica, under rocks, in 15.2-
24.4 m.
Etymology: We are pleased to name this species for
Donald R. Shasky, M.D., who collected the three type
specimens, and who generousK donated the holotype
and parat\pe 1.
DISCUSSION
This new species has a distinct morphology that easily
distinguishes it from all other eastern Pacific species of
A. D'Attilio and B. W. Myers, 1988
Page 109
the subgenus Murexiella. Favartia (Murexiella) lappa
(Broderip, 1833), a closely related species, differs in hav-
ing a higher spire, biconic form, and short, stubby, non-
recurved spines. Favartia (Murexiella) vittata (Broderip,
1833) also has a more or less biconic shape, but the body
whorl is more globose, and the spines, although recurved,
are short and stubby. Favartia (Murexiella) keenae (E.
H. Yokes, 1970) has somewhat similar sculpture, a larger
shell with a globose body whorl strongly constricted at
the base, and a strongly impressed suture. We have care-
fully studied and compared the 12 additional nominal
species of Murexiella, none of which are closely related
to this new species.
ACKNOWLEDGEMENTS
We are grateful to Dr. Donald R. Shasky for donating
the holotype and paratype and Kirstie Kaiser for the loan
of her two specimens. We thank David K. Mulliner for
the photography. M. G. Harasewych, EmiK H. Yokes,
William K. Emerson and Carole M. Hertz reviewed the
manuscript and made valuable suggestions.
LITERATURE CITED
Broderip, W. 1833. Characters of new species of Mollusca
and Conchifera collected by Mr. Cuming. Proceedings of
the Zoological Society of London 2:173-179.
Clench, W. J. and I. Perez Farf ante. 1945. The genus Mt/rei
in the western Atlantic. Johnsonia l(17):l-58.
Crosse, H. 1869. Diagnoses Molliiscorum novorum. Journal
de Conchyiiologie 17:408-410.
D'Attilio, A., B W. Myers, and D. R. Shasky. 1987. A new
species of Phyllonotus (Muricidac: Muricinae) from Isla
del Coco, Costa Rica. The Nautilus 101(4):162-165,
Jousseaume, F. 1880. Division methodique de la famille des
Purpurides. Le Naturaliste 42:335-336.
Montoya, M. 1983. Los Moluscos marines del la Isla del Coco,
Costa Rica. I. Lista anotada de especies. Brensia 21:325-
353.
Montoya, M. 1984. Marine mollusks of Cocos Island, Costa
Rica. I. Bibliographic compilation of species. Western So-
ciety of Malacologists Annual Report (1983) 16:33-44,
Radwin, G. E. and A. D'Attilio. 1971. Muricacean supra-
specific taxonomy based on shell and radula. Echo 4:55-
67.
Rafinesque, C. S. 1815. Analyse de la nature ou tableau du
univers et des corps organises. Barravecchia, Palermo, p.
136-149.
Sowerby, G. B., II. 1834. Conchological illustrations. Murex.
Sowerby, London, pis. 58-67.
Swainson, W. 1833. The zoological illustrations, ser. 2, vol. 3,
pi. 67.
Yokes, E. H. 1970. The west American species of Murexiella
(Gastropoda: Muricidae). Veliger 12(3):325-329, pi. 50.
THE NAUTILUS 102(3):110-114, 1988
Page 110
Conus baccatus G. B. Sowerby 111, 1877
A Panamic Fauna! Constituent
^ illiam K. Emerson
Waller E. Sage III
Department of Invertebrates
American Museum of Natural History
New York, NY 10024, USA
ABSTRACT
Specimens of a small species of Conus recent!) received from
Golfo de Chiriqui, Republica de Panama, are confirmed to be
referable to Conus baccatus Sowerby , 1877, a species originally
described from an unknown locality. The provenance and iden-
tity of this taxon had remained uncertain for more than 100
years until Rockel (198.5a) compared two specimens from Pa-
cific Panama in his collection with the holotype of Conus bac-
catus and determined the specimens to be conspecific. We
agree with Rockel's conclusion that this species has been con-
fused in the past w ith the pustulate form of the west American
Conus perplexus Sowerby, 1857, which is a somewhat similar,
but distinct species. For Conus perplexus, a lectotype is selected
and the type locality is restricted to the Golfo de Panama. For
Conus baccatus. the Golfo de Chiriqui is designated the type
locality.
INTRODUCTION
In 1877, G. B. So\verb%- III briefly described and illus-
trated in color Conus baccatus from an unknown locality.
Apparently based on a single specimen in the collection
of Dr. Prevost of Alen^on, France, the figured specimen
later was owned by J. C. Melvili (Sowerby, 1887:251)
and subsec}uently by J. R. Le B. Tomlin (1937:217). The
vast MeK ill-Tomlin Collection eventual!) w as deposited,
along with this type, in the National Museum of Wales,
Cardiff. Coomans et al. (1982:4) examined the holotype
and provisionally accepted Conus baccatus as a valid
species. They also rejected the Cjuestionable placement
by Walls (1979:726) of this ta.xon in the synonymy of C.
mindanus Hwass in Bruguiere, 1792, from the western
Atlantic. \'ink (1984:356) subsequently noted certain
similarities of the holotype of C. baccatus with C. selenae
Van Mol, Tursch, and Kempf, 1967, a Brazilian taxon
which we consider referable to the C. jaspidcus (Jmelin,
1791, species complex.
Through the generosit)- of Carol Skoglund and Robert
Koch, we have examined a series of well-preserved spec-
imens from Golfo de Chiriqui. Panama that are referable
to Coijus baccatus. Our findings confirm Rockel's (1985a:
29) conclusion that this taxon is a valid Pacific Panamic
species, which for many years has been masquerading
in collections as a pustulose form of Conns perplexus
Sowerb), 1857.
A specimen apparently referable to C. baccatus was
illustrated by Kiener (1846:56, 57, pi. 83, fig. 2) as "Conus
albomaculatus. Reeve ", in reference to Sow.[erby] (1833,
Conus fig. 2, which was named C. bicolor h\ Sowerby
in 1833 and renamed C. albomaculatus h\ Sowerb) in
1841). Kiener "s figured specimen, measuring 22 mm in
height, was attributed to the Dupont Collection from an
unknown locality. This specimen cannot be located in
the Museum National d Histoire Naturelle, Paris, and
the fate of the Dupont collection is not known (P. Bou-
chet, in lift.. March 4, 1987). Reeve (1849:3) considered
Kiener 's illustrations to represent the western Atlantic
species C. mindanus Hwass in Bruguiere, 1792. He also
stated that "C. albimaculalus [sic] [Sowerb) ] has more
resemblance with very young [pustulose] specimens of
C. lithoglyphus [.sir] [Hwass in Bruguiere, 1792] ', a con-
clusion accepted by Coomans et al. (1979:97) and Rocke!
(1985b, nr. 22). Sowerb)- "s figured specimen of C. albo-
maculatus has not been located. It is not in the Britisli
Museum (Natural History) (K, M. Way, in litt.. May 11,
Figures 1-7. Conus baccatus Sowerby, 1877. 1, 2. Holotype, NMW no. 1955.158.29; x 2 (courtesy of R. M Filmer) 3. 4.. Off
Isla Coiba, Panama, AMNH no, 221871a; x 2. 5. Off Isla Parida, Panama, AMNH no. 173698a; x 5 6, 7. Off Isla Coiba, Panama.
AMNH no. 221871; x 2, Figures 8-16. Conus perplexus Sowerby, 1857, 8. 9. Lectotype, BM(NH) no. 1978118; x 2. 10. Pla\a
de los Angeles, Bahia Tenacalita, Jalisco, Mexico, dredged, mud bottom, Aug., 1975, AMNH no. 221873a, ex C and P. Skoglund;
X 5. 11, 12. Paralectotype, BM(NH) no. 1978118a; x 2 13, 14. Paralectotype, BM(NH) no. 1978118b; x 2. 15. 16. "West
Panama" (note the distorted spire), AMNH no. 212547; x 2 Figures 17. 18. Conus puncticulatus form papillosus Kiener, 1849,
Cartagena, Colombia, Iseach, 1977, ex J. M. Bijur Coll., AMNH no. 225979; x 2. Figure 19. Conus yemanjae \aii Mol et al.,
1967, paratype, pi. 8, fig. lb, Fortaleza (Ceara), Brazil, ex pisce; x 2.
W. K. Emerson and W. E. Sage III, 1988
Page 111
Page 112
THE NAUTILUS, Vol. 102, No. 3
1987) nor is it in the National Museum of Wales (A.
Trevv, in litt.. Ma\, 1987). Therefore, the taxonomic
status of C. alboviaculatus Sowerby, 1841, cannot be
determined from the available data.
Although C. baccatus was not formally named by Sow-
erb\ ill until 1877, specimens were apparentl\ available
in European collections by the nii(l-I9th century, as sug-
gested by Kiener's figured specimen from the Dupont
collection. There is also a specimen of C. baccatus labeled
""Panama" in the F. A. Constable Collection (AMNH
47740), which dates from the latter part of the 19th
century and was received by the American Museum of
Natural History from the estate of Louise Constable in
1929.
Abbreviations for institutions used in text; AMNH =
.\merican Museum of Natural History; BM(NH) = Brit-
ish Museum (Natural History); Los Angeles County Mu-
seum of Natural History = LACMNH; NMW = National
Museum of Wales.
SYSTEMATIC ACCOUNT
Conus baccatus Sowerby, 1877
(figures 1-7)
? "Conus albomaculatus. Reeve"', Kiener, 1846:56, 57, pi. 83,
fig. 2 (apertura! and dorsal views; as "Conus albimaculatus
[sic] Sow.[erby] ), from unknown locaiit) , Dupont collec-
tion. Not C. bicolor Sowerby, 1833: pt 24, fig. 2, renamed
without explanation C. albomaculatus Sowerby, 1841.
Conus baccatus Sowerby, 1877:753, 754, pi. 75, fig. 5, from
unknown locality; Tryon, 1883: ser. 1, vol. 6, p. 22, pi. 6,
fig. 92 [copy of original fig.], "Habitat unknown "; Sowerby,
1887: vol. 5, p. 251, Conus pi. 29, fig. 660 [copy of original
fig.], "Habitat unknown. Coll. Meiviii"; Tomlin, 1937:217,
"Hab. ?, Type in coll. Tomlin"; Coomans et a/., 1982:4,
fig. 197, apertural and dorsal views of the holotype; Vink,
1984:354-358, pi. 20, fig, 1, dorsal view of holotype; Rock-
el, 1985a:29, dorsal view of holotype and dorsal and aper-
tural views of two recently collected specimens from "Pe-
rida Island ", Pacific Panama (figured in color).
Original description (in part): "Shell short, rather swol-
len, very minutely decussated, with regular rows of con-
spicuous granules, whitish, with large orange blotches
arranged in three bands; spire acute, short, whorls con-
cave, nearly smooth, last whorl biangulated. . . . Apart
from its somewhat stunted form, the delicacy of its mark-
ings and rows of gem-like granules, it is remarkable for
the double angle at the top of the body whorl. Long. 23,
lat. maj. 15 mill " [Holotype actually measures 22.2 by
14.2 mm.)
Type depository: Holotype in the National Museum of
Wales, no. 1955.158.29, fide Trew (1982:3); here illus-
trated (figures 1, 2).
Type locality: Off Isla Parida, Golfo de Chiriqui, Re-
piiblica de Panama, here designated.
Range: Known only from Pacific Panama.
Material examined: ,\MNH Collection: From Golfo de
Chiritpii, Pacific Panama: 6 specimens, no. 173698 and
8 specimens, no. 211778, off Isla Parida (8''05'N, 82°20'W),
in 5.5-9 m, sand bottom. May, 1972, ex R. E. Hubert; 2
specimens, no. 210849, off Isla Parida, sand, in 5.5 m,
March, 1974, ex R. E. Hubert; 3 specimens, no. 225981,
off Isla de Canal de Afuera, in 55 m, ex J. Ernest; 2
specimens, no. 221871, off Isla Coiba, in 24-40 m, March
28, 1986, ex C. and P. Skoglund; 2 specimens, no. 221872,
off S.E. Isla Rancheria, in 2-7.5 m, March 27, 1986, ex
C. and P. Skoglund; 2 specimens, no. 223269, off Isla
Rancheria (7°38'N, 81°44'W), in 3-9 m, coral sand and
debris bottom, February 24, 1987, ex R. Koch. From
"Panama"; 1 specimen, no. 47740, ex F. A. Constable
Coll.; 3 specimens, no. 201566, ex T. H. and V. B. Mun-
yan Coll.
Remarks: Specimens of Conus baccatus superficially
resemble small examples of the pustulose form of Conus
perplexus. There are several discrete differences in shell
morphology which serve to distinguish the two taxa. 1,
Nuclear whorls: In C. baccatus, I'i whorls, nucleus short,
terminating in a broad, well-defined mammillated pro-
cess. In C. perplexus, 2'/2 whorls, nucleus elongate, nar-
row, terminating in a needle-like projection (cf. figure 5
with figure 10). 2, Shoulder margin of bod\ whorl: In C.
baccatus, oblique, twofold-angled (i.e., biangulated). In
C. perplexus. monoangulated (cf. figures 6, 7 with figures
8, 9). 3, Spiral granulations; In C baccatus, rows of pus-
tules more widely spaced, some rows weakK developed.
In C. perplexus, rows consistently spaced, nearly uni-
formly developed (cf. figures 6, 7 with figures 15, 16).
4, Color pattern: In C. baccatus, spiral bands of broken
blotches, orange in faded specimens, reddish-broun in
fresh specimens. In C. perplexus, narrow spiral lines with
dark brown dots (cf. figures 1, 2 and 3, 4 with figures 8,
9). Additionally, the periostracum in C. baccatus is yel-
lowish tan, whereas in C. perplexus it is a brownish tan.
The largest specimen of C. baccatus examined (AMNH
221871a; figures 6, 7) is 26.9 mm in height, compared
to a maximum of 37.8 mm for C. perplexus (AMNH
206684) from Isla Cebaco, Golfo de Montijo, Panama.
Hanna (1963:39) reports a specimen of the latter taxon
from Isla San Jose, Golfo de Panama, with a height of
41.5 mm. Both of these taxa have shells with a distinct
posterior notch on the lip (cf. figure 5 with figure 10),
whereas C. xirnenes Gray, 1839, C. nmhogani Reeve,
1843, and C. tornatus Sowerb\, 1833, which also have
been confused with C. perplexus, have a narrow anal
notch (see Wolfson, 1962: figs. 6a-d; Tucker 1985: fig.
c; and Chancy, 1987: figs. 2, 3). All of these nominal
species have populations sympatric with those of C. bac-
catus, which is known only from Pacific Panamic waters.
Conus perplexus, with which C. baccatus is most likely
to be confused, ranges from the northern part of the Gulf
of California, south to Mancora, Peru, 4°05'S, 81°03.5'W
(LACMNH no. 70-101). Specimens of C. perplexus in
which the interrupted spiral lines of brown dashes or dots
are raised into prominent pustules are more commonly
found in the southern populations, from Panama south-
ward to Peru (figures 15, 16). .\ similar pustulose form
also occurs in populations of the western .\tlantic C
puncticulatus Hwass in Bruguiere, 1792, which is a twin
W. K. Emerson and W. E. Sage III, 1988
Page 113
species of C. perplexus. This form was named C. pa-
pillosus Kiener, 1849 (figures 17, 18).
On the basis of shell morphology, C. baccatus appears
to be most closely related to members of the western
Atlantic C. jaspideus species complex, especially the pus-
tulose, short-spired Brazilian forms which were given the
names C. yemanjae (figure 19) and C. selenae by Van
Mol et al. (1967: pi. 8); also figured by Vink (1984: pi.
20).
Conus perplexus Sowerby (1857:20, 1858: pi. 200, fig.
324) was briefly described in Latin, followed by the com-
ment; "This shell perplexes me, because there is a variety
of C. puncticulalus [Hwass in Bruguiere, 1792, a Carib-
bean twin species] which it nearly resembles. This, how-
ever, is quite smooth, and more angular' . A single spec-
imen was figured by Sowerby in a dorsal view, with the
statement: "Gulf of California, West Columbia [sic],
Cum.[ing]". Tomlin (1937:290) recorded the presence of
three "types" in the British Museum (Naural History),
which were kindh' loaned to us by Ms. K. M. Way
[BM(NH) 19781 18^ here illustrated, figures 8. 9, 11-14].
The original labels confirm these specimens as syntypes,
although none of them can be referred with certainty to
Sowerby s figured specimen. The drawing may represent
a composite. The three syntypes are somewhat faded,
but are otherwise well-preserved. We have selected as
the lectotype (figures 8, 9) the syntype that most closely
resembles the original figure. The lectot\pe is slightly
larger in height (26.8 vs. 26.1 mm) than the specimen
depicted in the original illustration.
The original labels accompanying the syntype lot of
Conus perplexus list the Gulf of California and West
Columbia [.sic], which are the same localities cited by
Sowerby (1858, caption to Conus, pi. 14). Hugh Cuming
is not believed to have collected in Mexican waters on
his voyage to the west coast of South America, 1828-30.
He is known, however, to have made extensive collections
in the "Gulf of Panama, the Pearl Islands, and the Gulf
of Chiriqui", as well as visits to Costa Rica, Nicaragua,
and Honduras (Dance, 1986:114). We here restrict the
type locality of C. perplexus to the Golfo de Panama, as
the Isthmus of Panama was part of Colombia at that
time.
In summary, Conus baccatus G. B. Sowerby III, 1877,
is recognized as a Panamic faunal constituent and as a
twin species of C. jaspideus Gmelin, 1791, from the
western Atlantic, especially the pustulose form named
C. yemanjae Van Mol et al., 1967. Conus baccatus is
compared with pustulose specimens of the sympatric C.
perplexus G. B. Sowerby II, 1857, with which it has been
confused, as well as with pustulose specimens of C. punc-
ticulalus Hwass in Bruguiere, 1792, a Caribbean twin
species of C. perplexus. A lectotype is selected and the
t\pe locality is restricted for C. perplexus. A type locality
is designated for C. baccatus.
ACKNOWLEDGEMENTS
We are indebted to Carol Skoglund and Bob Koch, Phoe-
nix, Arizona, for sending us comparative material, for
their observations on the differences in the nuclear whorls
of Conus perplexus and the specimens which were found
to be referable to C. baccatus. Their interest and coop-
eration, past and present, are greatly appreciated. We
acknowledge with thanks the assistance of Philippe
Bouchet, Museum National d'Histoire Naturelle, Paris;
Helen DuShane. Whittier, California; James Ernest, Bal-
boa, Repiiblica de Panama; R. M. Filmer, Chobham,
England; Robert A. Foster, Santa Barbara, California;
Barbara Good, San Diego, California; James H. McLean,
Los Angeles County Museum of Natural History; P. Gra-
ham Oliver and Alison Trew, National Museum of Wales,
Cardiff; Donald R. Shasky, Redlands, California; Kay C.
Vaught, Scottsdale, Arizona; and K. M. Way, British Mu-
seum (Natural History), London, for the loan of speci-
mens and/or photographs, or for providing information.
We also are indebted to Alan Kohn, University of Wash-
ington for providing data on the complicated nomen-
clatural history of the taxa Conus bicolor and C. albo-
maculatus. Stephanie Crooms, AMNH, word-processed
the manuscript.
We owe a special debt of gratitude to Henry W. Cha-
ney, Santa Barbara Museum of Natural History; James
H. McLean; and James Nybakken, Moss Landing Marine
Laboratories, for critically reading drafts of the manu-
script.
LITERATURE CITED
Born, I. von. 1778. Index rerum naturalium Musei Cesarei
Vindobonensis. Pars prima, Testacea. Vienna, xlii + 458
p., 1 pl.
Bruguiere, J,-G. 1792. Cone. In: Encyciopedie methodique.
Historie Naturelle des Vers 1:586-757, illus.
Chaney, H. W. 1987. A comparative study of two similar
Panamic cones: Conus ximenes and Conus mahogani. The
Veliger 29(4):428-436, 6 figs.
Coomans, H. E., R. G. Moolenbeek, and E. Wils. 1979. Al-
phabetical revision of the (sub)species in Recent Conidae
2. adansoni io atbuquerquei. Basteria 4;3(5-6):81-105, figs.
26-50.
Coomans, H. E., R. G. Moolenbeek, and E. Wils. 1982. Al-
phabetical revision of the (sub)species in Recent Conidae.
5. baccatus to byssinus, including Conus brettinghami
nomen novum. Basteria 46(l-4):3-67, figs. 172-292.
Dance, S. P. 1986. A history of shell collecting. E. J. Brill,
Leiden, xv -I- 265 pp., illus.
Gmelin, J. F. 1791. Systema naturae per regna tria naturae,
13th ed. 1(6), cl. 6, Vermes. Leipzig, p. .3021-3910.
Gray, J. E. 18.39. Molluscous animals and their shells. In: F.
W. Beechey. The zoology of Capt Beechey's Voyage . . .
to the Pacific and Behring's Straits in His Majesty's ship
Blossom. . . . London, p. i-xii -I- 103-155, pis. 33-44.
Hanna, G D. 1963. West American mollusks of the genus
Conus — II. Occasional Papers of the California Academy
of Sciences 35:103 p., 4 text figs., 11 pis.
Kiener, L.-C. 1846[-48], 1849[-.50]. Species general et ico-
nographie des coquilles vivantes. Vol. 2, Conus. Paris, p.
1-379, 111 pis. [Conus alhomaculatus, 1846, pp. 56, 57,
pl. 83, fig. 2; C. papillosus. 1849, p. 271, pl; 72, fig. 4]
Mol, J.-J. Van, B. Tursch, and M. Kempf. 1967. Resultats
scientifiques des campagnes de la 'Calypso' 32. Campagne
Page 114
THE NAUTILUS. \ol. 102, No 3
de la Calypso au large des cotes atlantiques de I'Amerique
de Sud (1961-1962) 16. Mollusques Prosobranches; les
Conidae du Bresil. Annales de I'lnstitute Oceanographique
Monaco 45(2):233-254, 6 pis.
Reeve, L. 1843. Conchologia iconica, \ ol 1 London Conus.
London, 47 pis., with text.
Reeve, L. [184S-]1849. Conchologia iconica, \'ol. .5. Conus,
emendations, London, 7 p., 9 pis , with te.xt.
Rtickel, D. 1985a. Conus Imccatus Sowerby III, 1877, a for-
gotten valid species from tropical west-America. La Con-
chiglia, Rome, yr. 17, nos. 192-193, p. 29, 8 figs.
Rockel, D. 198.5b. Die Familie Conidae, nr. 22 [Contis lito-
glyphus Hwass in Bruguiere, 1792]. Privately issued, 1 p.,
4 figs
Shaw, H. O. N. 1909. On the dates of issue of Sowerby s
"Conchological Illustrations." from the cop\ presented to
the Radcliffe Library. Oxford. Proceedings of the Mala-
cological SocietN of London 8(6):333-340.
Sowerby, G.B., I. 1833[-40], 1841. Conchological illustrations,
Conus, pt. 24, fig. 2 [published March 29, 1833]; list for
Conus section, p. 1-4 [published in 1841]. London. [See
Shaw, 1909:340]
Sowerby, G. B., II. 1857-58. Thesaurus conchyliorum. Vol.
3, pts. 17, 18. London, pp. 1-24, pis. 187-195, 196-210.
[Description of Conus perplexus, 1857, p. 20; 1858, pt. 18,
pis. 196-210; figure of C. perplexus, 1858, pi. 200, fig.
324, Conus pi 14]
Sowerby, G. B., III. 1877. Descriptions of six new species of
shells from the collections of the Marchioness Paulucci and
Dr Prevost. Proceedings of the Zoological Society of Lon-
don, for 1876, p. 752-755, pi. 75. [Issued Jan. 4, 1877]
Sowerby, G.B., III. 1887. /n.G. B. Sowerby II (ed.). Thesaurus
conchyliorum. Second supplement to Monograph of the
genus Conus, Vol. 5, pt. 44 London, p 249-279, pis. 507-
512, Conus pis. 29-36 [Conus baccatus, p. 251, pi. 507,
fig. 660, Conus pi. 29. Issued July. 1887]
Tomlin, J R le B. 1937. Catalogue of Recent and fossil cones.
Proceedings of the Malacological Societ) of London 22(4-
5):205-330.
Trew, A. 1982. The .Melvill-Tomlin Collection, part 10, Co-
nacea (Conidae). Handlists of the Molluscan Collections
in the Department of Zoology, National Museum of Wales,
Cardiff, Ser. 1:1-28 + i-ix. '
Tryon, G. W., Jr. 1883[-84]. Manual of concholog>, Ser. 1,
Vol. 6, family Conidae, pp. 1-1.50, 31 pis. [Part 21, pp. 1-
64, Dec. 27, 1883]
Tucker, J. K. 1985. Conus ximenes Gra\, 1839, Conus ma-
hogany [sic] Reeve, 1843 and friends. Shells and Sea Life
17(8):201-202, figs, a-e, 1-5. Bayside, California.
Vink, D. L. N. 1984. On the identity of Conus anaglypticus,
Conus baccatus, Conus selenae and Conus yemanjae.
Journal of Concholog\. London 31(6):353-.358, pi. 20.
Walls, J. G. 1979 Cone shells, a synopsis of the living Conidae.
T.F.H. Publications, Inc., Neptune Cit\. NJ, 1011 p, illus.
Wolfson, F. H. 1962. Comparison of two similar species of
Conus (Gastropoda) from the Gulf of California. Part 1:
a statistical analysis of some shell characters. The Veliger
5(l):23-28, 6 figs.
THE NAUTILUS 102(3):115-122, 1988
Page 115
References to Molluscan Taxa Introduced by Linnaeus in the
Systema Naturae (1758, 1767)
Kenneth J. Boss
Miiseuiii of Comparative Zoology
Harvard liniversity
Cambridge, MA 02138, USA
ABSTRACT
Bibliographic documentation is provided for the more than 90
published resources utilized by Linnaeus as citations for mol-
lusks in the 10th and 12th editions of his Systema Naturae.
INTRODUCTION
Linnaeus's 10th edition of the Systema Naturae of 1758
is the cornerstone of our system of binominal nomencla-
ture in zoology. Many species of mollusks were described
therein, and these have been the subject of individual
investigations such as revisions of particular taxa as well
as studies of the specimens themselves (Hanley, 1855;
Dodge, 1952-59; Dance, 1967).
Linnaeus also attempted to apply the binominal meth-
od to the references that accompanied his treatment of
the species. If he cited a reference that previously men-
tioned or figured the species under discussion, he usually
gave an abbreviated citation, which frequently embod-
ied a shortened name of the author and that of the in-
dividual work. Many of these are readily apparent to
someone with a knowledge of the pre-Linnaean litera-
ture; thus, "Gault. test." may be easily recognized as
being a reference to Gaultieri's Index Testarum Con-
chyliorum of 1742; others are more enigmatic.
Some years ago I thought that it might be interesting
to decipher these puzzling references in Linnaeus's Sijs-
tema, both the 10th (1758) and 12th (1767) editions (the
nth is a reprint of the 10th), much like Wheeler (1979)
has done for fishes. However, in correspondence with
the late Professor A. Myra Keen, I found that both she
and Dr. S. S. Berry also had an interest in this problem.
Professor Keen sent me a short draft of some 30 titles
that she had connected with Linnaeus's abbreviations,
and I supplied her with several that she had been unable
to find. Eventually, she published a narrative account of
certain authors whom Linnaeus had cited (see Keen,
1983a, b). In these papers, she noted that Linnaeus used
about 40 different references in his sections on mollusks.
However, when the different citations are tallied for both
soft bodied and shelled mollusks [including therefore
Linnaeus's categories with their genera, Vermes: Intes-
tina: Teredo, Vermes: MoUusca: Clio, Limax, Doris, Te-
thijs, Scyllaea, and Sepia; and Vermes: Testacea: (all
genera with the exception of Lepas and Serpula though
references to the two molluscan Serpula are included)],
the total exceeds 90. Dance (1967) also provided a short
list of volumes personally owned by Linnaeus which were
utilized in various editions of the Systema.
Unknown to any of us were the studies of Professor
John L. Heller, Emeritus, of the Department of Classics
of the University of Illinois on various botanical works
of Linnaeus (see Heller, 1983, for a collection of these
articles). He provided extensive bibliographic references
to similarly abbreviated citations by Linnaeus in some
of his botanical works such as the Species Plantarum
and the Hortus Cliffortianus. He also explicated certain
particularly equivocal abbreviations used by Linnaeus in
referring to various insects (Heller, 1961) and planned
to finish a complete bibliographical guide to zoological
works cited by Linnaeus under the proposed title Bib-
liotheca Zoologica Limmeana (Heller, 1968, 1979, 1983);
this work is currently in press (Heller, personal com-
munication) but will contain more than a malacologist
need know.
Although I am separately preparing a more extensive
treatment of the pre-Linnaean malacological citations
including comments on subsequent editions, translations
into other languages as well as annotations of the holdings
of the Harvard College Library system (Boss, in prepa-
ration), I present below the bibliographic abbreviations
utilized by Linnaeus in ihe Systema Naturae (1758, 1767)
and the full citation for each reference. Of all the ref-
erences, only one remains undeciphered: Linnaeus (1758:
708, 1767:1161) referred to "Tessin. epist. 1 n. 28 Cym-
bium" under Argonauta argo. This is presumed to have
been a letter of Count Carl Gustaf Tessin, on whose
collection Linnaeus worked. Despite a thorough check-
ing of apposite sources (i.e., items in the Literature Cited
as well as subsequent editions of the Systema and several
classical cephalopod authorities such as Chemnitz, Ke-
ferstein, Conrad, Naef, Robson, Tryon, and Ferussac-
Orbigny), this reference could not be located.
There are several bibliographic sources for pre-Lin-
naean works, and sometimes there are discrepancies in
Page 116
THE NAUTILUS, Vol. 102, No. 3
citations of the sources. These may arise due to different
bibliographic methods, through error, or due to the vari-
ability in the copies of pre-Linnaean works, which were
not always issued in uniform copies I have relied prin-
cipally on the Catalogue of the Books . . . British Mu-
seum .... Soulsby (1933), Engelmann (1846), Heimann
(1957), Nissen (1969), and the resources of \arious Har-
vard libraries. The birth and death dates of authors have
been included, mainly as listed in the British Museum
Catalogue; these do not always concur with such dates
given by other authors, principally Dance (1986) or Nis-
sen (1969). I have utilized my own method of punctua-
tion and have spelled out such words as "engraved" and
"portraits" when describing the texts; I have followed
standard procedure for the usage of brackets, etc., as
noted in the Catalogue of the Books . . . British Museum.
Further, the names of publishers or printers were added
when 1 knew them. With few exceptions, which are listed
as "not seen", examination oi the original publication
was possible.
ABBREVIATED CITATIONS UTILIZED
BY LINNAEUS FOR MOLLUSKS
Act. angl. 301, p. 2051 (see Breyn, 1705)
Act. helv. 4. p. 212, t.9, f.21, 22 (see Hofer, 1760)
Act. helv. 5. p. 283, n.4, t.3, f.25, 26 or n.5 t.3 f. 27, 28
(see Schlotterbeccius, 1762)
.\ct. paris. 1710, p. 463 (see Reaumur, 1710)
Act. paris. 1711, t.3, f.4,5 (see Reaumur, 1711a)
Act. paris. 1711, p. 199, t.6, f.5.7. or 9 (see Reaumur,
1711b)
Act. paris. 1712, p. 163 (see Reaumur, 1712)
Act. Petropol. 7, p. 321, t.U, 12 (see Koelreuter, 1761)
Act. Upsal. 2. p. 560, t.l52, f.4. idem or f.a. (see Bromell,
1729)
Adans. sen. or seneg. (see Adanson, 1757)
Aldr. exs. or exsang. or exangu. (see Aldrovandi, 1606)
Aldr. ins. (see Aldrovandi, 1602)
Amoen. acad. (see Amoenitates Academiae)
Amoen. acad. 1, p.325 (see Balk, 1746)
Amoen. acad. 3: 256 (see Odhelius, 1754)
Argenv. conch, (see Dezallier d'Argenville, 1742)
Barr. ic. or rar. or Barrel, icon, or rar. (see Barrelierus,
1714)
Baster subs, (see Raster, 1759-65)
Bell. aqu. or aquat. or Rellon. aqu. (see Belon, 1553)
Blank, ins. (see Blankaart, 1688)
Bocc. observ. (see Boccone, 1674)
Bohads. mar. (see Bohadsch, 1761)
Ronan. kirch. (see Buonanni, 1709)
Bonan. recr. or recreat. or Bonann. recr. (see Buonanni,
1681, 1684)
Brad, natur. or Bradl. nat. or Bradl. natur. (see Bradley,
1721)
Breyn. polyth. (see Breynius, 1732)
Brown, jam. (see Browne, 1756)
Bruckm. cent. 2 epist. (see Brueckmann, 1743)
Calceol. mus. (see Cerutus and Chiocco, 1622)
Chin. Lagerstr. (see Odhelius, 1754)
Column, aqu. or aquat. (see Colonna, 1606)
Column, phytob. (see Colonna, 1592)
(Column, purp. (see Colonna, 1616)
Crew. mus. (see Grew, 1681)
E.N.C. (see Francus de Frankenau, 1727)
Edw. av. (see Edwards, 1758-64)
Ellis cor. or corall. (see Ellis, 1755)
En. svec. (see Linnaeus, 1746)
Faun. svec. or En. svec. (see Linnaeus, 1746)
Frisch ins. (see Frisch, 1730)
Gesn. aqu. or aquat. (see Gesner, 1551-87)
Ginam. adr. (see Ginnani, 1755-57)
Ginan. or Ginan. adr. (see Ginanni, 1755-57)
Gmelin act. petrop. vol. 3, p. 246 (see Gmelin, 1729)
Gnalt. test, (see Gualtieri, 1742)
Grev. mus. or Grew. mus. (see Grew, 1681)
Gron. lap. (see Gronovius, 1740)
Gualt. test, (see Gualtieri, 1742)
Hasselq. or Hasselqv. itin. (see Hasselquist, 1757)
Hasselqv. act. ups. 1750. p. 33. (see Hasselquist, 1750)
Hevde .Ant. Anatome mvtulorum 1683 oct (see Heide,
1683)
Imperat. nat. (see Imperato, 1599)
It. gotl. (see Linnaeus, 1747)
It. oel. or oeland (see Linnaeus, 1745)
It. wgot. or wgoth. or wogth. (see Linnaeus, 1747)
Johnst. aquat. or exsangu. (see Jonstonus, 1650-53)
Jonst. aquat. (see Jonstonus, 1650-53)
Kaehl. or Kaehler act. Stockhb. 1754. p. 144, t.3, f.A-E
or A-F (see Kahler, 1754)
Kirch, mus. (see Buonanni, 1709)
Klein or Klen. ostr. (see Klein, 1753)
Kratz. or Kratzenst. Regenf. (see Regenfuss, 1758)
Labat. itin. (see Labat, 1722)
Lederm. micr. (see Ledermueller, 1760-61)
Lewenh. arcan. (see Leeuwenhoek, 1695)
List. angl. (see Lister, 1678)
List. angl. app. [appendices] (see Lister, 1685, under Lis-
ter, 1678)
List, conch, or hist, (see Lister, 1685-92[-97])
List. exer. 2 or exerc. 2 or exercit. 2 (see Lister, 1695)
List, exercit. anat. 1 (see Lister, 1694)
List, exercit. anat. 2 (see Lister, 1695)
M.L.U. or Mus. L. U. (see Linnaeus, 1764)
Mus. Ad. Fr. (see Linnaeus, 1754)
Mus. Tess. or Tessin (see Linnaeus, 1753)
Needham microsc. (see Needham, 1745)
Olear. mus. (see Olearius, 1674)
Osb. iter, or Osbeck. iter, (see Osbeck, 1757)
Pet. or Petiv. amb. or amboin. (see Petiver, 1713)
Pet. or Petiv. gaz. (see Petiver, 1764)
Pet. mus. (see Petiver, 1695)
Plane, conch, (see Plancus, 1739)
Reaum. act. paris. 1712, p. 163 (see Reaumur, 1712)
Regenf. conch, (see Regenfuss, 1758)
Roes. ins. or in.sect. (see Roesel von Rosenhof, 1746-61)
Rond. aqu. or aquat. (see Rondeletius, 1554-55)
Rond. or Rondel, pise, or test, (see Rondeletius, 1554-
55)
K. J. Boss, 1988
Page 117
Rumph. mus. (see Rumpf, 1705, 1711, 1741)
Scheuch. Diluv. (see Scheuchzer, J. J., 1716)
Scheuch. helv. (see Scheuchzer, j., 1708)
Seb. or Seba mus. (see Seba, 1734-65)
Sellii historia Teredinis. Traject. or Sellii monogr. ultra
or Sellii Tered. (see Sellius, 1733)
Sloan, jam. (see Sloane, 1696)
Stobaei Diss, epist. Lund, (see Stobaeus, 1732)
Strom, sondm. 173 (see Strom, 1762)
Swamm. or Swammerd. bibl. (see Swammerdam, 1737-
38)
Tessin. epist. 1, n. 28 (not found; apparently a letter to
Linnaeus)
Tournef. iter, (see Tournefort, 1717)
Tulp. obs. (see Tulpius, 1739)
Vallisn. nat. (see Valiisnieri, 1721)
Wolff, hass. (see Wolfart, 1719)
Worm. mus. (see Worm, 1655)
CITATIONS OF REFERENCES
Adanson, Michel [1727-1806]. 1757. Histoire naturelle du
Senegal. Coquiliages. Avec la relation abregee d un Vo-
yage fait en ce pays, pendant les annees 1749-53. Paris,
Bauche, 2 Parts: Voyage, p. [viii], 190; Coquiliages, p. xcvi,
275, 19 pis., 1 map, 4to.
Aldrovandi, Ulisse [1522-1607]. 1602. De Animalibus Insectis
libri septem cum singulorum iconibus ad \ i\ uni expressis.
Bonon.[iae], Jo. Bapt. BelJagamba, p [.\], 767 [4.3], text
illustrations {circa 720 figs.), 1 portrait, engraved title-
page, folio.
Aldrovandi, Ulisse [1522-1607]. 1606. U. Aldrovandi . . . de
reliquis .Animalibus exanguibus libri quatuor post mortem
ejus editi [with a preface by F. Aldrovandi]: Nempe de
Mollibus, Crustaceis, Testaceis, et Zoophytis. Bononiae apud
lo: Baptistum Bellagambam, p. [viii], 593 [26], 1 portrait,
text illustrations (circa 600 figs.), folio.
AmoenitatesAcademicae [1743-76]. [186 Dissertations by Stu-
dents of Upsala (or Uppsala) University, Linnaeus being
the Praeses from June 30, 1743 to June 22, 1776] 19 vol-
umes, illustrated. I'psaliae. 4to. [These are the 186 disser-
tations by students of Upsala University during Linnaeus's
praeses or administration from June 30, 1743 to June 22,
1776; these dissertations appeared in various editions, and
also were usually individually and separately issued; Lin-
naeus only cited two of these for mollusks (see Balk, 1746
and Odhelius, 1754); however, his citations utilized the
pagination in the collected so-called Linnaeus Edition of
the ,\moenitates ,\cademicae, volumes 1 and 3, respec-
tively, cited under Balk and Odhelius (see Soulsbv, 1933:
99 ff. and Heimann, 1957:83 ff]
Balk, Laurentius. 1746. Museum Adolpho-Fridericianum,
quod consensu ampliss. fac. medicae in Regia Acad. Up-
saliensi sub praesidio viri celeberrimi D D. Caroli Linnaei
speciminis academici loco publico bonorum examini
submittit Laurentius Balk Fil. . , . die XXXI Maji. Anno
MDCCXLVI Typis Laurentii Salvii, Holmiae, p. [vii],
48, [2], 2 pis. 4to. [See also: 1749. Museum Adolpho-
Friedericianum sub praesidio D.D. Car. Linnaei propo-
situm a Laurent Balk (May 31, 1746) Amoenitates Aca-
demicae 1:277-327, pis. XII-XIV, Holmiae et Lipsiae]
Barrelierus, Jacobus (or Barrelier, Jacques) [1606-73]. 1714
Plantae per Galliam, Hispaniam et Italiam observata . . .
Opus posdiumum, accurantc Antonio de Jussieu in lucem
cdilutn, et ad recentiorum normam digestuni; cui accessit
ejusdem auctoris specimen de insectis quibusdam marinis
mollibus, crustaceis et testaceis. Parisiis, apud Steph. Ga-
neau, p. [4], xxvi, [8], 8, 140, engraved title-page and atlas
of 334 pis. with 1327 figures.
Raster, Job [1711-75], 1759-65 [1762]. J. Basteri . .. Opuscula
subseciva, observationes miscellaneas de Animalculis et
Plantis quibusdam marinis, eorumque ovariis et seminibus
continentia. Harlemi, J. Bosch, vol. 1, p. 148, [6], pis. I-
XVI; vol. 2, p 100, pis. I-IX, 4to.
Belon, Pierre [1517-64], 1553. P. Bellonii . . De Aquatilibus,
libri duo. cum Eiconibusad vivum ipsorum effigies, cjuoad
eius fieri potuit, expressis. Parisiis, Car. Stephanus, p. [xxx],
448, with text illustrations (187 woodcuts), oblong 8vo.
Blankaart (or Blankaert), Steven [1650-1702]. 1688. Schou-
burg der Rupsen, Wormen, Ma'den, en Vliegende Dier-
kens daar uit voortkomende; door eigen ondervindinge by
en gebragt etc. (Aanhangsel van Brieven, &c. ). Amster-
dam, Jan Ten Horn, p. viii, 232, 22 pis. engraved title-
page, 8vo. [not seen].
Boccone, Paolo (afterwards Silvio) [1633-1704]. 1674. Re-
cherches et observations naturelles . . . Amsterdam, chez
Jean Jansson a Waesberge, p. (6), 328, 14 pis.
Bohadsch, Joannes Baptista [1724-72]. 1761. J.B.Bohadsch . . .
de quibusdam Animalibus marinis, eorumque proprieta-
tibus orbi litterario vel nondum vel minus notis liber, cum
nonnullis tabulis aeri incisis, ab auctore supervivus ani-
malibus delineatis. Dresdae, Walther, p. [xviii], 169, 12
pis., 4to,
Bradley, Richard [?-1732]. 1721. A philosophical account of
the Works of Nature ... To which is added an account of
the state of Gardening, as it is now in Great Britain, and
other parts of Europe, &c. London, W. Means, p. [xx], 194,
28 colored pis., 4to.
Breynius [Breyn], Joannes Philippus [1680-1764]. 1705 [1706].
II. Epistola D. Joannis Philippi Breynii M.D. & Societal
Reg. Soc. De Plantis & Insectis quibusdam rarioribus in
Hispania observatis. Philosophical Transactions [of the
Royal Society], giving some account of the present un-
dertakings, studies and labours of the Ingenious in many
considerable parts of the World, London, MDCC\ I, Vol.
XXIV (For the Years 1704 and 1705) No. 301, p. 2045-
2055, tabs. 1-2.
Breynius [Breyn], Joannes Philippus [1680-1764]. 1732. I. P.
Breynii . . . Dissertatio physica de Polythalamiis, nova tes-
taceorum classe . . . Huic adiicitur commentatiuncula de
Belemnitis Prussicis; tandemque Schediasma de Echinis
methodice disponendis. Gedani, Corn, a Beughem, p. [vi],
64, 12 pis., 4to.
Bromell, Magnus von [1679-1731 J 1729. Lithographicae sve-
canae. continuatio. caput secundum. Teslacea fossilia la-
pidesque conchiformes continens. Articulus primus de tes-
taceis marinis variis, quae Integra & immutata e tellure
effodiuntur. Acta Literaria (et Scientiarum) Sveciae. Up-
saliae & Stockholmiae. Trimestre Tertium, Anni
MDCCXXIX, p. 554-562, plate to accompany p. 557.
Browne, Patrick [1720?-90]. 1756. The civil and natural his-
tory of Jamaica, &c. London, p. viii, 503, 49 pis., 1 map,
folio.
Brueckmann, Franz Ernst [1697-17.5.3]. 1749. Centuria (II)
secunda Epistolarum itinerarium. Wolffenbuttelae, 1296
p., 44 pis. [not seenj
Buonanni, Filippo [1638-1725]. 1681. Ricreationedell'occhio
e della mente nell' osservation' delle Chiocciole, proposta
Page 118
THE NAUTILUS, Vol. 102, No. 3
a" curiosi delle opere della natura, &c. Roma, Varese, 4
parts, p. [xiv], 38-4 [1.5], 109 pis., 3 engraved title-pages, 3
engraved half-titles, 4to.
Buonaniii, Filippo [1638-1725]. 1684. Recreatio mentis, et
oculi in ohservatione .Animalium Testaceorum curiosis na-
turae inspectoribus. Italico sermone primum proposita . . .
nunc denuo ab eodem Latine oblata, centum additis Tes-
taceorum iconibus, circa quae varia problemata propo-
nuntur. Romae, Varese, p. xvi, 270 [10], 139 pis., 2 en-
graved titles, 3 engraved half-titles, 4to.
Buonanni, Filippo [16.38-1725]. 1709. Musaenm Kircheria-
num, sive Musaeum a P. .\thanasio Kirchero in (.'ollegio
Romano Societatis Jesu . . descriptum, & . . . illustratum,
&c. Romae. Geo. Plachi. p. (.\ii], 522, 173 pis., folio.
Cerutus, Benedictus [-1620] and .Andrea Chiocco [-1624]. 1622.
Musaeum F. Calceolarii [F'rancesco Calceolari or Francis-
cus Calceolarius or Calzolari, Francesco, the Younger], jun.
. . . a B. Ceruto . . . incaeptum et ab A. Chiocco ... lu-
culenter descriptum & perfectum, &c. Veronae, Ang. Ta-
mus, p. [46], 746, 43 pis., folio.
Colonna, Fabio [1567-1650]. 1592. Phytobasanos [in Greek],
si\e Plantarum aliquot Historia . . . .Accessit etiam Piscium
aliquot, PlantarunKpif novarum Hi.storia eodem auctore.
Neapoli, p. [xiv], 120, .32 [8], text illustrations, 4to.
Colonna, Fabio [1567-16,50]. 1606. Minus cognitarum Stir-
pium aliquot, ac etiam rariorum nostro coelo orientium
'Ekphrasis [in Greek] . . . Item de Aquatilibus. aliisque;
.\nimalibus quibusdam paucis libellus . . . opus nunc pri-
miim in lucem edituni Romae, apud Guil. Facciottum,
p. [vi], 340, Ixiii [7]. text illustrations, 4to.
Colonna, Fabio [1567-1650). 1616. F. Columnae. .. Purpura.
Hoc est de Purpura ab Animali testaceo fusa, de hoc ip.so
.'\nimali, aliisque rarioribus Testaceis quibusdam, &c. (De
Glossopetris dissertatio). Romae, Jac. Mascardus, p. [iv],
42, 8 pis., 4to.
Dezallier d'Argenville, Antoine Joseph [1680-1765]. 1742.
L'Histoire Naturelle eclaircie dans deux de ses parties prin-
cipales, la Lithologie et la Conchyliologie . . . Par M
[i.e.. A. J. Dezallier d Argeriville]. Paris, de Bure, 2 Parts,
p. [viii], 491, .36 pis., 4to.
Edwards, George F.R.S. [1694-1773]. 17.58-64. Gleanings of
Natural History, exhibiting figures of Quadrupeds, Birds,
Insects, Plants &c. (Glanures d'Histoire Naturelle . . . Tra-
duit de I'Anglois [Pt. 1 & II] par J. Du Plessis, [Pt. Ill, by
E. Barker].). London, printed for the author at the Royal
College of Physicians, 3 vols., text in English and French
in parallel columns, p. xxxv, vii, 347, colored pis. ccxi-
ccclxii, 4to
Ellis. John [1710r'-76]. 17.55. An essay towards a natural his-
tory of the C^orallines, and other marine productions of
the like kind, commonly found on the coasts of Great
Britain and Ireland. To which is added the description of
a large marine Polype taken near the north pole . . . in . . .
1753. London, A. Millar, p. xvii [x], 103, 39 pis., 4to.
Francus de PVankenau [Franck von F'ranckenau], Georg Fred-
eric [1644-1704] 1727 Observatio XXI. Dn D Georgii
Friderici Franci de Frankenau Calva serpentis americani
diademata. .'\cta Physico-medica Academiae (^aesareae
Lepoldino-Carolinae [full title: Acta Ph)sico-medica Aca-
demiae Caesareae [Leopoldino-Carolinac] naturae curio-
sorum exhibentia ephermerides sive observationes historias
et experimenta a celeberrimis germaniae et exterarum
regionum viris habita et communicata singulari studio col-
lecta], Norimbergae. Vol. 1, p. 63-64, plate 1, figure V,
nos. 1 and 2, 4to.
Frisch, Johann Leonhard [1666-1743]. 1720-38. Beschrei-
bung von allerley Insecten in Teutsch-Land, nebst niitz-
lichen Anmerckungen . . . von diesem . . . inlandischen
Gewijrme, &c. Berlin, Nicolae, 13 Parts [in 1 vol.], illus-
trated, 272 pis., 4to.
Gesner (or Gessner), Conrad [1516-65]. 1558. C. Gesneri . . .
Historiae Animalium. Liber IlII. qui est de Piscium &
Aquatilium Animantium natura . . . Continentur in hoc
volumine G. Rondeletii ... & P. Bellonii . . . de Aquatilum
singulis scripta. Tiguri, C. Froschoverus, p. [4], 1297, 616
woodcuts, folio.
Ginanni, Giuseppe. Count [1692-1753J 1755-57. Opere pos-
tume, &c. 2 Volumes: Tomo I. Cento quattordici piante
che vegetano nel Mare .Adriatico. N'enezia, G. Zerletti, p.
[xii], xix, 63, 55 pis.; Tomo 11. Testacei marittimi paludosi
e terrestri dell' Adriatico e del Territorio di Ravenna
(Memorie del Conte G. Ginanni, &c.), p. [viii], 72, 35 pis.,
folio.
Gmelin, Johann Georg [1709-.55]. 1729 Radiis .Xrticulatus
Lapideis Commentarii .Academiae Scientiarum Imperialis
Petropolitanae, \'ol. 3, p. 246-264, pis 11-12, 4to
Grew, Nehemiah [1641-1712]. 1681. Museum Regalis Socie-
tatis: or a description of the natural and artificial rarities
belonging to the Royal Society and preserved at Gresham
Colledge . . . Whereunto is subjoyned the comparative
anatomv of stomachs and guts. London, VV. Rawlins, 2
Parts, illustrated, p. [12], .386, [2]; 43 p.. 31 pis., folio.
Gronovius, Joannes Fredericus [1690-1760). 1740. Index su-
pellectilis Lapideae, quam collegit in classes et ordines
digessit specificis nominibus ac synon\mis illustravit Jo-
hannes Fredericus Gronovius. Lugduni Batavorum, 8vo.
[17.50. Editio altera., p. [xii], 106, [6].]
Gualtieri, Niccolo [1688-1747]. 1742. Index Testarum Con-
chyliorum quae adservantur in Museo N. Gualtieri . . . et
methodice distributae exhibentur tabulis C.X. (Introductio
ad historiam Testaceorum a ... P. Tournefortio in codice
MS. relicta). Florentiae, .\lbizzini, 5 Pt., p. xxiii, 110 pis.,
with descriptive letterpress, 1 portrait (dated 17.35), folio.
Hasselquist, Fredrik [1722-52J 17.50. Octopodia sepiae species
missa a Fr. Hasselquest. Smirnae 1749 .Acta Societatis
Regiae Scientiarum Upsaliensis, Upsaliae & Stockholmiae,
p. 33-34.
Hasselquist, Fredrik [1722-52). 1757. F. Hasselquists . . . Iter
Palaestinum, eller Resa til Heliga Landet fcirrattad ifran
ar 1749 til 17.52, med beskrifningar, ron, anmiirkningar,
ofver de markviirdigaste Naturalier . . utgifven af C Lin-
naeus. Stockholm, [Lars Salvii], p xiv. 619, 8vo.
Heide, .Antonius de. 1683 [1684]. Anatome Mytuli, Belgice
Mossel, structuram elegantem ejusque motum mirandum
exponens. Subjecta est centuria observationum medica-
rum. Jansson & Waesberge, Amsterdam, 2 Parts, p. 200,
8 pis. [Mytilus. p. 1-48, 8 pis], small 8vo.
Hofer. Joannis. 1760. Observatio Zoologica. Acta Helvetica
Phvsico-Mathematico-.Anatomico-Botanico-Medica, Basi-
leae, vol. 4, p 212-213. pi IX, figs. 21-22.
Imperato, Ferrante[ 1.5.50-1625]. 1.599 Dell'llistoria Naturale
. . libri XXX'lll. nella ipiale ordinatamente si tratta della
diversa coiulilion di Miniere. e Pietre. Con alcune historic
di Piante, & .Animali; sin hora non date in luce. Napoli,
Constantino Vitale, p. [xxiv], 791, 1 plate, text illustrated
(129 woodcuts), folio.
Jonstonus, Joannes [1603-75]. 16.50-53. Historiae naturalisde
Exanguibus .A<iuaticus — de Insectis — de Serpentibus. Libri
IV. I'rancofurti a. M , M. Merian, p. 78, [1], 20 pis., folio,
Kahler, Martin. 1754. .Angiiende en ny art af vatten-polyper
K. J. Boss, 1988
Page 119
som ata sten. Kungliga Svenska Vetenskap Academiens
Ilandlingar, L'ppsala and Stockliolm, vol. XV, p. 143-146,
plate III? figs A-t-:
Klein, Jakob Theodor [1685-1759] 1753. J T Klein Tenta-
men methodi Ostracologicae, sive di.spositio naturalis Con-
chlidum et Concharuni . . .Accedit lucubratiuncula de
formatione, cremeiito et coloribus Testarum . . . Turn com-
mentariolum in locum Plinii Hist. Nat. libr. IX, cap. XXXIII,
de Concharuni differentiis, Denique sciagraphia methodi
ad genus Serpentium ordinate digerendum. Lugduni Ba-
tavorum, Wishoff, p. [viii], 177, [35], 44, 16, [2], 12 pis.
text illustrated, 4to.
Koelreuter, Joseph Theophil. 1761. Polypi marini, russis ka-
rakatiza, recentioribus graecis o'ctapus [in Clreek] dicti des-
criptio. Novi Commentarii Academiae Scientiarum Im-
perialis Petropolitanae. Petropoli, vol. \'II, p. 321-343, pis.
11, 12.
Labat, Jean Baptiste [1663-1738]. 1722. Nouveau Voyage aux
isles de I'Amerique, contenant I'histoire naturelle de ces
pays . . . Avec une description exacte & curieuse de toutes
ces Isles, &c. Paris, Cavelier, 6 volumes, illustrated, 12vo.
Ledermueller, Martin Frobenius [1719-69]. 1760-61. M.F.
Ledermiiller's . . . mikroskopische Gemiiths- und Augen-
Ergotzung, &c. Niirnberg, 2 Parts, p. [xvi], 202 pp., 102
colored pis., 4to.
Leeuwenhoek, Anthony van [1632-1723]. 1695. Arcana Na-
turae detecta. Delphis Batavorum, Krooneveld, p. [vi], 568
[14], 27 pis., 1 portrait, engraved title-page, text illustra-
tions, 4to.
Linnaeus, Carolus [1707-78], after 1761 ennobled as Carl von
Linne. 1745. Carl Linnaei . . Oelandska och Gothland-
ska Resa pa Riksens hogloflige standers bef alining f orrattad
ahr 1741. Med anmiirckningar uti oeconomien, natural-
historien, antiquiteter, &c. [Half-title]. Carl Linnaei Goth-
landska Resa. Forrattad ahr 1741. Stockholm & Upsala,
Hos Gottfried Kiesewetter, p. xii, 344. [30], 1 plate, 2 maps,
text illustrated, 8vr
Linnaeus, Carolus [1707-78]. 1746. Caroli Linnaei ... Fauna
Svecica, sistens Animalia Sveciae Regni: Quadrupedia, Aves,
Amphibia, Pisces, Insecta, Vermes, distributa per Classes
& Ordines, Genera & Species. Cum differentiis Specierum,
synonymis Au torum, nominibus Incolarum, locis Habi-
tationum, descriptionibus Insectorum. Sumta & literis Lau-
rentii Salvii, Stockholmiae, p. [xxvii], 411, 2 pis., engraved
frontispiece, 8vo.
Linnaeus, Carolus [1707-78]. 1747. Carl Linnaei . . . Wast-
giita-Resa, pa Riksens hogloflige Standers befallning for-
rattad ar 1746. Med anmarkningar uti OEconomien, Na-
turkunnogheten, Antiquiteter, Inwanarnes Seder och
Lefnads-satt, med tilhcirige figurer. Stockholm, Uplagd pa
Lars Salvii kostnad, p. xi, 284. [20], 5 pis., 8vo.
Linnaeus, Carolus [1707-78]. 1753. Museum Tessinianum,
&c., opera . . . Comitis Dom. Car. Gust. Tessin . . . collec-
tum. Hans Excellence . . . Gr. Carl. Gust. Tessins naturalie-
samling. Apud L. Salvium: Holmiae, p. [vi], 123 [9], 12
pis., folio.
Linnaeus, Carolus [1707-78]. 1754. Museum S:ae R:ae M:tis
Adolphi Friderici ... in quo Animalia rariora imprimis,
et exotica: Quadrupedia, Aves, .Amphibia, Pisces, Insecta,
Vermes describuntur et determinantur, Latine et Svetice,
cum inconibus, jussu Sac Reg Maj:tis a Car. Linnaeo, Equ.
Hans Maj:ts Adolf Frideriks . . . naturalie samling inne-
hallande salsvnte och frammande djur, som bevaras pa
Kongl. Lust-Slottet Ulriksdahl. Beskrefneochafritadesamt
pa nadig befallning utgif ne af Carl Linnaeus. Typographia
regia Direct. Pet. Momma, Holmiae, p. xxx, 96 [8], 33 pis.,
folio.
Linnaeus, Carolus [1707-78]. 1764. Museum Sae R:ae M:tis
Ludovicae Ulricae Reginae Svecoruni, Gothorum, Van-
dalorumque ... in quo Animalia rariora, exotica, imprimis
Insecta & Conchilia describuntur & determinantur. Pro-
dromi inslar editum, &c. Holmiae, Literis & impensis Di-
rect. Laur. Salvii, p. vi, 720 [2], 8vo.
Lister, Martin (c. I638-17I2]. 1678. M. Lister . . . Historiae
Animalium Angliae tres tractatus. Unus de Araneis. Alter
de Cochleis turn terrestribus turn fluviatilibus. Tertius de
Cochleis marinis. Quibus adjectus est quartus de Lapidibus
ejusdem insulae ad cochlearum quandam imaginem fi-
guratis. Londini, J. Martyn, p. [vi], 250, 9 pis., 4to. [Ap-
pendix, &c. p. 23, 1 pi. Eboragi, 1681, 4to.; Appendicis
. . . altera editio, &c., p. [ii], 45, 2 pis., Londini, 1685, 8vo.]
Lister, Martin [c. 1638-1712]. 1685-92[-97]. M. Lister, His-
toriae sive Synopsis methodicae Conchyliorum . . . liber
primus (-liber IV. — Appendix ad . . . librum IV. — [Tabu-
larum Anatomicarum explicatio.]) . . . S. et A. Lister figuras
pin. Londini, Autor, 6 Parts [in 2 volumes], folio.
Lister, Martin [c. 1638-1712]. 1694. Lister exercitatio ana-
tomica. In qua de Cochleis, maxime terrestribus & Li-
macibus, agitur. Omnium dissectiones . . . illustrantur, &c.
Londini, Smith and Walford, p. xi, 208, 8 pis. 8vo.
Lister, Martin [c. 1638-1712]. 1695. M. Lister exercitatio ana-
tomica altera, in qua maxime agitur de Buccinis fluviati-
libus & marinis, &c. Londini, Autor, p. [xiv], 7, 267, 128,
6 pis., 8vo.
Needham, John Turberville (as N., T.) [1713-81]. 1745. An
account of some new microscopical discoveries found on
an examination of the Calamary . . . Also observations on
the Farina /ort'!/)u/ans of Plants . . , Likewise observations
on the supposed embryo Sole-fish fixed to the bodies of
Shrimps ... A description of the Eels or Worms in blighted
Wheat; and other curious particulars relating to the Nat-
ural History of Animals, Plants, &c. London, Needham,
p. viii, 126, 6 pis., 8vo.
Odhelius, Johannes Laurentius. 1754. Specimen academicum
sistens Chinensia Lagerstromiana, quod . . . Praeside . . .
Carolo Linnaeo . . . submittit Johannes Laurentius Odhe-
lius ... d [23] Decembr. MDCCLIV. Impressit Jacob
Merckell, Holmiae, p. [iii], 36, 1 plate, 8vo. [See also 1759.
LXI. Chinensia Lagerstromiana {sic), praeside D.D.Car.
Linnaeo proposita a Johann Laur. Odhelio (December 23,
1754). Amoenitates Academicae 4:230-260, 1 pi., Hol-
miae.]
Olearius, Adam [1599-1671]. 1674. Gottoffische Kunst-Kam-
mer worrinnen allerhand ungemeine Sachen, so theils die
Natur, theils kiinstliche Hande hervorgebracht und be-
reitet, &c. Schlesswig, [Gottfredt Schulkens], p. [x], 80, 36
pis., engraved title page, 4to.
Osbeck, Pehr [1723-1805]. 1757. Dagbok ofver en Ostindisk
Resa aren 1750-52, med anmarkningar uti Naturkun-
nigheten, frammande folkslags sprak, &c. (En Ostindisk
Resa til Suratte, China, &c. Fran 1750 ... til 1752 . . .
forrattad af O. Toren, &c.) Stockholm, [Lor. Ludv. Gre-
fing], p. [vi], 376, [16], 12 pis., 8vo.
Petiver, James [c. 1658-1718]. 1695(-1703). MuseiPetiveriani
centuria prima (secunda-decima), Rariora Naturae con-
tinens: viz. Animalia, Fossilia, Plantas, ex variis mundi
plagis advecta, ordine digesta, et nominibus propriis sig-
nata. London, p. 93 [3], 2 pis., 8vo.
Petiver, James [c. 1658-1718]. 1713. Aquatilium Animalium
Amboinae, &c. Icones & nomina. Containing near 400
Page 120
THE NAUTILUS, Vol. 102, No. 3
figures ... of aquatick crustaceous and testaceous Animals;
. . . found about Amboina. and the neighboring Indian
shores, with their Latin, English. Dutch, and native names,
London, J. Bateman, p. 4, 20 pis., folio.
Petiver, James [c. 16.58-1718]. 1764. J. Petiveri Opera, his-
toriam naturalem spectantia; or Gazophylaeium, &c. Lon-
don, 3 vols, illustrated, 8vo. and folio.
Plancus, Janus (pseudonym for Simon Giovanni Paolo Bianchi)
[1693-177.5]. 1739. J. Planci . . . de Conchis minus notis
liber, cui accessit specimen aestus reciproci maris superi
ad littus portumque .•Kriminl. V'enetiis, Pasquali aeri au-
toris, p. 88. 5 pis.. 4to.
Reaumur, Rene ,\nton Ferchault de [1683-1757]. 1710. Du
mouvement progressif, et de quelques autres mouve-
men[t)s de diverses espeees de Coquilles, Orties & Etoiles
de mer. Histoire de I'Academie Royale des Sciences, Avec
les Memoires de Matheinatiques & de Physiques ....
Memoires . . . Annee MDCC;.\ [printed MDCC.XX.XIl],
Paris, p. 439-490 [p 809-877 in 1777 4to. reprint], pis.
IX-XII, 8vo
Reaumur, Rene Antciii IVrcluuilt de [1683-1757]. 1711a. Des
Differentes Manieres Dont phisieurs espeees d animaux de
mer s'attachent au .sable, au.v pierres et les uns aux autres.
Histoire de lAcademie Royale des Sciences, Avec les Me-
moires de Mathematiques & de Phvsiques .... Memoires
. . . Annee MDCC;XI [printed MDCicXXX], Paris, p. 108-
134 [p. 140-175 in 1777 4to. reprint], pis. 1-3, 8vo.
Reaumur, Rene .'Knton Ferchault de [1683-1757]. 1711b. De-
couverte dune nouvelle Teinture de Pourpe, & diverses
experiences pour la comparer avec celle que les anciens
tiroient de quelques espeees de coquillages que nous trou-
vons sur les Cotes de I'Ocean. Histoire de I .\cademie Ro-
yale des Sciences, Avec les Memoires de Mathematiques
& de Phvsiques .... Memoires . . . Annee MDCCXI [print-
ed MDCCXXX], Paris, p. 166-196 [p. 218-258 in 1777
4to. reprint], pi. 6, 8vo.
Reaumur, Rene .\nton Ferchault de [1683-1757]. 1712. Sur
le mouvement progressif de quelques coquillages de mer,
sur celui des herissons de mer, & sur celui, d une espece
d etoile. Histoire de 1 .'\cademie Royale des Sciences, Avec
les Memoires de Mathematiques & de Physiques ....
Memoires . . . Annee MDCCXII [printed MDCCXXXI],
Paris, p. 115-145 [p. 148-191 in 1777 4to. reprint], pis. [I-
III] 6-8, 8vo.
Regenfuss, Franz Michael [1713-80]. 1758. .'Kuslerlesne
Schnecken Muschelii uiid andre Schaalthiere . nach den
Originalen gemalt, in Kupfer gestochen, und mit natiir-
lichen Farben erleuchlet — (Jioix de Coquillages et de
Oustaces [with an introduction by J. A. Cramer and text
by C. G. Kratzenstein, P. Ascanius and L. Spengler], &c.
Kopenhagen, A. H. Godiche, p. [xiv], xiv, 22, Ixxxvii, 12
colored pis., engraved title-page, folio.
Roesel von Rosenhof, August Joliaini[ 1705-59]. 1746-61. Der
monatlich-herausgegebenen Insecten-Beliistigung erster (-
vierter) Theil (. , nebst einer zuverliissigen Naehricht von
den Lebensumstiinden des seel. Verfassers, beschrieben
und herausgegeben von C.F.C Kleemann), &c. Niirnberg,
4 Parts [vol. 1 (1746); vol. 2 (1749); vol. 3 (1755); vol. 4
(17.55)], I engraved portrait, 3 engraved frontispieces, 357
colored illustrations, 4to.
Rondeletius, Gulielmus (Rondelet, Guillaume) [1.507-66]. 1554-
55. G. Rondelelii . l.ibri de Piscibus Marinis, in quibus
verae Piscium effigies expressae sunt, &c; Universae aqua-
tilium historiae pars altera, cum veris ip.sorum Imaginibus
. . . Lugduni [Lyons], apud Matthiam Bonhomme, 2 Vols.:
Vol. 1 (1554), p. [16], 583, [23], circa 250 woodcuts; Vol.
2 (1555), p. [12], 242, [9], circa 170 woodcuts, folio.
Rondeletius, Gulielmus (Rondelet, Guillaume) [1507-66]. 1558.
C. Gesneri . . historiae .Aninialiuni . liber IIII . Con-
tinentur in hoc volumine, G. Rondeletii . . . de Aquatilium
singulis scripta, &c. (see Gesner, 1558), folio.
Rumphius, Georgius Everhardus (Rumpf, Georg Eberhard)
[1627-1702]. 1705. D'Amboinsche Rariteitkamer, be-
helzende eene beschryvinge van allerhande zoo weeke als
harde schaalvisschen, te weeten raare Krabben. Kreeften,
en diergelyke Zeedieren, als mede allerhande Hoorntjes
en Schulpen. die men in d'.'Vmboinsche Zee vindt: daar
beneven zommige Mineraalen, Gesteenten, en soorten van
Aarde, die in d .Amboinsche, en zommige omleggende Ei-
landen gevonden worden. Verdeelt in drie booken, &c.
Amsterdam, Fr. Halma, p. [xxviii], .340 [43], 60 pis., 1
portrait, text illustrations and engraved title-page, folio.
Rumphius. Georgius Everhardus (Rumpf, Georg Eberhard)
[1627-1702] 1711. Thesaurus imaginum Pi.scium Tes-
taceorum; quales sunt Cancri, Echini, Echinometra, Stel-
lae Marinae, Aic, ut & Conchlearum quibus accedunt
Conchylia ut Nautilus, Cornu Ammonis, &c., Conchae
univalviae & bivalviae . . . denique Mineralia; uti Metalla,
Lapides & Argillae, variis in locis reperta, &c. Lugduni
Batavorum, p. [ii], 15 [8], 60 pis., 1 jX)rtrait, engraved title-
page, folio. [This consists of the pis. of the 'D'.Amboinsche
Rariteitkamer' with their explanations, see Rumphius
(1705).]
Scheuchzer, Johann [I684-I738]. 1708. .Agrostographiae Ilel-
veticae Prodromus, sistens binas Graminum .Alpinorum
hactenus non descriptorum, & quorundam ambiguorum
decades. [Zurich], p. 28, 8 pis., folio.
Scheuchzer, Johann Jacob [1672-1733]. 1716. Museum Di-
luvianum quod possidet J.J. Scheuchzer. Tiguri, p [xii],
107 [2], 1 pi., 8vo.
Schlotterbeccius[Schlotterbeck]. Philippus Jacobus 1762. Ob-
servatio physica de cochleis quibusdam, nee non de tur-
binibus nonullis, ut & de cochlea quadam petrafacta. .Acta
Helvetica Physico-Mathematico-.Anatomico-Botanico-
Medica, Basileae, vol. 5, p. 275-288, pi. Ill [actually pis.
Ill A and III B with a total of 37 figs.].
Seba, Albertus (1665-1736J 1734-65. Locupletissimi rerum
naturalium thesauri accurata descriptio, et iconibus arti-
ficiosis.simis expressio, per universam physices historiam,
&c. Amstelaedami, Jansson-Waesberg. 4 volumes, 449 pis.,
folio.
Sellius, Godofretlus (Sell. Godefried) [■:'-1767] 1733 C; Sellii
. . . Historia naturalis Teredinis, seu .X\ lophagi marini tu-
bulo-conchoidis speciatim Belgici, &c. Trajecti ad Rhen-
um, Besseling, p. [xxxiv], 353 [11], 2 colored pis., 4to.
Sloane, Sir Hans. Bart. [1660-1753]. 1696. Catalogus Plan-
tarum quae in Insula Jamaica sponte pro\eniunt. vel vulgo
coluntur, cum earundeni synonymis & locis natalibus; ad-
jectis aliis (juibusdam quae in Insulis Maderae, Barbados,
Nieves. & .Sancti (^hristophori nascuntur. Seu Prodromi
Historiae Naluralis Jamaicae par.s prima. Londini, p. v,
232 [43]. English translation: 1707-25. A Voyage to the
Islands Madera, Barbados, Nieves, S. Christophers and Ja-
maica, with the natural history of the Herbs and Trees,
Four-footed Beasts, Fishes, Birds, Insects, Reptiles, &c. of
the last of tho.se islands London, 2 vols., illustrated, folio.
Stobaeus. Kllian [1690-1742] 1732. Dissertatio epistolaris ad
Th W (irothaus de nummulo Brattensburgensi, nee non
de noiuiullis aliis ad banc historiae naturalis patriae partem
pertinentibus, imprimis frondosis Cornu .\mnionis majoris
K. J. Boss, 1988
Page 121
fragmentis. Londini Gothorum [Goteborg], G Riurnian,
p. 22, 1 pi, 4 to. [not seen],
Strom, Hans [1726-97]. 1762. Physisk eg oeconomisk beskri-
velse over Fogderiert Sondmor. beliggende i Bergens stift
i Norge. 2 Deele, Soriie, [Rothteste], p [xvi], 570, [ii], -1
pis , 4to.
Swammerdam, Jan [1637-80]. 1737-38. Bybel der Natuure
door J. Swammerdam of Historie der Insecten, &c. —
J Swammerdamii . . Biblia Naturae; sive Historia Insec-
torum . . . Insertis numerosis rariorum naturae observa-
tionibus. Omnia lingua Batava . . . conscripta. Accedit
praefatio, in qua vitam auctoris descripsit H. Boerhaave
. . . Latinam versionem adscripsit H.D. Gaubius. Levdae,
Severin and van der An. 2 \ols., p 550, [56]; 551-910.' [36],
124, 53 pis,, folio.
Tournefort, Joseph Pittonde[1656-1708], 1717 Reiationd'un
Vovage du Le\ant . . Contenant I'histoire , . . de plusieurs
Isles de I'Archipel, de Constantinople, des Cotes de la Mer
Noire, de I'Armenie, de la Georgie. des frontieres de Perse
& de I'Asie Mineure . . Enrichie . . de plusieurs obser-
vation touchant I'Histoire naturelle. ([To which is prefixed
an] Eloge de M de Tournefort. Par M de Fontenelle.). 2
\'ols., illustrated Paris, 4to.
Tulpius, Nicolaus [1593-1674]. 1739. N. Tulpii . . . Obser-
vationes medicae. Editio sexta, prioribus emendatior &
auctior, cum oratione funebri . . . L. Wolzogeni, &c. Lug-
duni Batavorum, p. [viii], 51 [4], .392 [7], 18 pis., 1 portrait,
engraved title, 8vo, [not seen].
X'allisnieri, Antonio [1661-1730]. 1721. De'Corpi marini che
su' Monti si trovano; della loro origine . . . lettere critiche,
&. Venezia, Dom. Lovisa, p. [ix], 254 [10], 4 pis., 4to.
Wolfart, Peter [1675-1726]. 1719. Historiae naturalis Hassiae
inferioris pars prima, &c. Der Natur-Geschichte des Nie-
der-Fuerstenthums Hessen. Erster Theii. Latin & German.
Cassel, Harmes, p. 52, 26 pis. folio.
Worm, Ole [1588-1654]. 1655. Museum Wormianum. Seu
historia rerum rariorum. tarn naturalium, quam artifici-
alium, tarn domesticarum, quam exoticarum. quae Haf-
niae Danorum in aedibus authoris servantur. Variis & ac-
curatis iconibus illustrata Lugduni Batavorum, Elsevir, p.
[x], 389, 2 pis., text illustrated (139 figures), folio.
ACKNOWLEDGEMENTS
I wish to thank Dr. Elizabeth A. Shaw% Bibliographer
and Research Taxonomist in the Gray Herbarium, Mrs.
Geraldine C. Kaye of the Fariow Herbarium Library,
Ms. Barbara Callahan of the Harvard University Her-
barium, and Eva Jonas, Dana Fisher, Roxane Coombs,
and Mina Brand of the library of the Museum of Com-
parative Zoology. Mr. Richard I. Johnson also made avail-
able rare malacological volumes, and valuable criticisms
were offered by Mr. Richard E. Petit and two anonymous
reviewers.
LITERATURE CITED
Boss, Kenneth J. In preparation. The bibliographic sources
for Linnaeus's knowledge of mollusks; the authors, edi-
tions, and translations.
Catalogue of the Books. Manuscripts. Maps and Drawings of
the British Museum (Natural History) London. Hazell,
Watson and Viney, Ld., Vol. 1. A-D (1903), p. viii, 1-500;
Vol, II. E-K (1904). p. [vi], .501-1038; Vol. III. L-O (1910),
p. [vi], 1039-1494; Vol. IV. P-Sn (1913), p. [vi], 1495-
1956; Vol. V. So-Z (1915), p. [vi], 1957-2403; London.
William Clowes and Sons, Vol. VI. Supplement A-I (1922),
p [vi]. 1-512; ."Addenda and Corrigenda to Vols. I and II.
A-Hooker. p. 1-48; Oxford. Universitv Press. Vol. VII.
Supplement J-O (1933), p. [vi]. 513-968; Vol. VIII. Sup-
plement P-Z (1940), p. [vi], 969-1480.
Dance, S. Peter. 1967 Report on the Linnaean shell collection.
Proceedings of the Linnean Society of London 178(1):1-
24, pis. 1-10.
Dance, S. Peter. 1986. A history of shell collecting. Revised
edition of Shell Collecting. E. J. Brill-Dr. W. Backhuys,
Leiden, p. xv, 265, 32 pis., 17 text-figures.
Dodge, Henry. 1952-59. A historical review of the mollusks
of Linnaeus. Parts 1-7. Bulletin of the American Museum
of Natural Historv, 100(1): 1-264 (1952); 103(1): 1-134
(1953); 107(1):1-158 (1955); Ill(3):153-312 (1956); 113(2):
73-224 (1957); 116(2): 153-224 (1958); 118(5):207-258
(1959).
Engelmann, Wilhelm. 1846. Bibliotheca Historico-naturalis.
Verzeichniss der Bijcher iiber Naturgeschichte welche in
Deutschland, Scandinavien, Holland, England, Frank-
reich. Italien und Spanien in den Jarhren 1700-1846 er-
schienen sind. Erster Band. Bijcherkunde. Hiilfsmittel, all-
gemeine Schriften, vergleichende Anatomic und
Physiologic. Zoologie. Palaeontologie. Verlag von Wilhelm
Engelmann, Leipzig, p. vii, 786.
Hanley, S>lvanus C. T. 1855. Ipsa Linnaei Conchy lia. Wil-
liams and Norgate. London. 556 p.. pis. I-V.
Heimann. Willy. 1957. .\ catalogue of the works of Linnaeus.
Issued in commemoration of the 250th anniversary of the
birthday of Carolus Linnaeus 1707-1778. Sandbergs An-
tikvariatsforteckning. Nr. 12 Varen. 1957. Sandbergs Bok-
handel, Stockholm. 179 p.
Heller. John L. 1961. List. Loqu ' and List Mut.'. two puz-
zling Linnaean abbreviations Proceedings of the Linnean
Society of London 173:61-64, pis. 1-4.
Heller. John L. 1968. Zoological sources of Linnaeus. Journal
of the Society for the Bibliographv of Natural History 5(1):
76-77.
Heller, John L. 1979. Bibliotheca Zoologica Linnaeana. Sven-
ska Linnesallskapets arsskrift. argang 1978. Yearbook of
the Swedish Linnaeus Society. Commemorative Volume.
Uppsala, p. 240-264.
Heller, John L. 1983. Studies in Linnaean method and no-
menclature. Marburger Schriften zur Medizingeschichte,
(Verlag Peter Lang, Frankfurt am Main, Bern, New York. )
7(IX):328 p.
Keen. Angeline Mvra. 1983a. (January). On Linnaeus" book-
shelf. The Festivus (San Diego Shell Club) 15(1):5-15. 24
figs.
Keen. Angeline Myra. 1983b (August 30). On Linnaeus' book-
shelf [abstract]. .Annual Report of the Western Society of
Malacologists 15:15.
Linnaeus. Carolus [1707-78]. 1758. Caroli Linnaei . . . Sys-
tema Naturae per Regna tria Naturae, secundum classes,
ordines, genera, species, cum characteribus. differentiis,
synonymis, locis . . . Editio decima reformata. Impensis
Direct. Laurentii Salvii: Holmiae, Tom. 1. Animalia. p.
[iv], 823.
Linnaeus. Carolus [1707-78]. 1767. Caroli a Linne . . . Sys-
tema Naturae per Regna tria Naturae, secundum classes,
ordines. genera, species, cum characteribus. differentiis,
synonymis, locis . . . Editio duodecima, reformata. Impen-
Page 122
THE NAUTILUS, Vol. 102, No. 3
sis Direct. Laurentii Salvii: Holmiae, Tom. 1 Regnum
Animale. Pars II. Classis V-VI, Nomina Generica. Nomina
specierum propria: Nomina trivialia. Nomina trivialia
Papiliomim and Phalaenarum Synonyma Termini Artis.
Appendi.x Svnonvniorum. Addenda. Errata, p. 533-1327
[36].
Nissen, Glaus. 1969. Die Zoologische Buchillustration. Ihre
Bibliographie und Geschichte. Band I. Bibliographie. An-
ton Hiersemann, Stuttgart, p. 666.
Soulsby, B. H. 1933. A catalogue of the works of Linnaeus
(and publications more immediately relating thereto) pre-
served in the libraries of the British Museum (Bloomsbury)
and the British Museum (Natural History) (South Ken-
sington). Second Edition Trustees of the British Museum,
London, p. xi, 246; .\ddenda and Corrigenda, 68 p., 7
plates.
Wheeler, A. 1979. The sources of Linnaeus knowledge of
fishes. Svenska Linnesallskapets arsskrift. argang 1978.
Yearbook of the Swedish Linnaeus Society. Commemo-
rative Volume. Uppsala, p. 156-211.
THE NAUTILUS 102(3):123-124, 1988
Page 123
Frederick Benjamin Isely: Biographical Sketch and
Malacological Contributions
Mark E. Gordon
Department of Zoology
University of Arkansas
Fayettevi'lle, AR 72701, USA
Frederick B. Isely was known primarily for his pioneer-
ing research in orthopteran ecology (mainly Acrididae
and Tettigoniidae) and as an educator/administrator.
However, he contributed significantly to the basic knowl-
edge of unionacean biology and ecology. Isely began his
studies with freshwater mussels in the Chikaskia River
of north-central Oklahoma during 1906. These initial
observations soon expanded to include aspects of union-
acean ecology, growth, migration, and distributional sur-
vey of mussels within the Arkansas and Red river basins
of eastern Oklahoma. In these endeavors, he was finan-
cially aided through an appointment as a scientific as-
sistant with the U.S. Bureau of Fisheries during the sum-
mers of 1910-13. As a result, five manuscripts were
published between 1911 and 1931. Isely's innovative con-
cepts for ecological investigations are reflected by his
discussion in the 1914 growth paper of the use and value
of replicate samples and of the problems encountered
using mark/recapture techniques. One such problem was
predation by small bo\s interested in collecting the brass
tags used to mark the mussels. The distributional survey,
although funded by and conducted for the U.S. Bureau
of Fisheries, was delayed in publication due to govern-
ment reassessment of funding priorities — World War I.
This manuscript was published independently by Isely
after the war. His last malacological publication resulted
from the recovery of one of the "lost" growth study
specimens 15 years following the completion of that in-
vestigation. Further evidence for the value of replication
in research.
Frederick was born June 20, 1873 of Swiss parents
(Christian Isely and Elise Dubach) at Spring Grove farm
near Fairview, Brown County, Kansas. His early edu-
cation was gained locally, culminating with his gradu-
ation in 1894 from Hiawatha Academy (Hiawatha, Kan-
sas). He entered Fairmount College (now Wichita State
University) in its founding year 1895. He was quite active
in college activities (e.g., debate team, class president
1895-99) and sports (e.g., track, football) and was award-
ed a B.S. in 1899, the first 4-year graduating class of the
college. In 1909, he earned an M.S. from the University
of Chicago and continued that summer with further study
at the Marine Biological Laboratory. Later academic
endeavors were pursued during the summers of 1929
and 1931 at the University of Chicago and University of
Colorado, respectively.
His teaching career began with an appointment at
Franklin Rural School, Brown County, Kansas (1894-
95). Following his graduation from Fairmount College,
Figure 1. Frederick B. Isely during his Trinity University
years. Reproduced from a photograph given to the Department
of Biology, Trinity University by Mrs. F. B. Isely (courtesy of
H. D. Murray).
Page 124
THE NAUTILUS, Vol. 102, No. 3
he assumed the principalship of Central School, Hia-
watha (1899-1901) followed by stints as teacher of bi-
ology at Wichita High School (1901-06) and University
Preparatory School, Tonka wa, Oklahoma (1906-12).
Equipped \s ith his M.S., he served as professor of biolog\'
at Central College, Fayette, Missouri (1912-20) with so-
journs during the summers of 1915-17 as instructor of
biology at the L'niversity of Missouri. His administrative
career was initiated with an appointment as dean and
professor of biology at Ckilver-Stockton College, Canton,
Missouri (1920-22) followed by his assumption of the
same roles at Texas Women's College, Fort Worth (1922-
31). He returned to a strictK teaching position as pro-
fessor of biology at Trinity Universit\, Wa.xahachie and
later San Antonio, Texas (1931-47) where he developed
the majority of his orthopteran research. He also served
as department chairman from 1931 to 1946, retiring from
active teaching in 1946. In recognition of his achieve-
ments. Trinity University awarded him the Sc.D. honoris
causa upon his retirement.
Isely married Mary E. Nickerson of Clearwater, Kan-
sas, on May 8, 1901. They had four children and he
appears to have influenced his famiK , as he did his stu-
dents; biologists continue in the famih to date. In spite
of a heavy teaching load, administrative duties, and re-
search, he was active in community organizations
throughout his life and still managed time for personal
pleasures {e.g.. camping, gardening, keeping up with the
latest episodes of Joe Palooka and Little Orphan Annie).
Isely was a well-loved and enthusiastic instructor in ad-
dition to being an active and respected scientist. He
received grants-in-aid for his orthopteran research from
the National Research Council (1935, 1936) and the
American Philosophical Society (1937, 1941). He was a
fellow and founding member of the Oklahoma Academy
of Science (secretar\ 1901-12) and the North Texas Bi-
ological Society (president 1924-27); a fellow of AAAS
and the Texas Academy of Science (president 1938); and
a member of the Society of American Zoologists, Eco-
logical Society of America, Entomological Society of
America, Kansas Academy of Science, Texas Entomo-
logical Society, and Texas State Teachers Association.
Frederick B. Isely died December 30, 1947 still active
in research. Additional information may be acquired
from two memoria (Alexander, 1949; Geiser, 1949).
Recently, a remnant of Isely s malacological collection
was rediscovered at the Stovall Museum, University of
Oklahoma (Shepard, 1982). About 450 specimens of
unionids, representing 37 species remain. Some inade-
quately labeled lots may also be referable to Isely. Thirty-
nine specimens including seven species represent vouch-
ers for the 1914 growth paper. Fourteen detached brass
tags are also present. The remainder of his collection is
from the 1925 distribution sur\e\. Little else of Isely's
malacological material, other than some papers in the
archives of the Biology Department at Trinity University
(H. D. Murra\, personal communication), remain.
A small malacological collection of his brother, Dwight
IseK , was redisco\ered at Wichita State L'niversity (Met-
calf and Distler, 1984). Further information pertaining
to D. Isely may be obtained in Miner (1976) and Metcalf
and Distler (1984).
I thank the following people for their kind assistance
in providing data for this note: Mrs. Katherine Isely
McGuire and Drs. Douglas G. Alexander, Harle\ P.
Brown, Donald A. Distler, Duane Isely, Charles G. Lin-
coln (deceased), Harold D. Murra\-, Robert W. Pennak,
William D. Shepard, and Gary D. Snell.
LITERATURE CITED
Alexander, G. 1949. Frederick B. Isely (1873-1947), Ento-
mology News 60:29-30.
Geiser, S. W. 1949. Proceedings of the forty-fifth annual
meeting of the American Society of Zoologists: 3. Memorial
resolutions, Frederick B. Isely 1873-1947. Anatomical
Record 103:233-234.
Metcalf, A. L. and D. A. Distler. 1984. Gastropods collected
from eastern Oklahoma by Dwight Isely in 1911. The
Nautilus 98:135-137.
Miner, F. 1976, Dwight Isely 1887-1974 Journal of Economic
Entomology 69:289-290.
Shepard, W. D. 1982. Rediscovery of a portion of the Isely
unionid collection. The Nautilus 96:8.
MALACOLOGICAL BIBLIOGRAPHY OF
FREDERICK B. ISELY
1911. Preliminary note of the ecolog> of the early juvenile life
of the Unionidae. Biological Bulletin 20:77-80.
1914. Experimental stud> of the growth and migration of fresh-
water mussels. U.S. Bureau of Fisheries Document no.
792, 26 p.
1914. Mussel streams of eastern Oklahoma. U.S. Bureau of
Fisheries Economic Circular no. 9, 6 p
1925. The fresh-water mussel fauna of eastern Oklahoma. Pro-
ceedings of the Oklahoma Academy of Science 4:43-
118.
1931. A fifteen vear growth record in fresh-water mussels.
Ecology 12:616-619,
THE NAUTILUS 102(3): 125-126, 1988
Page 125
Rediscovery of Planorhella magnifica (Pilsbry) in
Southeastern North Carohna
William F. Adams
Environmental Resources Branch
U.S. Armv Corps of Engineers
P.O. Bo.x 1890
Wilmington, NC 28402, USA
Andrew G. Gerberich
Division of Fishes
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560, USA
Early in this century, Planorbis magnificus was de-
scribed by Pilsbry (1903) from the lower Cape Fear River
region of North Carolina. The type localitv for the species
was given simply as "lower Cape Fear River". Bartsch
(1908) correctly surmised that the species is an inhabitant
of lentic environments and found it living in Greenfield
Lake, a manmade impoundment lying to the south of
what were then the city limits of Wilmington, NC, as
well as in Orton Pond, then called Sprunt's Pond, ap-
proximately 16 km south of Wilmington. Baker (1945)
figured and studied portions of the anatomy of specimens
collected b\' Bartsch and concluded that P. magnificus
should be reassigned to Planorhella. This ta.xon repre-
sents the largest known planorbid.
Planorhella magnifica has not been reported since the
accounts b\ Pilsbry and Bartsch and has been considered
extinct by some (Opler, 1976; Imlay, 1977; Palmer, 1985).
Fuller (1977) hypothesized that the species may still sur-
vive in Orton Pond, a manmade pond similar to Green-
field Lake in age and physiography, based on an obser-
vation of large planorbid egg masses there by J. P. E.
Morrison.
The Greenfield Lake watershed has been almost totally
developed since Planorhella magnifica was described,
and now undergoes intensive management for control of
nuisance algae. Management measures include appli-
cation of algicides during the growing season and oc-
casional drawdown during winter months. Fuller (1977)
mentioned fruitless attempts by himself and others to
find Planorhella magnifica in Greenfield Lake. Our in-
vestigations of that lake indicate that it no longer provides
a suitable habitat for this species.
During July 1986, Planorhella magnifica was seren-
dipitously rediscovered in Orton Pond, Brunswick Coun-
ty, NC, while collecting water samples. Three living
specimens were obtained in 20 minutes. Four empty
shells were also collected from the drift line on the shore.
During another water-sampling trip in January 1987, an
attempt was made to collect live specimens for captive
propagation. This effort was unsuccessful as much of the
aquatic vegetation had died back. However, two more
empty shells were found washed up on the shore.
Our samples of Planorhella magnifica vary consid-
erably in size. The maximum shell diameters of the spec-
imens collected alive are 35.5, 21.5, and 16.3 mm. Vouch-
er specimens from our collection have been deposited in
the collections of the North Carolina State Museum of
Natural History (NCSM P468-P471) and the National
Museum of Natural History (USNM 857935). Because of
restricted access, additional trips to Orton Pond have not
been made.
Live specimens of Planorhella magnifica were found
on the stems and undersides of the floating leaves of
Spatterdock, Niiphar liiteum (Sibthorp & Smith, 1809),
and Fragrant Waterlily, Nymphaea odorata (Aiton, 1789).
Water depth where living specimens were collected was
approximately 1 meter, and the bottom substrate was
organic. Orton Pond closely matches Bartsch's (1908)
description of habitat suitable to P. magnifica in Green-
field Lake.
Like Greenfield Lake, Orton Pond is a manmade im-
poundment. Both were created early in the last century
to serve as a fresh-water source for rice agriculture. Orton
Pond exemplifies a type of lentic waterbody unique in
southeastern North Carolina. Although blackwater lakes
and ponds in the region typically are acidic, Orton Pond's
waters have a circumneutral pH, ranging from 6.2
through 7.9 (Smock and Lenat, 1978). We suspect that
Orton Pond may have a direct connection to waters of
the Castle Hayne aquifer, an Eocene limestone deposit.
The molluscan fauna of Lake Waccammaw (Columbus
County, NC), the only other sizable circumneutral-pH
waterbody in the region, has been investigated frequent-
ly (Fuller et al, 1976; Porter, 1985), but Planorhella
magnifica has not been found in that system. Lake Wac-
camavv differs from Orton Pond in being much larger,
a natural waterbody, and is not a blackwater system.
The owners of Orton Pond undertake little manage-
ment of the pond and manage the lands that surround
it for timber and wildlife. With a continuation of this
type of management, the near future for Planorhella
magnifica seems secure. However, not all of the pond s
watershed is protected in this way. Therefore, potentially
adverse developments could occur quite some distance
from the pond.
Because of the apparent uniqueness of the waters of
Page 126
THE NAUTILUS, Vol. 102, No. 3
Figure 1 . Shells of Planorbella magnifica collected alive from
Orton Pond, Brunswick County, North Carolina, July 1986
(apprcximately natural size).
Orton Pond and absence of Planorbella magnifica in
Greenfield Lake and Lake W'accamaw, it appears that
the species is extremely restricted in distribution and can
be found today only in Orton Pond. We believe that any
changes in the quality of the surface waters in Orton
Pond's drainage basin or in its underlying groundwater
regime may have a deleterious effect on P. magnifica.
Due to the uncertainty of the future of the pond, an in-
depth study of the autecology of the species is needed
so that potential conservation measures can be developed
and implemented should they prove necessary.
LITERATURE CITED
Baker, F. C. 1945. The molluscan family Planorbidae. Uni-
versity of Illinois Press, Urbana, p. 1-530.
Bartsch, P. 1908. Notes on the fresh-water mollusk Planorbis
magnificus and descriptions of two new forms of the same
genus from the southern states. Proceedings of the United
States National Museum 33:697-700.
Fuller, S. L. H. 1977 Freshwater and terrestrial mollusks. /n:
Cooper. J. E . S S Robinson, and J B Funderburg (eds. ).
Endangered and threatened plants and animals of North
Carolina. North Carolina State Museum of Natural His-
tory, Raleigh, p. 143-194.
Fuller, S. L. H., M. J. Imlay, and J. D. Williams. 1976. En-
dangered or threatened fresh-water mussels (Mollusca:
Bivalvia; Unionidae) of the Waccamaw River basin of the
Carolinas. Association of Southeastern Biologists Bulletin
23:60.
Imlay, M. J. 1977. Competing for survival Water Spectrum
9:7-14.
Opler, P. A. 1976. The parade of passing species: a survey of
extinctions in the U.S. Science Teacher 43(9):30-34.
Palmer, S, 1985. Some extinct molluscs of the U.S.A. .Atala
13(l):l-7.
Pilsbry, H. A. 1903. The greatest American Planorlns. The
Nautilus 17:75-76.
Porter, H. J. 1985. Rare and endangered fauna of Lake Wac-
camaw, North Carolina watershed s\ stem, \'ols. 1 and 2.
North Carolina Endangered Species Restoration: Job Title
No. VI-7. North Carolina Wildlife Resources Commission,
Raleigh, p. 1-358.
Smock, L. A. and D. R. Lenat. 1978. Preliminary stud\ of
the effects of salt water intrusion on the macrobenthic
community of Orton Pond, North Carolina. U.S. Army
Engineer District, Wilmington, p. 1-48.
THE NAUTILUS 102(3):127-128, 1988
Page 127
Bite by Octopus joubini: A Case Report
Donald M. McKinstry
Division of Science, Engineering, and
Technology
The Pennsylvania State University
at Erie
The Behrend College
Erie, PA 16563-1200, USA
While octopuses are capable of delivering toxic bites,
human case reports are scarce (Halstead, 1978). This
report describes a bite by Joubin's octopus (Octopus jou-
bini Robson, 1929) in a 12-year-old boy. The incident
occurred on Sanibel Island, Florida, on January 2, 1987
at about 0900. A storm had deposited a variety of marine
animals on shore from the Gulf of Mexico. Not realizing
that octopuses are venomous, the boy, Patrick Reed
McKinstry, picked up a small specimen and was prompt-
ly bitten on the dorsal surface of the right thumb prox-
imal to the interphalangeal joint. He pulled the octopus
away from his thumb within 10 seconds. The actual bite,
described as a "sharp pinch", was quickly followed by
a burning sensation of greater intensit% than a bee-sting.
A tiny red spot surrounded by a white circular area 6
mm in diameter was then noticed. Bleeding was minimal,
a few small drops. Over the next se\ eral hours the thumb
became moderately swollen and felt sore, stiff, and, at
times, somewhat numb. These symptoms, to a lesser de-
gree, extended to the wrist. Medical attention was not
sought because only local effects, which gradually di-
minished by day one, were noticed. By day three, the
thumb was slightly swollen, reddened, and sore, and a
tiny hole was noticed at the site. Over the next several
days, the thumb became itchy and was treated with 3%
hydrogen peroxide solution. This was the first treatment
of the bite. By day eight, the swelling and redness had
disappeared but some soreness in the metacarpopha-
langeal joint was experienced. The hole had enlarged to
2 mm in diameter and displayed a slightly irregular
border. During the period up to day 12, the hole became
darker and a red circular area of up to 7 mm in diameter
developed around it. Itching continued but soreness in
the joint area decreased. On day 12, a small amount of
clear yellow fluid oozed from the hole. The area was
then treated with hydrogen peroxide solution and anti-
biotic ointment. The signs and symptoms gradually di-
minished, and by day 32 the hole was replaced by a
slightly indented circular scar 1 mm in diameter, sur-
rounded by a slightly darkened firm area 6 mm in di-
ameter. By da\- 100, onK' the indented scar was evident
and the wound was considered completeK healed. .A
shiny, hard circular scar, 1 mm in diameter, was still
evident by 1 year. While in excellent health, Patrick is
allergic to hymenopteran venoms and to penicillin. How-
ever, this was his first exposure to octopus venom, thus,
an allergic reaction to explain the bite effects was dis-
counted. Secondary bacterial infection could have con-
tributed to the effects experienced, however, signs of
infection, e.g., pus, lymphangitis, and fever, were not
observed.
The octopus died after about 2 hours in a pail of sea
water and was preserved in 70% isopropyl alcohol. It was
identified as a nearly reproductively mature male spec-
imen of O. joubini by Dr. Ronald Toll. This species, the
smallest of the western Atlantic octopuses, ranges from
southern Florida to the Bahamas and the West Indies.
Specimens are often washed ashore by storms on the Gulf
coast of Florida (Meinkoth, 1981). The dorsal surface of
the specimen is deep tan, while the arms and ventral
surface are light tan. The mantle is 3.2 cm in length. I
have donated this specimen to Dr. Toll s collection.
Two accounts of envenomation following the bite of
O. joubini were found. In the first (Anonymous, 1965),
a physician was bitten on the finger while handling a
specimen washed ashore on Sanibel Island, Florida. He
experienced a "sharp" bite with a little bleeding, quickly
followed by '"the most excruciating pain which rapidly
became almost intolerable." Within a few hours, the
finger became very swollen, hard, and deep red. By the
next day, the inflammation and pain had extended to
the hand. Later, some white discharge was noted from
the bite wound. The swelling and hardness gradually
subsided over a period of 2 months. The only systemic
effect was a mildly elevated temperature. He noted that
"there was never any evidence of bacterial invasion or
infection, only an extreme degree of cellulitis. '
The second case (Wittich, 1968), occurred on a re-
search vessel off Egmont Key, Florida. A Florida Board
of Conservation worker was bitten on the back of the
hand while sorting marine specimens. In this case, '"a
sharp, piercing sensation " was felt, and a "severe pain"
extended to the upper arm. The bite wound bled slightK
and almost instantK became surrounded by a "'pure white
Page 128
THE NAUTILUS, Vol. 102, No. 3
welt of about 25 mm diameter." In the following hour,
the victim experienced pain and sw elling around the bite,
accompanied by nausea, headache, and fever. Improve-
ment gradually occurred, but after 3 days the area was
still sensitive and swollen. The swelling persisted, and
some itching and a serous discharge from the wound
occurred. After 1 month, a 2 mm wound surrounded by
a 6 mm red periphery was present. The wound then
healed quickK.
In the case I report, the bite effects were not as severe
as those noted above even though the victim was a child
with less tissue mass to dilute the venom. However, the
amount of venom delivered could have been minimal.
ACKNOWLEDGEMENTS
I am grateful to the following for their helpful assistance
and suggestions: Ronald Toll, The L'niversity of the South,
Sewanee, TN; Sherman A. Minton, Jr , Indiana Univer-
sit\ , School of Medicine, Indianapolis, IN; Harry N. Cun-
ningham, Jr., Kath\' Mauro, and Robert Rose, all of The
Pennsylvania State University at Erie, The Behrend Col-
lege, Erie, PA. A special thanks is due Patrick R. Mc-
Kinstrv for his account of the octopus bite.
LITERATURE CITED
.\nonymous. 1965. Florida octopus bite. Sea Secrets 9(7):5.
Halstead, B. W 1978. Poisonous and venomous marine
animals of the world The Darwin Press, Princeton, NJ,
1043 p.
Meinkoth, N. A. 1981. The Audubon Society field guide to
North American seashore creatures. .\if red .\. Knopf, New
York, NY, 799 p.
Wittich, A. C. 1968. Account of an octopus bite. Florida
Academy of Science Quarterly Journal 29l4):265-266.
THE NAUTILUS 102(3):129, 1988
Page 129
Occurrence of Mites in Mexican Land Snails
0§car J. Polaco
Departamento de Prehistoria
Institute Nacional de Antropologia
e Historia
Moneda #16
Mexico 06060, Mexico
Wolfgang Mendl
Lab. Acarologia
Departamento Zoologia
Escuela Nacional de
Ciencias Biologieas
Apartado Postal 42186
Mexico 11340, Mexico
To date there has only been a single report (Baker, 1945)
of mites infecting the mantle cavity of a land snail from
Mexico (Helix pomatia Linne, 1758, an introduced
species). The mite was identified as Riccardoella oude-
mansi (Thor, 1932), however, the mite shown by Baker
(1945) is similar to the figure published by Fain and Van
Goethem (1986) for Riccardoella limacum (Schrank,
1776) (Prostigmata: Ereynetidae). This mite has been
studied by several European authors (Turk & Phillips,
1946; Baker, 1970) and reported as cosmopolitan.
In this note, we report findings of mites parasitizing
the mantle cavities of two species of land snails of the
famiK Bulimulidae and one species referable to the fam-
ily Helicidae. The bulimulid Bidimulus unicolor (Sow-
erby, 1833), collected in Tomas Garrido, in the state of
Quintana Roo, was parasitized by Riccardoella limacum.
with one protonymph and one deutonvmph collected
from nearly 50 specimens of the snail. Of 79 specimens
of Rhabdotus alternatus (Say, 1830), another bulimulid
collected in Xicotencatl in the state of Tamaulipas, that
were examined, one contained three mites of the genus
Boydaia (Ereynetidae) and another contained a single
specimen of Eupodes voxencollinus (Sig Thor, 1934).
This is the first report of the last two mite species on
land snails.
Examination of other land snail species of the families
Helicidae, Achatinidae, Succinidae, Oleanicidae, and
Polygyridae revealed no evidence of parasitism by mites,
with the exception of a single specimen of Helix aspersa
(MiJller, 1774), an introduced species collected in Mexico
City, which was infected with two specimens of E. vox-
encollinus. As no mites were found in any Mexican land
snails from the above families, the family Bulimulidae
remains the only new record of a host for mites.
The examined snails are deposited in the Mollusk Col-
lection of the Departamento de Prehistoria, I.N.A.H.,
and the mites in the Acarology Collection of the Labo-
ratorio de Acarologia of the E.N.C.B.
We acknowledge two anonymous reviewers for their
critical comments to improve this paper.
LITERATURE CITED
Baker, E. W. 1945. Five mites of the family Ereynetidae
from Mexico. Journal of the Washington Academy of Sci-
ences .35:16-19.
Baker, R. A. 1970, The food of Riccardoella limacuni (Schrank)
(Acari: Trombidiformes) and its relationships with pul-
monate mollusks. Journal of Natural History 4:521-530.
Fain, A. and J. L, VanGoethem. 1986. Les acariens du genre
Riccardoella Berlese, 1923, parasites du poumon de Mol-
lusques gasteropodes pulmones terrestres. Acarologia 27(2):
125-140.
Turk, F. A. and S. M. Phillips. 1946. A monograph of the
slug mite Riccardoella limacum (Schrank). Proceedings of
the Zoological Society of London 115:448-472.
THE NAUTILUS 102{3):130, 1988
Page 130
Axelella, New Name for Olssonella Petit, 1970, a Preoccupied
Taxon (Mollusca: Cancellariacea)
Richard E. Petit
Research Associate
Department of Invertebrate Zoology
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560, USA
The cancellariid genus Olssonella Petit, 1970, was pro-
posed for a rather compact and well-defined group of
species from the Later Tertiary and Recent faunas of
the Americas. An Eastern .Atlantic species was subse-
quently placed in this genus by Bouchet and Waren
(1985;263), and the morpholog\ of the type species was
published b\ Harasew\ch and Petit (1984). The ta.xon
has been used b\ a number oi other authors since it was
proposed. Unfortunately, it has just been noted that
Olssonella is preoccupied, and a replacement name is
here proposed.
Axelella, new name for Olssonella Petit, 1970:83.
Not Olssonella Glibert & \'an de Poel, 1967:121.
Type species, by original designation of Olssonella
Petit, 1970, Cancellaria smithii Dall, 1888, Recent,
Western .\tlantic.
This new name honors the late A.xel \. Olsson, as did
the name it replaces, and the name which makes the
replacement necessary.
Appreciation is expressed to Dr. Philippe Bouchet,
Museum National d'Histoire Naturelle, Paris, for calling
the prior use of Olssonella to m\ attention.
LITERATURE CITED
Bouchet, P. and A. Waren. 1985. Revision of the Northeast
Atlantic bathval and ab\ssal Neogastropoda excluding
Turridae (Mollusca: Gastropoda). Bollettino Malacologico,
Supplemento 1:121-296
Dall, W. H. 1888. Mollusks. In: Alexander Agassiz, Three
Cruises of the United States Coast and Geodetic Survey
Steamer Blake 2:62-75.
Glibert, M. and L. Van de Poel. 1967, Les Bivalvia fossiles
du Cenozoique etranger des collections de I'lnstitut Royal
des Sciences Naturelles de Belgique. \". Oligodontina. In-
stitut Royal des Sciences Naturelles de Belgique, Memoires
(2nd ser.') 83:1-152.
Harasew ych, M. G. and R. E. Petit. 1984 Notes on the mor-
phology of Olssonella smithii (Gastropoda: Cancellari-
idae). The Nautilus 98(l):37-44.
Petit, R. E. 1970. Notes on Cancellariidae (Mollusca: Gastrop-
oda) — II. Tulane Studies in Geologv and Paleontology 8(2):
83-88.
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Volume 102, Number 4
December 21, 1988
ISSN 0028-1344
A quarterly devoted
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Marine Biological Laboratory
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Division of Mollusks
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THEt^NAUTILUS
CONTENTS
Volume 102, Number 4
December 21, 1988
ISSN 0028-1344
Nidia S. Romer
Donald R. Moore
A new species of Alvania (Rissoidae) from the West Indian
region
131
Charles N. D'Asaro
Micromorphology of neogastropod egg capsules
134
Philippe Bouehet
Two new species of Metula (Gastropoda: Buccinidae) with
a description of the radula of the genus
149
Kurt Auffenberg
Harry G. Lee
A new species of intertidal Terebra from Brazil
154
Douglas C. Smith
Notes on the biology and morphology of Margaritifera
hemheli (Conrad 1838) (Unionacea: Marearitiferidae)
159
Alan R. Kabat
Richard E. Petit
The two printings of J. F. Gmelin's Systema Naturae. 13th
Edition (1788-96)
164
Marine Biological Laboratory
LIBRARY
JAN 3 1989
Woods Hole, Mass.
THE NAUTILUS 102(4):131-133, 1988
Page 131
A New Species of Alvania (Rissoidae) from the
West Indian Region
Nidia S. Ronier
Donald R. Moore
Rosenstit'l School of Marine
and Atmospheric Science
l'niversit> of Miami
4600 Bickenbacker Causeway
Miami, FL 33149, USA
ABSTRACT
Alvania (Alvania) colombiana new species was found from 45
to 261 m (24 to 143 fms) depths off the Caribbean coast of
Colombia to South Florida, the Gulf of Mexico, and Puerto
Rico. Alvania colombiana differs from Rissoa xanthias (Wat-
son, 1886), in having a protoconch with a single spiral zig-zag
line of tiny nodules just above the suture, and by its smaller
shell size. Alvania colombiana also differs from Rissoa precip-
itata (Dall, 1889), by its protoconch and size.
INTRODUCTION
Exploration of the deep sea a century ago was the equiv-
alent of the space age today. The famous "Challenger"
expedition sampled two offshore stations in the tropical
western Atlantic in which small mollusks were numerous.
These were described by Watson (1886). Among the
Rissoidae were two species, Rissoa pyrrhias (Watson,
1886) and Rissoa xanthias (Watson, 1886), that were
similar in size and shape. Dall (1889), reporting on the
"Blake" material, described two more species of Rissoa,
Rissoa acuticostata and Rissoa precipitata. Rissoa acu-
ticostata was similar to Rissoa xanthias. and finally Dall
(1927) acknowledged that with more specimens it was
impossible to separate the material into two species. Ris-
soa precipitata, on the other hand, is known principally
from the original description and illustration.
There is another small western Atlantic rissoid which
at first glance appears similar to the illustration of Rissoa
precipitata. However, there are a number of features
that differentiate them. Size is one: the small species has
an average adult size of 1.2 mm while Rissoa precipitata
(holotype) has an adult size of 3.92 mm. Protoconch is
the other: the small species has a projecting protoconch
while Rissoa precipitata has a depressed protoconch. One
of us (D.R.M) had obtained specimens of the small species
during the MAFLA (Mississippi, Alabama, and Florida)
study (1974 to 1975) in the eastern Gulf of Mexico. This
small species was first reported by one of us (N.S.R.) as
Alvania sp. 1 (see literature cited; Rodriguez, 1983).
ABBREVIATIONS
USNM = National Museum of Natural History, Smith-
sonian Institution.
FSBS I = Florida Department of Natural Resources,
Marine Research Laboratory, St Petersburg.
MCZ = Museum of Comparative Zoology, Harvard Uni-
versity.
UMML = University of Miami Marine Laboratory.
CNMS = Colombian Natural Museum of Science, Na-
tional University, Bogota
MATERIALS AND METHODS
All the specimens to be photographed in the SEM were
cleaned in 100% sodium hypochlorite for 2 minutes, then
rinsed in distilled water twice. If the specimens still
showed residual material (e.g., sand grains), they were
sonicated for 60 seconds. Finally, the specimens were air
dried and put on a SEM stub with double-sided Scotch
tape.
Alvania (Alvania) colombiana new species
(figures 1, 2)
Description: Shell 1.0 to 1.3 mm in length. Protoconch
multispiral, glossy, light brown, with 3'/2 whorls that are
sculptured with 1 spiral zig-zag line of tiny nodules just
above the suture and 1 undulating line of the same nod-
ules in the middle of the whorl. Nodules irregularly ar-
ranged over entire protoconch. Teleoconch of approxi-
mately 2 whorls, with numerous axial ribs. Adult shell
translucent, very light brown in color.
Axial ribs slightly curved, 26 ribs on the body whorl
fading out on the base of the shell. Body whorl with a
narrow spiral groove just below the suture and another
7 to 9 grooves between the periphery of the final whorl
and the base of the shell. Varix terminal with a narrow
extension forming outer lip of semilunate aperture. Some
specimens weakly umbilicate.
Page 132
THE NAUTILUS, Vol. 102, No. 4
Figures I, 2. Alvania colomhiana new species. I. Hololype, USNM miriiber 8.59339, 1.2S mm in length. 2. Protoconch, 280 x.
Figures 3, 4. Rissoa xanthias (Watson, 1886), from off Miami, in the junior author's collection, 2.26 mm in length. 4. Protoconch,
220 X.
Type locality: Off the west coast of Florida at 29°35'N,
87°20'06"W, depth 107.3 m.
Holotype: USNM number 859339. Length, 1.28 mm.
W'idtii, U.78 mm.
Paratypes: One from off the west coast of Florida at
29°35'N, 87°20'06"W, depth 107.3 m, USNM number
859340; 2 from off the west coast of Florida at 28°24'N,
85°15'06"W, depth 164.6 m, FSBC I number 33113; and
off the west coast of Florida at 29°49'30"N, 86°25'30"W,
depth 82.3 m, FSBC 1 number 33114; 2 from off Puerto
Rico at 17°53'24"N, 66°35'10"W, depth 221 m, MCZ
number 297220; and off Miami at 25°47'N, 80°01'30"W,
depth 137 m, MCZ number 297219; 2 from off Miami
N. S. Romer and D. R. Moore 1988
Page 133
Figure 5. Drawing of Alvania precipitata (Dall, 1889), ho-
lotype, MCZ 7470, 3.92 mm in length.
at 25°47'N, 80°01'30"W, depth 137 m, UMML number
8349; and off Miami at 25°46'30"N, 80°00'08"W, depth
76.81 m, UMML number 8350; 2 from off the Caribbean
coast of Colombia at 09°52'35"N, 75°47'25"W, depth 72
m, CNMS number ICN-MHN(MO)522; and off Miami
at 25°46'30"N, 80°00'08"W, depth 76.81 m, CNMS num-
ber ICN-MHN(MO)523.
Distribution: This species was found in 31 samples from
off the Caribbean coast of Colombia collected between
1979 and 1981. The specimens were found at depths
ranging from 45 m to 261 m, but were most abundant
at depths between 65 m and 160 m. Other specimens of
this species were found from off the west coast of Florida,
off southern Puerto Rico, and the Straits of Florida east
of Miami. The bottom in almost all cases consisted of
sandy mud. The species is probabK' found in these depths
throughout the Caribbean, the southern and eastern Gulf
of Mexico, and the Bahamas. All the specimens were
found dead.
Discussion: Alvania colombiana appears to be a com-
mon widespread species throughout the Caribbean and
adjacent areas. This is a very small compact species that
should not be confused with any other in its range. Some
features, however, are lacking or lost on the protoconch
of some specimens. These include the fine nodules dis-
tributed all over the protoconch, and the undulating line
of nodules on the periphery. These features are seen only
under high magnification, so are difficult to observe for
routine identification. Details of the teleoconch sculp-
ture, however, should be easy to make out under low
power, even in a somewhat worn specimen.
Alvania colombiana has been confused with Rissoa
precipitata (Dall, 1889) (figure 5). This was due to the
fact that specimens of Rissoa precipitata were not avail-
able for comparison. The holotypc and only known spec-
imen, MCZ 7470, is a thin shell v\itli no terminal varix.
In fact, the sculpture fades away almost completely on
the last half whorl (except for the sub-sutural tubercles).
The protoconch is of the form seen in gastropods without
a planktonic larval stage: large, rounded, unsculptured,
and with a smooth transition from protoconch to teleo-
conch. The type locality is Yucatan Strait at a depth of
640 fathoms (not 670 as is given by Dall, 1889:280).
Another similar species is Rissoa xanthias (Watson,
1886) (figures 3 and 4). It, however, is about twice the
size, has just over half as many axial ribs, and has a
different and very distinctive protoconch. In this species
the protoconch has 3'/2 whorls, in which the sutures are
deep and ornamented with a line of spirally arranged
vertical pustules. Just above the suture there is a wide
canal whose edges are formed by an undulating line with
vertical pustules equally spaced and directed downward.
Each canal also has in the middle dots and small nodules
randomly distributed. Watson reported it from Chal-
lenger Sta. 24 off Culebra Island, 18°38'30"N, 65°05'30"W,
depth 715 m, and from Sta. 122, 9°10'S, 34°49'W to
34''53'W, depth 640 m.
Finally, Rissoa pyrrhias (Watson, 1886) is another sim-
ilar species. It is slightly larger than Rissoa xanthias, and
its spiral grooves are not confined to the base of the shell.
According to Watson (1886), the protoconch has 2'/2 whorls
which are scored with coarse but feeble spiral threads.
ACKNOWLEDGEMENTS
We are very thankful to Dr. Patricia Blackwelder and
her assistant team at the SEM lab for their collaboration
in the use of and picture taking on the SEM. We also
thank Dr. Kenneth J. Boss for the loan of the holotype
of Rissoa precipitata.
LITERATURE CITED
Dall, W. H. 1889. Reports on the results of dredging ... in
the Gulf of Mexico (1877-78) and in the Caribbean Sea
(1879-80) by the U.S. Coast Survey Steamer "Blake" . . .
29. Report on the Moilusca, part II. Gastropoda and Sca-
phopoda Bulletin of the Museum of Comparative Zoology
18:1-492. pis. 10-40.
Dall, W. H. 1927 Small shells from dredgings off the southeast
coast of the United States b\ the United States Fisheries
Steamer "Albatross" in 1885 and 1886. Proceedings of the
L'nited States National Museum 70(2667);1-134.
Rodriguez, (Romer) N. S. 1983. Micromoluscos recientes de
la plataforma Caribe Colombiana entre Bocas de Ceniza
e Isla Fuerte. Thesis, Universidad Jorge Tadeo Lozano,
247 p.
Watson, R. B. 1886. Report on the Scaphopoda and Gaster-
opoda collected by H. M. S. Challenger during the years
1873-1876. Voyage of H. M. S. Challenger, Zoology 15:
1-675; appendix A, 677-680; geographical distribution,
691-722; index, 723-756; pis. 1-50.
THE NAUTILUS 102(4):134-148, 1988
Page 134
Micromorphology of Neogastropod Egg Capsules
Charles N. D'Asaro
Department of Biology
L iii\ersit\ of West Florida
Pensacola, FL 32514, USA
ABSTRACT
Egg capsule micromorphology of eight species of neogastropods
{Chicoreus florifer dileclu.s. PhijHonotus pomum. Cantharus
inuUangiilus, C. cancellariti.s. Conus floridanus floridensis, C.
ju.spidcus stearnsi. Cranulina ovuliforniis. and Marginella au-
reocincta) was studied with light microscopy hy examining
very thick, toluidine blue stained sections Laminae exposed
hy fracturing the sections, and their reactions to the stain,
provided the characters used to describe micromorphology.
The results showed that muricaceans and buccinaceans have
complex but microstructurally similar egg capsules, while co-
nids and marginellids have egg capsules with taxonomically
distinct rnicrostructural characteristics. In the Muricacea and
Buccinacea, four structural laminae of similar origin and func-
tion are usually present in the capsule wall, the second from
the outermost having the most complex pattern of fibers and
the greatest thickness. The third outermost lamina is continuous
with one or more components sealing the escape aperture. In
some buccinaceans, only a trace of the third lamina exists in
the wall. Four layers, including one or two mucoid plugs, close
the escape aperture. Conid egg capsules differ in that they
include only three structural laminae in the capsule wall and
three layers, including a riuicoid plug, sealing the escape ap-
erture. Microstructurc and fiber pattern in the middle lamina
are probably unique to this family. Egg capsules of marginellids
have a distinctive thin, dense wall not separated into laminae
and lack an escape aperture closed by a mucoid plug. There
is a preformed suture that fractures at hatching in the wall of
most marginellid egg capsules.
Ke\i words: Reproduction; egg capsules; Neogastropoda; Chi-
coreus. Phyllonotiis. Cantharus, Conns, Cranulina: Margi-
ncllti
INTRODUCTION
Encapsulation of early ontogenetic stages is typical of
higher gastropods, especially neogastropods. The highly
refractory, layered envelopes of carbohydrate and pro-
tein are structurally and chemicalK complex (Hunt, 1971;
Flower, 1973; Goldsmith et ai, 1978; Gruber, 1982; Sul-
livan and Maugel, 1984). Encapsulation and the for-
mation of egg masses provides protection and has con-
siderable survival value (Tamarin and C^arriker, 1967;
Pechenik, 1979, 1983; Abe, 1983; Lord, 1986). Although
there are thousands of neogastropod species, it still has
not been clearly established how the egg capsules are
formed in the oviduct and ventral pedal gland. Nor is it
known if a common capsular microstructurc exists in
various neogastropod taxa.
Macromorphology of the often species-specific egg
capsules and egg masses produced by neogastropods has
been studied more frequently than any other aspect of
neogastropod egg encapsulation. This was illustrated in
the review of prosobranch reproduction by Webber
(1977), in a more recent review of egg encapsulation for
all mollusks by Pechenik (1986), and in descriptive re-
ports on the external structure of neogastropod egg cap-
sules by Bandel (1975. 1976a,b,c, 1982) and D'Asaro
(1986a).
In early reports on neogastropod capsule-wall micro-
morphology (e.g., .\nkel, 1937; Hancock, 1956), three or
four laminae were described, including specific patterns
of fibers. However, it was not established if these are
structural relationships common to the species studied
or to higher taxa. Fretter (1941) provided for Nttcella
lapillus (Linne, 1758), Ocenebra erinacea (Linne, 1758),
Nassarius reticulatus (Linne, 1758), and Buccinum un-
datiim Linne, 1758 the most frequently cited descrip-
tions of how these neogastropods construct, in the oviduct
and ventral pedal gland, three or four lay ered capsules.
To characterize the laminated egg capsule of Uro-
salpinx cinerea follyensis B. Baker, 1951, Tamarin and
Carriker (1967) published the first comprehensive study
employing light microscopy — including polarized light —
and electron microscopy. They described a capsule with
four laminae composed of asymmetrical protein-like
molecules bound into collagen-like matrices. They also
noted that the great structural complexity of the capsular
wall could only be partialh explained b\ Fretter's (1941)
description of the formative process. Bayne (1968) con-
ducted a histochemical study of several gastropod egg
capsules, including those of Nucella lapillus, and found
mucopolysaccharides in at least three capsular laminae
of that species. He was one of the first to use toluidine
blue for this purpose. Flower et al. (1969) and Flower
(1973) described the origin and the ultrastructure of neo-
gastropod capsular proteins. In another histochemical
study, (Jruber (1982) described six structural layers in
the capsule wall of Eupleura caudata etterae B. B Baker,
1951, and three additional layers associated with tlie
escape aperture and la\ ers of albumen.
C. N. D'Asaro 1988
Page 135
Table 1. Comparative micromorphology of muricacean egg capsules stained in most cases with toluidine blue (meta = meta-
chromatic, mp = mucoid plug, nr = not reported, ortho = orthochromatic, pre = present, tt = two types of albumen cells in the
oviduct, z = zone).
Lasers
Width
closing
Lavers ot
Species
of wall
LI
L2
L3
L4
aperture
albumen
Author
Chicoreus florifer
64 ^m
meta.
meta.
meta.
meta.
LI, mp, L3
two meta
this report
dilectus
4 ^m
54 ixm
2 Mm
4 Mm
(mp), L4,
315 Mm
Phylloiiotus po-
51 nm
meta.
ortho in
meta.
meta.
LI, L3 (mp).
two? meta
this report
imini
1-2 MITl
part.
47 /um
1-2 Mm
2 Mm
mp, L4,
429 Mm
S'uccllu lapillus
nr
meta
pre
meta
nr
Li'' (mp)
meta matrix
Ankel, 1937;
Fretter,
1941;
Bayne, 1968
Urosalpinx cine-
109 Mm
ortho.
ortho.
pre, 2 Mm
ortho.
outer z, dense
pre
Hancock,
rea
30 Mm
75 Mm
2 um
z, L3 or dif-
fuse z, L4,
500 Mm
19.56; Tam-
arin and
Carriker,
1967
Ocenebra erinacea
nr
nr
pre, two fi-
brous
lavers
pre, mu-
coid lay-
er
nr
mp
tt
Fretter, 1941
Eupleura caudata
nr
meta, (1)*
(2-5)
(6?-7)
meta (8)
(1, 6?, 7 =
two meta
Gruber, 1982
ctterac
mp, 8)
* Ten lasers were described; possible relationships for eight are indicated in parentheses.
In the most comprehensive and detailed study to date,
Sullivan and Maugel (1984) used transmission and scan-
ning electron microscopy, histochemistry, and poly-
acrylamide gel electrophoresis to examine the egg cap-
sule of Ihjanassa obsoleta (Say, 1822). They were able
to determine the physical and chemical structure of the
capsule wall and confirm the origin of four structural
laminae in the oviducal capsule gland. A fifth, nonstruc-
tural layer was identified as a product of the ventral
pedal gland.
Since the previous work did not consider egg capsule
micromorphology in a taxonomic perspective, the pur-
pose of this stud) was to describe the complex laminar
structure of egg capsules from two species in each of
four diverse neogastropod taxa: Muricacea, Buccinacea,
Marginellidae, and Conidae. These descriptions, as well
as those in a few papers with similar data, were used to
compare neogastropod egg capsule micromorphology in
order to determine if there is a common wall structure
in the selected taxa, and indirectly, if a common mech-
anism to produce neogastropod egg capsules exists.
MATERIALS AND METHODS
The eight species from St. Joseph Bay in northwest Flor-
ida selected for study are listed in tables 1-3. Egg capsule
macromorphology of each species is known (refer to the
Results for citations and museum voucher numbers).
Specimens for sectioning were preserved in 10% buffered
seawater formalin, a fixative considered acceptable be-
cause egg-capsule proteins are very stable in a range of
acids and bases and are only slightly soluble when au-
toclaved in water (Hunt, 1971). Capsules were taken
from near the center of well-formed egg masses, or if
deposited individually, they were selected to represent
the average size and shape for that species.
To determine the number of egg capsules per species
that must be sectioned to account for possible variations
in staining and micromorphology, preliminary histolog-
ical procedures were performed on two egg capsules
from each of five egg masses of Cantharus multangulus
(Conrad, 1846) deposited by different females. Although
the sectioned capsules were not microstructuralh iden-
tical, a recognizable pattern of structural laminae existed
among all capsules produced by the same and different
females of this species. Because the pattern for C. mul-
tangulus could be recognized in each sectioned capsule
and the literature suggested that it existed for other neo-
gastropod species, sections for the remaining seven species
were prepared from only two or three egg capsules pro-
duced by different females.
.After fixation, the larger egg capsules were opened
along one edge in the longest axis with a scalpel to ac-
celerate diffusion of the embedding fluids. Following
dehydration in ethyl alcohol, capsules were cleared in
toluene and embedded in paraffin (melting point = 56
°C).
Capsules were sectioned at 20-22 um. All sections were
parallel to the longest axis and included some that passed
through the sealed escape aperture and the basal plate.
To separate structural laminae, the capsule wall was frac-
Page 136
THE NAUTILUS, Vol. 102, No. 4
Figure 1. Schematic drawing of a longitudinally sectioned
egg capsule of Cbicurciis florifer dilectiis. Figure 2.
Drawing, like figure 1, of a PhijUonoius pomum egg capsule.
Numbers with arrowheads indicate positions at which photo-
graphs for figures 3-12 were taken. Empty spaces in figures 1
and 2 are artifacts produced during dehydration. A = adhesive,
DA = dense albumen, E = embryo, IMP = inner mucoid plug,
LI = metachromatic first lamina — solid line, L2 = second
lamina — light stipple, L3 = metachromatic third lamina — solid
line, L4 = metachromatic fourth lamina — broken line, OMP
= outer mucoid plug, T.'\ = thin albumen. L2 is metachromatic
in figure 1 and partially orthochromatic in figure 2.
turcd and partially delaminated by varying the speed at
\\ liicli the microtome blade cut a section.
To demonstrate mucopolysaccharides, sections were
stained with toluidine blue for 1.5-2.0 minutes according
to the method of Gurr (1962:440). They were then rap-
id!) delnclrated in ethyl alcohol, cleared in toluene, and
mounted in methyl methacrylate.
Egg capsule microstructure was examined using light
microscopy. Observation of the preliminary sections sug-
gested that toluidine blue metachromasia would produce
a range in color from pur[)le through heliotrope (reddish
purple) but rarely red. Also, some layers would be or-
thochromatic (blue) or colorless. In the intact capsule
wall, structural laminae were difficult to trace by follow-
ing metachromatic la\ers alone. However, when a struc-
tural lamina was first isolated and defined in a fractured
or delaminated section, then the lamina could be rapidly
and comparably traced in whole, stained sections. If a
capsule fractured and delaminated repeatedK' in a con-
sistent pattern, then the parts were considered distinct
structural laminae produced by different types of cells
or produced at different times. Those less than a few
micrometers wide could be recognized when they sep-
arated from the sections and folded to one side. Very
thick sections also allowed microsculpture on the outer
laminar surface to be examined {e.g., figure 3). Following
Tamarin and Carriker's (1967) method, structural lam-
inae were identified from outermost to innermost as Ll-
L4. Diagrams of median sections were prepared with a
drawing tube mounted on a compwund microscope, while
photomicrography was used to record microstructure at
selected positions on a section
Measurements of the capsule wall, structural laminae,
and the sealed escape aperture were taken from the
middle regions of each (tables 1-3). These dimensions
should be considered approximations, as Sullivan and
Mangel (1984) suggested, because of possible shrinkage
introduced by the method and the wide range in width
occurring at approximately the same position on different
capsules of the same species.
RESULTS
Chicoreus florifer dilectus (A. Adams, 1855)
(figures 1, 3-8; table 1; USNM 860426)
Egg capsules of this muricid are distinctly vasiform in
section (figure 1). Macromorpholog\ was described by
D'Asaro (1970:420, fig. 3).
The capsules have a thick, highK fibrous wall in which
four structurally distinct components can be identified:
LI, a thin, fineK fibered, metachromatic outer lamina;
L2, a metachromatic lamina, with layered, coarse fibers
forming the bulk of the wall; L3, the thinnest meta-
chromatic lamina, closely applied to the innermost layer;
and L4, a metachromatic lamina surrounding the al-
bumen (figures 3, 4; table 1). LI is so transparent that
the coarser L2 fibers can be seen through it. Except on
the stalk and basal plate, LI is folded in a corrugated
maruier (figures 3, 4, os). L2 may have fibers directed at
right angles or parallel to the long capsule axis and ar-
ranged with regional differences in two or three fused
layers. When three fibrous components are present, the
middle one has the coarser fibers arranged parallel to the
long axis (figure 3, L2). Where the wall was folded during
formation, L2 fibers separate, forming large \acuoles
(figure 4, v). These vacuoles are not stained and are
probably filled with a nonmucoid liquid. L2 fibers fuse
with a more or less homogeneous inner component in
Figures 3-8. Sections of the Chicoreus florifer dilectus egg capsule. 3. Wall showing laminae 4. Wall just below the apical plate.
5. .Albumen fibers. 6. Mucoid plugs in the escape aperture at the junction with 1.2 7. Wall in the stalk 8. Basal plate and adhesive.
Positions at which photographs were taken are indicated on figure 1. a = adhesive, da = dense albinnen, f = fracture, fl,3 =
C. N. D'Asaro 1988
Page 137
w. .. - V *;^ * A*" I 'fly J -^ V .
•ts
%T:y ta
m
50 u m
• ' — I
fragment of L3, fiL2 licmmm luuus component of L2, imp = inner mucoid plug, isz = intensely stained zone, LI = metachromatic
first lamina, L2 = metachromatic second lamina, L3 = metachromatic third lamina, L4 = metachromatic fourth lamina, os =
outer surface of LI, omp = outer mucoid plug, sL2 = outer portion of L2 in the stalk, ta = thin albumen.
Page 138
THE NAUTILUS, Vol. 102, No. 4
Figures 9-12. Sections of the Phyllonotuti pnmum egg capsule. 9. Wall showing all laminae. 10. Escape aperture toward the
base. 11. Escape aperture toward the ape.x 12. Stalk and basal plate. Positions at which photographs were taken are indicated on
figure 2. a = albumen, cf = circular fibers, e = embryo, hL2 = homogeneous component of L2, imp = inner mucoid plug, Li =
metachromatic first lamina, L2 = partially orthochromatic second lamina, L3 = metachromatic third lamina, L4 = metachromatic
fourth lamina, omp = outer mucoid plug, os = outer surface of LI. sL2 = outer portion of L2 in the stalk, r = ridge.
contact with L3 (figures 3, 4, hL2). L3 stains less inten-
sively than L4. In the apical plate, L3 appears to fuse
with the inner mucoid plug and may have similar com-
position, .^s described by Ankel (1929: fig. 1) for Nucella
lapilhis and Franc (1940: fig. 2) for Ocinelmna aciculata
(Lamarck, 1822), L4 completely surrounds the albumen
and probably contains it during assembly of the capsule
while other components are more fluid. Hereafter, L4
will be called the albinnen retaining layer.
Two structurally distinct types of albumen occur: an
outer layer including metachromatic fibers with random-
ly attached granules in which all embryos arc situated
(figures I, TA; 5, ta) and, an inner core including denser
strongly metachromatic fibers (figures I, DA; 5, da). Fi-
bers of dense albumen ma\ fuse in a continuous layer
where the albuminous components meet; however, the
fused layer did not delaminate during sectioning. The
thin outer albumen does not envelop the denser core
basally (figure 1).
The apical escape aperture is closed by a structuralh
complex barrier composed of four la\ers (figures 1. 6).
LI iscontiguouswith the outer layer. The metachromatic
(purple) second layer or outer mucoid plug is larger than
the aperture and interdigitates with but is clearly sepa-
rated from the homogeneous iimer component of L2
(figure 6, omp, hL2). Fractures across the w idth of the
outer mucoid plug apparently caused during sectioning
occurred (figure 6, f). The third layer is a lenticular,
heliotrope mucoid plug, larger than the aperture, w hich
sometimes fractures during sectioning. L3 is fu.sed to the
lenticular or inner plug in an intensely stained zone (fig-
ure 6, isz). L4 is the innermost layer closing the escape
aperture and is fused to L3 in the same intenseK stained
zone.
The stalk and basal plate are formed from the capsule
wall and an adhesive material (figures 1, A; 7, 8, a). The
outer portion of both includes LI and up to the longi-
tudinal, fibrous component of L2 (figure 7, sL2), while
C. N. D'Asaro 1988
Page 139
the base of the capsular lumen is formed from the re-
maining parts of L2 as well as L3 and L4. Mucoid ad-
hesi\e, which has orthochromatic and metachromatic
zones and occasional L2 fibers, fills the center of the stalk
(figures 7, 8, a).
Phyllonotus pomtim (Gmelin, 1791)
(figures 2, 9-12; table 1; USNM 860425)
The somewhat tongue-shaped capsules and the rather
massive, communal egg masses were described by Perr\
and Schwengel (1955: fig. 338), D'Asaro (1970:422, fig.
3), Radwin and Chamberlin (1973:107, fig. 1), Bandel
(1976a:10, fig. 4), and Moore and Sander (1978:253, fig.
2). Unlike Chicoreus florifer egg capsules, those of Phyl-
lonotus pomum may have two points of attachment w ith
a supporting substratum that nearly always is a conspe-
cific capsule. Sections of such capsules are very variable
in outline.
Phyllonotus pomum capsules have a distinctly fibrous
wall that is only partially metachromatic (table 1). Four
structural parts e.\ist: LI, a thin, metachromatic outer
lamina; L2, a thick, coarsely fibered and partially or-
thochromatic lamina; L3, a strongly metachromatic com-
ponent; and L4, a metachromatic albumen retaining lay-
er (figures, 2, 9). LI is as transparent as the homologous
lamina in Chicoreus florifer dilectus and is corrugated
on or near folds in the wall (figures 10, 11, os). L2 remains
almost unstained. It has a core of longitudinal fibers bor-
dered in most areas by a few circular fibers (figure 9,
L2, cf). Each side of the fibrous zone may be bordered
by a narrow, faintly orthochromatic, homogeneous zone
(figure 9, hL2). L3 has no obvious fibers and appears
fused to the outer mucoid plug of the escape aperture,
as noted later. L4 has distinct circular fibers arranged in
a single layer.
Albumen, including fibrous and amorphous material,
is present in sections only near the retaining layer (figure
10, a). This distribution is an artifact, as noted in Can-
tharus cancellarius, caused by loss of the more fluid
contents during dehydration. Embryos are distributed in
both components of the albumen.
Positioned on one side of the capsule, the sealed escape
aperture includes four layers (figures 10; 11, LI, omp,
imp, L4). The outer component is formed from LI. The
second outermost layer, including intensely stained
metachromatic regions, is continuous with but distinct
from the homogeneous proximal part of L2, and appears
to be fused or continuous with L3. This pattern is dif-
ferent from that of Chicoreus florifer dilectus. A third
layer, which is metachromatic, mucoid with small ves-
icles, and exceedingly fragile, bulges into the capsular
lumen, where it is bordered by L4, the albumen retaining
layer. The more apical and lateral borders of the escape
aperture, formed from L2, have an inward projecting
ridge (figure 11, r).
The basal plate is formed from LI and the outer half
of L2. A separation occurs in a dense reticulum or vac-
uolated zone near the midline of the lamina (figure 12,
Figure 13. Schematic drawing of a longitudinally sectioned
egg capsule of Canlharus multanguliis. Figure 14. Drawing,
like figure 13, of a Cantharus cancellarius egg capsule. Empty
spaces in figures 13 and 14 are artifacts produced during de-
hydration. Numbers with arrowheads indicate positions at w hich
photographs for figures 15-23 were taken. A = adhesive, .\D
= apical depression, BM = basal mucoid material, D.A = dense
albumen, E = embryo, LI = orthochromatic first lamina —
solid line, L2 = second lamina — light stipple, L4 = metachro-
matic fourth lamina — broken Une, MP = mucoid plug, TA =
thin albumen
sL2). Where the outer half of L2 was in contact with the
substratum, a homogeneous zone having some fibers ex-
tending through it toward the substratum is present. The
inner half of L2, L3, and L4 form the floor of the capsular
lumen. No adhesive layer was observed on capsules that
had been attached to those of conspecifics.
Cantharus multangulus (Philippi, 1848)
(figures 13, 15-19; table 2; USNM 947143)
Macromorphology of the egg capsule was described by
Perrv and Schwengel (1955: fig. 340), Radwin and Cham-
berlin (1973:110, fig. 6), and D'Asaro (1986a:85, fig. 3).
Median sections through the ribbed and spined capsule
are roughK vasiform in outline and have a rounded,
apical depression partly covered by a transverse ridge
Page 140
THE NAUTILUS, Vol. 102, No. 4
Figures 15-19. Sections of the Canlhartis mullai^gulus egg capsule 15. Wall showing laminae 16. Escape aperture adjacent
to apical depression 17. Escape aperture on side opposite apical depression. 18. .Alliunien layers 19. Stalk and adhesive. Positions
at which photographs were taken are indicated on figure 13. a = adhesive, ad = apical depression, bm = basal mucoid material,
da = dense albumen, LI = orthochromatic first lamina, L2 = second lamina, L2m = metachromatic component, L2o = ortho-
chromatic component, mp = mucoid plug, os = outer surface of LL ta = thin albumen.
C. N. D'Asaro 1988
Page 141
Table 2. Comparative niicromorplinlogy of buccinacean egg capsules stained in
c'hronialic, nip = mucoid plug, nr = not reported, ortho = orthoclironiatic, pre =
oviduct)
most cases with toluidine blue (mcta = meta-
present, tt = two types of albumen cells in the
Species
Width
of wall LI
L2
L3
L4
Layers
closing Layers of
aperture albumen
Author
Cantharus ntul-
langulus
23 iim ortho, ortho and vestige meta,
1-2 /im meta, 1-2 /tm
21 ^m
Cantharus cancel- 22 fim ortho, ortho and vestige meta,
larius 2 ^m meta, 3— i ^m
17 urn
Btisycon carica nr pre pre pre pre
and B. canalicu-
latum
Nassarius reticiila- nr three layers of uncertain relationship mentioned
tus
Buccinium undo- nr pre two fibrous mucoid nr
turn layers layer
Ilyanassa obsoleta 11-22 meta pre, meta, 1 ^m pre, 60 nm
(LO pre) Mm 2 nm 9-10 ^m
LI, L2, mp.
two
meta
this report
L4,
210 /im
LI, L2, mp,
two
meta
this report
L4,
117 Mm
nr
two
Harasewych,
1978
nr
tt
Fretter, 1941
nr
tt
Fretter, 1941
LI, L2, L3
nr
Sullivan and
(mp), L4,
Maugle,
110-150
1984
Min
(figure 13, AD). In the intact capsule, the depression is
not obvious when viewed apically. Cantharus cancel-
larius (Conrad, 1846) has similar sculpture on its apical
plate (figure 14, AD).
Capsules of C. miiltangulus have a wall in which the
fibers appear fused in most regions when viewed with
light microscopy. There are three distinct components
that appear homologous to three of the four muriccacean
laminae: LI, a dense, orthochromatic outer lamina; L2,
a mostly homogeneous central lamina, including vacuoles
in some regions and fibers in others; and L4, a retaining
layer surrounding the albumen (figure 15; table 2). LI
stains intensely and may appear black. L2 may have up
to four layered but fused components with the outermost
being metachromatic and the innermost orthochromatic.
The innermost component, like LI, is intensely stained
and may appear black (figure 15, L2o). L2 has somewhat
tubular vacuoles in the metachromatic area, especially
near the apical depression and where the wall is folded
into ridges or spines (figures 16, ad, L2m; 17, L2m). L3,
which in many muricaceans is continuous with a mucoid
plug in the escape aperture, is not a distinctive feature
in the capsule wall (see the following comments con-
cerning the escape aperture). The albumen retaining
layer is characteristically wrinkled when it separates from
the wall (figure 15, L4).
Albumen occurs in two metachromatic components:
a dense portion, broadest near the apical plate, containing
short, closely packed fibers with attached, irregular gran-
ules, and a thinner, more diffuse portion with coarser
fibers and granules, filling most of the lumen (figures 15,
da; 18, da, ta). Embryos are positioned in the latter.
The escape aperture is closed by four layers. LI, the
outer component, is in continuous contact with the or-
thochromatic part of L2 (figures 16; 17, L2o). The or-
thochromatic part of L2 is somewhat folded, intensely
stained, and may fracture vertically during sectioning.
The broadest layer closing the aperture is a metachro-
matic, mucoid plug lying between the orthochromatic
part of L2 and L4 (figures 13, MP; 16, 17, mp). Apically,
the mucoid plug lies between L2 and L4 in the homol-
ogous position for L3 in muricaceans, but it does not
extend to the base of the lumen as a continuous lamina
(figure 13). However, above the stalk between L2 and
L4, there is some metachromatic mucoid material with
chromotropic characteristics similar to those of the apical
plug (figures 13, BM; 19, bm). This mucoid material
could have been formed at the same time as the plug
and would be homologous to L3. L4 is the innermost
layer closing the escape aperture.
The stalk and basal plate are formed from LI and
most of L2, while the base of the capsular lumen is
formed from the orthochromatic part of L2, some mu-
coid material, and L4 (figure 19, L2o, bm, L4). The stalk
has a core of metachromatic mucoid adhesive (figure 19,
a).
Cantharus cancellarius (Conrad, 1846)
(figures 14, 20-23; table 2; USNM 847140)
The macromorphology of the egg capsules of this species
was described by Radwin and Chamberlin (1973:110,
fig. 8) and D'Asaro (1986a:84, fig. 2). Most median sec-
tions of the capsules are cylindrical to vasiform in outline
(figure 14). A rounded depression in the apical plate,
similar to one observed in C. multangulus, is prominent
and partially covered by a raised transverse ridge folded
toward the escape aperture.
As in C. multangulus, much of the capsule wall ap-
pears to be composed of fused fibers. Three structural
Page 142
THE NAUTILUS, Vol. 102, No. 4
Figures 20-23. Sections of the Cantharus cancellarius egg capsule. 20. Wall showing laminae 21. Escape aperture opposite
apical depression. 22. Escape aperture adjacent to apical depression. 23. Basal plate and adhesive. Positions at which photographs
were taken are indicated on figure 14. a = adhesive, ad = apical depression, bm = basal mucoid material, da = dense albumen,
LI = orthochromatic first lamina, L2 = second lamina, L2m = metachromatic component, L2o = orthochromatic component, mp
= mucoid plug, OS = outer surface of LI, sL2 = outer portion of L2 in the stalk, ta = thin albumen.
laminae, homologous to three of the four muricacean
laminae, are present. These are: LI, a very thin, ortho-
chromatic outer lamina; L2, an orthochromatic and
metachromatic central lamina with tubular vacuoles; and
L4, a metachromatic albumen retaining layer (figure 20;
table 1). LI is often denser on the apical plate. Distinct
tubular vacuoles are present in the central portion of L2,
especially where ridges e.xist in the wall and apical plate.
Longitudinally directed vacuoles are positioned nearer
to the outer surface, while more circularly directed vac-
uoles are nearer to the ituier surface. Apically, where the
wall is folded, fibers in the lamina are visible (figure 21).
As shown for C multangulus, L3 is not present as a
distinct and continuous lamina throughout the whole
capsule. The metachromatic albumen retaining layer is
extremely thin and can be identified only where it is
pulled away from the albimien or the capsule wall and
folded to one side (figure 20, L4). Dense metachromatic
albumen with fine fibers surrounds more coarsely fibered,
less dense albumen in which embryos are positioned (fig-
ure 20, da, e, ta).
The escape aperture is closed by four layers arranged
in a pattern essentially identical to that of C. multan-
gulus. Apically, LI forms the outermost layer (figures
21, 22). The expanded, orthochromatic part of 1,2 is the
second component. It has \ertical folds or fractures (fig-
ures 21; 22, L2o). The broadest component is a meta-
chromatic mucoid plug lying between the orthochro-
matic part of L2 and L4 (figures 21; 22, mp). This plug
tapers basally between L2 and L4, but does not extend
to the base as a continuous lamina equivalent to L3.
There is a layer of similar mucoid material positioned
between L2 and L4 at the base of the lumen (figures 14,
BM; 23, bm).
Basally, the plate is formed from LI and most of L2
(figures 14; 23, sL2). Only the orthochromatic, proximal
C. N. D'Asaro 1988
Page 143
part of L2 extends across the base, where it, some mucoid
material, and L4 form the floor of the lumen. Unlike
most specimens of C. multangulus. the lumen is sepa-
rated from the substratum b\ only a narrow layer of
metachromatic adhesive (figure 23, a).
Contis jioridanus flohdensis Sowerby, 1870
(figures 24, 26-29; table 3; USNM 847141)
Median capsular sections typically have the longitudinal
axis angling away from the axis of the stalk and show a
slightly irregular outer surface (figure 24). Macromor-
phology was described by Perry and Schwengel (1955:
fig. 360, as C. spurius atlanticits Clench, 1942) and
D'Asaro (1986a:88, fig. 4).
The capsule wall has three distinct components: LI, a
fibrous metachromatic outer lamina; L2, a finely fibered,
orthochromatic central lamina; and L4, a thin, meta-
chromatic albumen retaining layer (figure 26; table 3).
LI is very transparent and has fibers circular in cross
section. Occasional corrugations mark this lamina, es-
pecially near the escape aperture and external ridges
(figure 27, LI). Fibers in L2 tend to form a cross-hatched
pattern in the upper three quarters of the capsule, but
some align with the long axis in the stalk and the base
(figures 26-29). Globular structures buried in the fibrous
layer occur frequently (figure 27). No homologue of L3
is present as a distinct layer, nor is there evidence that
the mucoid plug in the escape aperture is homologous
to L3. The albumen retaining layer (L4) is not obviously
fibrous. Although this layer is the third in sequence, it is
designated as L4, the albumen retaining layer, because
its position and probable function appear the same as L4
in Vluricacea and Buccinacea.
in this species, only one layer of fibrous, metachro-
matic albumen is visible (figure 26, al). Granules are
attached to the widely separated fibers. Embryos are
randomly positioned in the albumen.
The sealed escape aperture is less complex than similar
structures formed by muricaceans or buccinaceans. It is
lined on its outer surface by LI and on its inner surface
by L4. L2 is replaced by metachromatic, layered mucoid
plug with which it interdigitates extensively (figure 27,
mp). The degree of interdigitation suggests that L2 and
the mucoid plug were formed at the same time.
The stalk is composed of LI and almost all of L2 from
both sides, while the basal plate on each side is composed
of LI and L2 from the same side (figures 28, 29, sL2).
A fibro-mucoid, metachromatic adhesive, which may ex-
tend into the stalk, attaches the basal plate to the sub-
stratum (figure 29, a). Fibers from L2 radiate widely into
the adhesive, especially where it extends into the stalk.
Conus jaspideus stearnsi Conrad, 1869
(figures 25, 30-32; table 3; USNM 847148)
When sectioned, the lamellate capsules, described by
D'Asaro (1986a:88, fig. 4), appear pointed at the apex
with a broad stalk and an escape aperture on one side.
Figure 24-. Schematic drawing of a longitudinally sectioned
egg capsule of Conus floridanus floridensis. Figure
25. Drawing, like figure 24, of a Conus jaspideus stearnsi
egg capsule. Numbers with arrowheads indicate positions at
which photographs for figures 26-32 were taken. \ = adhesive,
.\L = albumen, E = embryo, LI = metachromatic first lamina
— solid line, L2 = second lamina — light stipple, L4 = meta-
chromatic albumen retaining layer — broken line, MP = mucoid
plug.
In this plane, the basal plate is the widest part (figure
25).
Three laminae were identified: LI, a finely fibered,
metachromatic outer lamina; L2, a complex orthochro-
matic and metachromatic lamina with fibrous and mu-
coid components; and L4, a metachromatic albumen re-
taining layer (figure 30; table 3). LI, with circular fibers,
is tightly fused to the fibrous portion of L2. Except near
the escape aperture and on the stalk, this layer is uni-
formly corrugated (figures 30; 31, os). L2 has an ortho-
chromatic outer component with coarse, circular fibers
(figure 30, L2). The inner component of L2 consists of
at least three metachromatic layers of mucoid material
containing scattered fibers arranged parallel with the
long axis (figures 31, 32, mL2). Of the species studied,
only C. /. floridanus has a similar arrangement of fibers
in mucoid material, and that occurs only in the stalk. No
distinct lamina equivalent to L3 is present. L4 may have
fragments of albumen fused to its inner surface (figure
32, is).
Albumen in this species is metachromatic and includes
scattered transparent spherules and irregular granular
material that may form layers (figures 30-32). Embryos
are distributed throughout it.
Page 144
THE NAUTILUS, Vol. 102, No. 4
LI
n.
Figures 26-29. Sections ot the Conu:> jioridanm floridensis egg capsule. 26. Wall showing laminae. 27. Escape aperture. 28.
Stalk 29. Basal plate and adhesive. Positions at which photographs were taken are indicated on figure 24 \ = adhesive, al =
albumen, is = inner surface of L4, LI = metachromatic first lamina, L2 = orthochromatic second lamina, L4 = metachromatic
albumen retaining layer, mp = mucoid plug, as = outer surface of LI, sL2 = outer portion of L2 in the stalk.
Table .'{. Comparative micromorphology of conid and marginellid egg capsules stained in most cases with toluidine blue (meta
irutai hromatic. mp = mucoid plug, ortho = orlhochromatic. pre = present).
Species
Width
of wall
LI
L2
L.3
L4
Lasers
closing
aperture
Layers of
albumen
Author
Conus floridanus 49 ^m meta,
floridensis .5 nm
Conus jaspidius 25 niu meta,
slearnsi 2-3 Mm
Granulina ovuli-
formia
\r> fim
Marginella aureo- 12 ^m
cincta
ortho, not pre
40 M'li
ortho and not pre
meta,
22 nm
one lamina present
one lamina present
meta, LI, mp, L4, one meta
3-4 nn\ 330 ^m
meta, I.I, mp. L4. one meta
1-2 ^m I N ^m
suture in wall two meta
suture? two meta
this report
this report
this report;
D Asaro,
1986
this report;
D'.Asaro,
1 9.Sfi
C. N. D'Asaro 1988
Page 145
LI forms the outer covering of the escape aperture,
w hile the inner lining is L4. A metachromatic, mucoid
plug replaces L2 (figure 31, mp). The edges of the mucoid
plug are separated from L2 b> an interdigitating meta-
chromatic boundary zone, similar but less complex than
that of C. /. floridensis. None of the scattered fibers in
the metachromatic mucoid layers of L2 enter the mucoid
plug.
The two outer laminae form the stalk and basal plate,
w hile the base of the capsule is composed of the albumen
retaining layer and the inner, mucoid part of L2 (figure
32, mL2). The stalk varies in length; therefore, in some
specimens the lumen of the capsule may be nearly level
with the substratum. A dense mucoid adhesive is present
basalK (figure 32, a).
Granulina ovuUjormis (Orbigny, 1841)
(figure 33; table 3; USNM 836973)
Macro- and micromorphology of the egg capsules, based
in part on eosinophilic features, were described by D'Asaro
(1986b: 196, figs. 3-5). Because of structural differences,
marginellid capsule micromorphology is not describable
with reference to Tamarin and Carriker's (1967) system
of enumerating capsule laminae.
The simple, pustulate capsules are constructed from
an inner component, enclosing two layers of albumen
and an embryo, and basement and outer components
that sandwich and fuse the previously mentioned struc-
ture between them (D'Asaro, 1986b). Because these com-
ponents were uniformly unstained and did not delami-
nate during sectioning, the capsule wall is defined as
having a single structural lamina of uniform composition,
possibK formed by the same portion of the oviduct that
produced the layers of L2 in muricaceans and bucci-
naceans (figure 33, cw). The outer surface of the capsule
is coated with metachromatic mucus. No escape aperture
with a mucoid plug exists. The point at which the capsule
wall fractures at hatching is marked by a distinct meta-
chromatic suture (figure 33, s). Dense granular albumen
lies just below the capsule wall, while a less dense com-
ponent immediately surrounds the single embryo. Both
albuminous layers stain metachromatically. The adhesive
on the basal layer is not stained (figure 33, a, bl).
Marginella aureocincta Stearns, 1872
(figure 34; table 3; USNM 836974)
Macro- and micromorphology of the egg capsule were
described by D'Asaro (1986b: 195, figs. 3-5). 'With ap-
plication of toluidine blue, all parts of the capsule are
metachromatic, except the unstained basement adhesive.
The capsule wall does not delaminate and stains intensely
to the point of obscuring its layered structure suggesting
that, as in Granulina ovuliformis, it should be defined
as a single structural lamina (figure 34, cw, osw). Dis-
tinctive granulations, described by D'Asaro (1986b: 195,
fig. 3), are obvious on the outer surface. No specific
fibrous layer surrounds the albumen. The metachro-
matic, finely granular albumen occurs in two compo-
Figures 30-32. Sections of the Conus jaspideus stearnsi egg
capsule. 30. Wall showing laminae 3 1 . Escape aperture toward
apex. 32. Stalk, basal plate, and adhesive. Positions at which
photographs were taken are indicated on figure 25. a = ad-
hesive, al = albumen, is = inner surface of L4, LI = meta-
chromatic first lamina, L2 = orthochromatic and metachro-
matic second lamina, L4 = metachromatic albumen retaining
ia\er, mL2 = mucoid component of L2, mp = mucoid plug,
OS = outer surface of LL
Page 146
THE NAUTILUS, Vol. 102, No. 4
Figure 33. Section of the GranuHna ovitlifurmis egg capsule attached to a Thalassia leaf. Figure 34. Section of the Marginella
aureocincta egg capsule, a = adhesive, bl = basement layer, cw = capsule wall, da = dense albumen, osw = outer surface of
capsule wall, s = suture, ta = thin albumen.
nents, a narrow band of dense material and a more fluid,
central zone in which the embryo lies (figure 34, da, ta).
DISCUSSION
Separation of structural laminae by fracturing and to-
luidine blue staining allows differentiation and identifi-
cation of homologous components of neogastropod egg
capsules. Laminae can be identified in this manner be-
cause the fibers of which they are composed appear to
have been secreted and chemically bound together dur-
ing separate phases of assembly. Separate or combined
functions related to protection, structural support, release
of larvae or juveniles, and albumen retention are inferred
for the structural laminae.
,\mong the iieogastropods studied, three microstruc-
tural patterns of laminae were identified, each charac-
terizing specific higher taxa. Complex, four-layered cap-
sules with an escape aperture sealed by four layers,
including two that are mucoid, were found in nearly all
muricaceans and buccinaceans examined. Three-layered
capsules having an escape aperture sealed by three layers,
including one that is mucoid, were found in the Conidae.
Uncomplicated, single-layered capsules that may only
have a suture in the wall to facilitate hatching appear
to be characteristic of the Marginellidae.
Muricacea and Buccinacea are species-rich taxa hav-
ing family or species-specific egg capsule morphologies
with a final shape that results from a molding process in
the ventral pedal gland (.Ankel, 1929; Gruber, 1982; Sul-
livan and Maugcl, 1984). Micromorphological similari-
ties suggest that oviducal mechanisms common to both
superfamilies are u.sed to construct structural laminae
that serve the same respective functions in these taxa
(tables 1, 2). Some species may add additional nonstruc-
tural layers in the ventral pedal gland (table 2, LO; Sul-
livan and Maugel, 1984).
Protection (sealing the fibrous wall) appears to be a
function of the outer structural lamina, LI. In most
species, LI is thin (5 ^m or less), usually dense and finely
fibered, and reflects the final shape (corrugations and
ribbing) imparted by the ventral pedal gland (Gruber,
1982; Sullivan and Maugel, 1984). LI seals the surface
of the w hole capsule including the escape aperture, but
not the basal plate in contact with the adhesive. Since
the contents of some prosobranch egg capsules are axenic
but not bacteriostatic (Lord, 1986), the fine structure of
this lamina could serve as a ph\sical barrier to invasion
by microorganisms, especially in species with a vacuo-
lated and fluid-filled L2. (The basal adhesive may also
serve as a barrier.) In Urosalpinx cinerea, LI is thick
with separated fibers, but the interstices are filled with
dense mucus (Tamarin and Carriker, 1967). LI could
also serve as a reactive substrate during the molding and
hardening process in the ventral pedal gland which fixes
the capsule in its final shape (see Gruber, 1982 and Sul-
livan and Maugel, 1984).
L2 comprises the internal skeleton of the capsule wall
in all neogastropod taxa studied except Marginellidae,
which has a single structural lamina (tables 1-.3). In some
taxa, extensive cross-linkages between the protein fibers
form a homogeneous L2. In others, there are dense ho-
mogeneous zones on inner and outer surfaces gradually
.separating into directionally oriented fibers forming most
of the lamina. In ribs or other sculpture, these fibers are
loo.sely packed. The homogeneous zones, in addition to
skeletal support, could pro\ide another ph\sical barrier
to invasion by microorganisms.
In Muricacea and Buccinacea, L2 may have fibrous
components with different axial orientation often de-
scribed as separate layers (Fretter, 1941; Gruber, 1982).
Resistance to delamination and cohesiveness suggest that
these layers were constructed from the same capsular
protein during a continuous process; thus, the\ should
be described as parts of a single capsular lamina. If all
similar fibrovis components anil contiguous homogeneous
C. N. D'Asaro 1988
Page 147
zones from previously described egg capsules of these
superfamilies are viewed in this manner (as organized
in tables 1, 2), then more meaningful comparisons be-
tween taxa can be made.
L3 is distinct in most muricaceans and buccinaceans
and appears to be a mucoid extension of the apical plug,
a part of the hatching mechanism (tables 1, 2). In both
species of Cantharus, L3 is not present in most of the
capsule wall; however, the mucoid plug in the escape
aperture tapers gradually into the wall in a homologous
position. There is also mucoid material with chromo-
tropic characteristics similar to the apical plug in a ho-
mologous basal position suggesting that a thin layer of
L3 mucopolysaccharide may be present throughout the
wall but could not be identified with the histological
technique applied. In the egg capsule of Eupleura cau-
data etterae, which was described by Gruber (1982: fig.
14), the singular mucoid plug extending as a layer into
the capsule wall and surrounding the lumen can be de-
scribed as L3. Lying between the outer structural layers
and L3 in E. c. etterae, is another layer (Gruber s sixth
layer). The position of this layer, in contact with the
equivalent of L2 and surrounding the outer edge of the
mucoid plug formed by L3, and its composition suggest
that it is homologous with the first mucoid plug in Chi-
coreus florifer dilectus (table 1).
L4. the albumen retaining layer, is the primary lamina
in muricaceans and buccinaceans completely surround-
ing albuminous fluids and embryos (tables 1, 2). Often
it is not reported in the literature, perhaps because it is
4 nm or less in thickness and usually bound tightly to
other components. A function of the layer could be to
prevent nonrefractory albumen from mixing with re-
fractory capsular proteins when the outer structural lam-
inae are assembled by the ciliary mechanism Fretter
(1941) described.
Albumen in egg capsules of muricaceans and bucci-
naceans is stratified to some degree at oviposition (tables
1, 2). This was rather obvious in capsules of Chicoreus
florifer dilectus, which had a core of dense albumen,
free of embryos, that was structurally different from the
surrounding, less dense material. More typically in other
species, the core albumen contained the embryos. Dif-
ferent layers of albumen in newly deposited capsules
suggest that two or more kinds of albumen producing
cells exist in the oviducts of the species studied, as Fretter
(1941) has demonstrated for several neogastropods.
Muricaceans and buccinaceans have structurally sim-
ilar barriers closing the escape apertures, which are five
to nine times as thick as the capsule wall (tables 1, 2).
The increased thickness of the less refractory mucoid
plugs possibly serves to prevent premature hatching of
a capsule. In muricaceans, no fibers from the more re-
fractory parts of L2 extend into the mucoid plugs. During
sectioning, mucoid plugs in the escape aperture fractured
across the width of the capsule wall, while the more
refractors parts of the capsule delaminated lengthwise.
Boundary laminae in the wall (LI and L4) appear to
hold the mucoid plugs in position.
Conid egg capsules are less complex than those of the
Muricacea and Buccinacea and contain smaller structural
fibers, usually embedded in a mucoid component. The
capsule wall has two laminae (LI and L2) and an al-
bumen retaining layer (L4). Rather than having ribbon-
like fibers in the laminae (see Flower et ai, 1969, for a
discussion of fiber ultrastructure), there are very fine,
short, and folded or twisted fibers that overlap into a
cross-hatched pattern. A singular mucoid plug lying be-
tween the outer laminae, LI, and the albumen retaining
layer, L4, and interdigitating extensively with the fibrous
middle lamina seals the escape aperture. Extensive in-
terdigitation would require simultaneous formation of
L2 and the mucoid plug. There is no evidence in the
conids studied of a lamina equivalent to L3 associated
with the mucoid plug. These differences in micromor-
phology suggest that important differences in oviducal
structure and function exist between conids and muri-
caceans and buccinaceans.
Pustulate marginellid egg capsules are microstruc-
turally the least complex of the taxa studied, and differ
markedly from capsules of other neogastropods. Each is
constructed of three homogeneous components fused al-
most indistinguishably into the others to form a capsule
in which the parts do not delaminate during sectioning.
Resistance to delamination and a uniform response to
toluidine blue suggest that only one protein was used to
form the capsule wall. There is no complex escape ap-
erture, but in many marginellids it is possible to identify
a preformed suture at which the capsule will break dur-
ing hatching. The suture is visible at one end and on the
sides, which corresponds exactly to the position where
the capsule wall breaks during hatching. These features
also suggest that the structure and function of the mar-
ginellid oviduct is different from that of the major su-
perfamilies studied.
Although this study does show that there are micro-
morphological features common to the egg capsules of
some prosobranch taxa, it does not provide clear evidence
that can be used to explain exactly how capsules are
formed in the oviducal glands. It can be inferred that
L4, the albumen retaining layer, is deposited around the
albumen and embryos to hold them in a central position
\\ hile the more refractory parts of the capsule are formed.
It can also be inferred that for most species the remaining
parts of the process involve sequential deposition of struc-
tural laminae, as Fretter (1941) described for Nassarius
reticulatits, with L3 and formation of the innermost mu-
coid plug being the second part of the process. The third
part of assembly would involve formation of the main
structural lamina, L2, and the outer mucoid plug. LI
would be the last structural lamina added. Sculpturing,
hardening, addition of nonstructural la\ ers, and attach-
ment with an adhesi\e are functions of the ventral pedal
gland (Sullivan and Maugel, 1984).
ACKNOWLEDGEMENTS
I wish to acknowledge the revisions to the manuscript
suggested by a colleague. Dr. Paul V. Hamilton. Some
of the specimens sectioned were provided by Mr. Larry
Page 148
THE NAUTILUS, \'ol. 102, No. 4
Dilmore. This project was financed in part with funds
from the U.S. Environmental Protection Agency (CR-
811649) and the University of West Florida.
LITERATURE CITED
Abe, .Naoya. 1983. Breeding of Thais clavigera (Kiister) and
predation of its eggs by Cronia margariticola (Broderip).
In: Morton, B. and D. Dudgeon (eds.). Proceedings of the
Second International Workshop on the Malacofauna of
Hong Kong and Southern China Hong Kong University
Press, Hong Kong, p. 381-392.
Ankel, W. E. 1929. Lber die Bildung der Eikapsel bei
Nassa-Arten. V'erhandlungen der Deutschen Zoologischen
Gesellschaft, Zoologischer Anzeiger, supplement 4:219-
230.
Ankel, W. E. 1937. Der feinere Bau des Kokons der Purpur-
schnecke Nucella lapiUus (L.) und seine Bedeutung fiir
das Laichleban. Verliaiidlungen der Deutschen Zoolo-
gischen Gesellschaft, Zoologischer Anzeiger, supplement
10:77-86.
Bandel, K. 1975. Embryonalgehause Karibischen Meso- und
Neogastropoden (Mollusca). .\kademie der Wissenschaf-
ten und der Literalur. Mainz. Abhandiuiigen der Math-
ematisch-Xaturwissenschaftlichen Klasse. 1975(1): 1-133.
Bandel, K. 1976a. Morphologic der Gelege und okologische
Beobachtungen an Muriciden (Gastropoda) aus der siid-
lichen Karibischen See. Verhandlungen der Naturfor-
schenden Gesellschaft in Basel 85(1/2): 1-32.
Bandel, K 1976b Spawning, development and ecology of
some higher Neogastropoda from the Caribbean Sea of
C;olombia (South America), X'eliger 19:176-193.
Bandel, K. 1976c Morphologic der Gelege und okologie Beo-
bachtungen an Buccinacean (Gastropoda) aus der siidlich-
en Karibischen See. Bonner Zoologische Beitraege 27:98-
133,
Bandel, K. 1982. Morphologic und Bildung der friihontoge-
netischen Gehause bei conchiferen Mollusken Facies (Er-
langen) 7:1-153.
Bayne, C. J. 1968 Histochemical studies on the egg capsules
of eight gastropod molluscs. Proceedings of the Malacol-
ogical Society of London 38:199-212.
D'Asaro, C.N. 1970. 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.
D'Asaro, C. N. 1986a. Egg capsules of eleven marine proso-
branchs from northwest Florida Bulletin of Marine Sci-
ence 39(1):76-91.
D'Asaro, C.N. 1986b. Laboratory spawning, egg membranes,
and egg capsules of 14 small marine prosobranchs from
Florida and Bimini, Bahamas. American Malacological
Bulletin 4(2): 18.5- 199.
Flower, N. E. 1973. The storage and structure of proteins
used in the production of egg capsules by the mollusc
CominvUa maculosa. Journal of l'ltra.*tructure Research
44:1.34-145.
Flower, N. E., A. J. (;eddes, and K M Kudall 1969 I'ltra-
structurc of the fibrous prolcin from the egg capsules of
the whelk Buccinium undatum Journal of Ultrastructure
Research 26:262-273.
Franc, A. 1940, Recherchessur ledeveloppement d'Ocinefcra
ackulala Lamarck (Mollus()uc gasteropode). Bulletin Biol-
ogicjue de la France ct dc la Belgique 74327-345.
Fretter, V. 1941. The genital ducts of some British stenog-
lossan prosobranchs. Journal of the Marine Biological As-
sociation of the I iiited Kingdom 25173-211
Goldsmith, L. A., H, Hanigan, J M, Thorpe, and K. A. Lind-
berg. 1978. Nidamental gland precursor of the egg cap-
sule protein of the gastropod mollusc Busycon carica.
Comparati\e Biochemistr\ and Physiology 59B:I33-138.
Gruber, G. L. 1982. The role of the ventral pedal gland in
formation of an egg capsule by the muricid gastropod
Euplcura caudata etterae B. B Baker. 1951: an integrated
belia\ ioral. morphological, and histochemical study Mas-
ter s thesis. University of Delaware, 142 p.
Gurr, E. 1962. Staining animal tissue, practical and theoret-
ical. Leonard Hill, London. 631 p.
Hancock, D. A. 1956. The structure of the capsule and the
hatching process in Urosalpinx cinerea (Say). Proceedings
of the Zoological Society of London 127565-571.
HarasewNch, M. G. 1978. Biochemical studies of the hatching
process in Busycon. Master's thesis. University of Dela-
ware, Newark, 52 p
Hunt, S. 1971. Compari.son of three extracellular structural
proteins in the gastropod mollusc Buccinum undatum L.,
the periostracum, egg capsule, and operculum. Compar-
ative Biochemistry and Physiology 40B:37-46.
Lord, .\ 1986. .\Te the contents of egg capsules of the marine
gastropod Nucella lapillus (L.) axenic':* American Mala-
cological Bulletin 4(2):201-204.
Moore, E. A. and F Sander 1978 Spawning and early life
historv of Murex pomum Gmelin, 1791. N'eliger 20:251-
259.
Pechenik, J. .\. 1979. Role of encapsulation in in\ertebrate
life histories. American Naturalist 104(6):859-870.
Pechenik, J. A. 1983. Egg capsules of Sucella lapillus (L.)
protect against low-salinit\ stress. Journal of Experimental
Marine Biology and Ecologv 71:165-179.
Pechenik, J .A,. 1986. The encapsulation of eggs and embryos
by molluscs: an overview .American Malacological Bul-
letin 4(2): 165-172.
Perry, L. M. and J. S. Schwengel. 1955. Marine shells of the
western coast of Florida. Paleontological Research Insti-
tution, Ithaca, New York, 318 p.
Radwin, G. E. and J. L. Chamberlin. 1973. Patterns of larval
development in stenoglossan gastropods. Transactions of
the San Diego Society of Natural History 17(9): 107-1 18.
Sullivan, C. H. and D. B. Bonar 1984. Biochemical charac-
terization of the hatching process of llyanassa ohsoleta.
Journal of Experimental Zoology 229(2 ):223-234.
Sullivan, C. H. and T. K Maugel 1984. Formation, organi-
zation, and composition of the egg capsule of the marine
gastropod llyanassa obsoleta. Biological Bulletin 167:378-
389.
Tamarin, A. and M H C^arriker. 1967. The egg capsule of
the muricid gastropod I'rnsalpinx cinerea an integrated
stud) of the wall b\ ordinarv light, polarized light, and
electron microscop\ Journal of Ultrastructure Research
21:26-40.
Webber, H. H. 1977. Gastropoda, Prosobranchia. /n- Giese.
J. S. and J. S. Pearse (eds). Reproduction of marine in-
vertebrates. Vol. IV, molluscs: gastropods and cephalopods.
Academic Press. New York, 114 p.
THE NAUTILUS 102(4):149-153, 1988
Page 149
Two New Species of Metula (Gastropoda: Buccinidae) with a
Description of the Radula of the Genus
Philippe Bouchet
Museum National d'Histoire Naturelle
55, rue Buffoii
75005 Paris, France
ABSTRACT
Two species of Metula H. and A. Adams, 1853, lack a radula,
and it is very small in a third species. The morphology of the
teeth is significantly different from that in Pisaiiia, and the two
genera are probably not closeK related Metula crosnicri new
species, from 400-450 m off SW Madagascar, is a large, broad
species, with \ery convex whorls and a deep suture. Metula
africana new species, from the deep continental shelf off West
Africa, is considered the descendant of the Mediterranean Plio-
cene M. milraeformis (Brocchi, 1814). This lineage cannot be
taken as evidence for Mediterranean-Indo-Pacific connections
in the lower Pliocene as claimed by Grecchi (1978).
INTRODUCTION
The chequered taxonomical history of the generic name
Metula H. and A. Adams, 1853, has recently been sta-
bilized by Emerson (1986), who clarified the identity of
its type species, Buccinum clathratitm Adams and Reeve,
1850. Additional information was provided by Beu and
Maxwell (1987).
The familial position of the genus has been the subject
of a controversy between Ponder (1968, 1973) and Cer-
nohorsky (1971). Ponder (1968) described the anatomy
of Ratifitsus Iredaie, 1919, and Iredalula Finlay, 1927;
he concluded that their peculiar glandular mid-esoph-
agus as well as other features of the anterior alimentary
canal justified their inclusion in the family Colubrariidae,
which he considered to be anatomically distinct from
the Buccinidae. Cernohorsky (1971) regarded the pres-
ence of a vestigial radula in Ratifusus and Iredalula to
indicate placement in the Buccinidae, since the species
of Colubrariidae have no radula at all, and he suggested
a placement in the buccinid subfamily Pisaniinae. This
view has been accepted by most subsequent authors deal-
ing with Metula (Olsson and Bayer, 1972; Kilburn, 1975;
Houbrick, 1984; Emerson, 1986), who apparently over-
looked Ponder s (1973) refutation of Cernohorsky s point
of view. Ponder (1973) confirmed that Ratifusus, Ire-
dalula. and Colubraria shared the same anatomical char-
acters that separate them from the Buccinidae, and com-
mented on the radular differences: "It thus appears that
some Colubrariidae have lost the radula and that it is
relatively small or vestigial in the remainder. It is possible
that the whole Metula-Ratifusus series discussed by Cer-
nohorsky (1971) belongs in the Colubrariidae as they all
have similar shell features" (Ponder, 1973:328).
The family Colubrariidae is treated as a synonym of
Buccininae by Ponder and Waren (1988), while Beu and
Maxwell (1987) recognize a subfamily Pisaniinae, where
they include Metula, Colubraria, and a number of other
genera.
The purpose of the present paper is to provide a name
for the West African species that has been known in the
recent literature as Metula clathrata Adams and Reeve,
and to describe another new Metula from the upper
continental slope in the Mozambique channel. Several
additional Indo-Pacific species of Metula, now under
study, can be distinguished only on the basis of their
protoconch, which has already been emphasized by Al-
tena (1949) as a taxonomical character.
SYSTEMATICS
The radula of a species of Metula is figured here for the
first time (figure 1). It is very small for a buccinid (ribbon
25 ;um wide; central tooth 6.5 ^m wide, lateral teeth 12
^m wide) and very similar to the radulae of Ratifusus
and Iredalula figured by Ponder (1968): the central tooth
has a narrow arched basal plate with 3 long, slender, and
equal cusps; the lateral teeth also have a narrow basal
plate and 3 long slender cusps, the outermost one being
longest.
A radula has been looked for, but not found in Metula
amosi Vanatta, 1913, and M. cumingi (Adams, 1853);
several specimens were examined in each case by A.
Waren (personal communication). I do not consider pres-
ence or absence of this very reduced radula to be of
generic importance.
The radula (figure 2) of Pisania striata (Gmelin, 1791),
tvpe species of Pisania, is 150 ^m wide; it differs in
having a central tooth with a large square basal plate
and 5 short and broad cusps, the outermost 2 being small-
er; the lateral teeth are more strongly built, with 3 un-
equal cusps.
In view of the small variation of radular types in buc-
Page 150
THE NAUTILUS, Vol. 102, No. 4
cinids, this difference is remarkable and probabK indi-
cates that the two genera are not closeK- related. What-
ever rank (subfamily or tribe) the Pisania group is given
in Huccinidae, additional research is needed before the
Colubraria group is considered a mere s>nonym of it.
DESCRIPTIONS
[For a diagnosis of the genus see Altena (1949) as An-
temetula]
Metula crosnieri new species
(figures 3-5, 9)
Description: Shell solid, fusiform, consisting of 2.5 pro-
toconch and 6.2 teleoconch whorls. Spire high, body whorl
comprising 64% of total shell height. Protoconch (figure
9) with large nucleus of two smooth convex whorls
abruptK demarcated from teleoconch. Teleoconch whorls
convex, without sutural ramp, with deeply impressed
suture. Sculpture of raised spiral cords and curved op-
isthocline axial ribs; cords and ribs producing beaded
intersections and pitted intervals. Eight spiral cords on
spire whorls; 4 minor, intermediate cords on penultimate
whorl; about 18 cords above periphery of body whorl,
principal and secondary cords alternating rather regu-
larly, and 33 cords below periphery, of which about 15
are set close together in siphonal region. Axial ribs about
equal in strength to spiral cords on spire whorls; weaker
on penultimate whorl, with main sculpture being spiral
on body whorl. In addition to axial ribs, several incre-
mental scars are obvious, especially on body and pen-
ultimate whorls. Aperture ovate; inner lip thin, smooth,
adherent to body whorl, thicker in columellar region;
outer lip bearing 12 very weak teeth that do not corre-
spond with position of external spiral cords; two most
apical teeth slightly stronger. Peristome thickened, form-
ing broad varix, also covered by spiral cords. Siphonal
canal long, broad, widely open, and recurved. Fasciole
indistinct.
Colour light tan, with a very indistinct darker spiral
band at periphery of body whorl; incremental scars light-
er; aperture white.
Dimensions of the holotype: Height 51 3 mm, width
19.4 mm; height of the aperture 24.5 mm, width 8 mm;
height of the body whorl 33 mm.
The largest paratype is 55.4 mm high.
Type lf>cality: Mozambique channel, SW Madagascar,
off Baie de Faiiemotra, 22°15'S, 43°05'E, 470-475 m.
Type material: Holotype and paratype 1 (MNHN) from
the type locality, collected by A. Crosnier, Dec. 2, 1973
aboard R.V. "N'auban"; paratype 2 (MNHN), Mozam-
bique channel, SW Madagascar, 22°17'S, 43°04'E, 400-
450 m, collected by R. v. Cosel, Nov. 30, 1986 aboard
trawler "Mascareignes III".
Distribution: Known only from llic type material, off
SW Madagascar.
Figures 1, 2. Scanning electron niicrographi ol raciulac I.
Metula africana new species, scale bar lO/iiii. 2. Fisania striata
((Jnielin. 1791), scale bar .50 nm.
Remarks: Metula crosnieri has remarkably convex
whorls and a deep suture when compared with its con-
geners. The protoconch (figure 9) indicates non-planc-
totrophic larval development. The combination of these
two characters distinguishes it from all known Indo-Pa-
cific Metula.
In the Atlantic, Bartschia significans Rehder, 1943,
type species of Bartschia Rehder, 1943, has even more
convex whorls, and a multispiral protoconch. I cannot
find characters other than the convexit\- of the whorls
that sharply distinguish Bartschia from Metula, and con-
clude that Bartschia should be considered at most a sub-
genus of Metula.
p. Bouchet 1988
Page 151
I am naming this species after my colleague Alain
Crosnier, who first collected it during a survey of deep
water shrimp populations oft Madagascar.
Metula ajricana new species
(figures 1, 6-8, 10)
Metula clathrata Knudsen, 1956;39, plate 1, figure 1; non M.
clathrata (Adams and Reeve, 1850).
iKin Metula knudseni Kilburn, 1975:592 (replacement name
for Buccinum clathratum Adams and Reeve, 1850: see
Emerson, 1986).
Description: Shell solid, fusiform, consisting of 7 teleo-
conch whorls. (Protoconch of holotype partly broken, on
a paratype consisting of large nucleus and 1.5 smooth
convex whorls, figure 10.) Teleoconch whorls convex,
with faint but distinct sutural ramp on early whorls; ramp
indistinct on penultimate whorl, body whorl evenly con-
vex. Teleoconch sculpture of raised spiral cords and
slightly curved opisthocline axial ribs; intersections dis-
tinctly beaded in sutural ramp area, only slightly nod-
ulous below ramp. Spiral and axial sculpture of similar
strength on early whorls; later spiral sculpture gradually
dominates. Nine primary spiral cords per whorl on spire
whorls, first (adapical) and third cords stronger, limiting
sutural ramp. Fine spiral threads present between cords
on penultimate and bod\ whorl, several eventually de-
veloping into secondary spiral cords, with one present in
sutural ramp area of body whorl. About 22 primary cords
below periphery of body whorl, plus another 10 in si-
phonal area. Incremental scars distinct; growth lines very
distinct between axial ribs. Aperture ovate, narrow. Inner
lip thin, smooth, adherent to body whorl, thicker in col-
umellar region. Outer lip regularly convex, bearing small
teeth that correspond at least partly with position of spiral
cords on peristome; a group of 5 teeth forms a small
callus in apical portion of outer lip, delimiting small anal
canal. Peristome forming a thickened varix, over which
spiral sculpture extends. Siphonal canal short, narrow,
open, only slightly recurved.
Ground colour of shell beige cream, with 3 brown
spiral bands; 3 adapical cords brown, interval between
them beige cream. Below, uniformly beige band extends
over next 2 spiral cords, occupies central position on spire
whorls. Darkest band occupies next 4 spiral cords and
intervals between them; this band occupying suprasu-
tural position on spire whorls, and a central position on
body whorl. Below brown band a second beige band
extends over 4 spiral cords; next 2 cords brown, with
brown colour fading towards base of shell. Aperture cream
colored.
Dimensions of the shell: Height 54.5 mm, width 17.5
mm; height of the aperture 27 mm, width 7 mm; height
of the body whorl 35 mm.
Type locality: Off Saint Louis, Senegal, in 300-600 m.
Type material: Holotype (MNHN), paratype 1 (MNHN)
and paratype 2 (AMNH 198755), all from the type lo-
cality, collected by M. Pin on the trawler "Louis Sauger".
Figures 3-8. Back, side, and front views. 3-5. Metula cros-
nieri new species, holotype, 51.3 mm. 6-8. Metula ajricana
new species, holotype, 54.5 mm.
Other material examined: SENEGAL: off Saint Louis,
"deep water", 4 shells (MNHN, leg. M. Pin), 10 shells
(coll. M. Pin, Dakar); IVORY COAST: off Abidjan, 1
shell, coll. Marche-Marchad (MNHN), and 1 shell, P. Le
Loeuff coll. (MNHN); EQUATORIAL GUINEA: Atlan-
tide Sta. 120, 02°09'N, 09°27'E, 250-850 m, 1 specimen
Page 152
THE NAUTILUS, Vol. 102, No. 4
Figures 9. 10. Protoconchs. 9. Metula crosnieri. 10. Metula
ajricana. Scale line 1 mm.
(Kiiudsen, 1956) (ZMC); CONGO: West of Pointe-Noire,
100 m, 1 shell, A. Crosnier coil. 1962 (ANSP 333810).
Distribution: Deep continental shelf and upper slope
of West Africa, from Senegal to Congo.
Remarks: Metula africana has been figured three times
in the literature: by Knudsen (1956: plate 1, figure 1)
under the name Metula clathrata; by Emerson (1986:
figures 4, 5) as Metula sp.; finally by Kaicher (1987: card
4851) as Metula sp. I refer to Emerson (1986) who re-
viewed the nomenclature of this West African Metula
and concluded that it represents a new species.
The protoconch (figure 9) indicates non-planctotro-
phic larval development.
There may be clinal variation in adult size along the
West African coast: The 18 shells examined from Senegal
have a mean height of 41.7 mm; the 2 shells from Ivory
Coast, although fully adult, measure 32 and 36 mm; that
from Equatorial Guinea is 30.6 mm high, and that from
Congo 25.5 mm.
The tiny rachiglossate radula (figure 1) was prepared
from the specimen taken during the .-Ktlantide expedition
(ZMC).
The close connection between Mediterranean Pliocene
and Recent West African marine faunas has been dem-
onstrated in a number of paleontological papers (for re-
cent reviews see Ruggieri, 1967; Marasti and Raffi, 1979;
Sahelli and Taviani, 1984). With a single species of Me-
tula present in the Pliocene of Italy and a single Recent
species in West Africa, it is reasonable to assume that
Metula mitraeformis (Brocchi, 1814) (for figures and
references see Pelosio, 1966) is the direct ancestor to M.
africana. I have examined material of the fossil species
and found it to differ from the Recent one by its much
weaker axial sculpture, which on the body and penul-
timate whorls is limited to growth lines. In M mitrae-
formis, there is a very broad sutural ramp that extends
over the adapical third or half of early teleoconch whorls,
and becomes obsolete on the penultimate and body whorls;
the siphonal canal is also broader.
Because he was mistaken about the identity, type lo-
calit\, and distribution of M. clathrata, Grecchi (1978)
speculated that the presence of its presumed ancestor M.
mitraeformis in the Mediterranean Pliocene was an in-
dication of Mediterranean-Indo-Pacihc connections in
the lower Pliocene after the Messinian salinity crisis.
With M. clathrata now known to be a West American
species distinct from M. africana, the history of the M.
mitraeformis-africana lineage can not be taken as an
indication of such connections. This lineage probably has
an Eastern .Atlantic history dating to the Miocene; al-
though the Neogene West African fossil record is lacking,
it is far more probable that M. mitraeformis reinvaded
the Pliocene Mediterranean from West Africa rather than
from the Indo-Pacifie through unproven maritime con-
nections.
ACKNOWLEDGEMENTS
I thank Dr. A. Waren who prepared the radula of Metula
africana and commented on the manuscript. A. Crosnier,
M. Pin, and R. von Cosel collected most of the material
cited in this paper. Dr. W. Emerson generousK refrained
from describing M. africana when he learned 1 was
working on it. Photography is b\ P. Lozouet.
LITERATURE CITED
Alteiia, C O van R 1949. The genus Antemetula Rehder
in the Indo-West Pacific area, with the description of two
new fossil species. Bijdragen tot de Dierkdunde 28:385-
393.
Beu, A. G. and P. A. Maxwell. 1987. A revision of the fossil
and living gastropods related to Plesiotriton Fischer, 1884
(Family Cancellariidae, Subfamily Plesiotritoninae n.
subfam.) with an appendix: genera of Buccinidae Pisani-
inae related to Colubraria Schumacher. 1817. New Zea-
land Geological Survey Paleontological Bulletin 54:1-140,
Cernohorsky, W. 1971. Indo-Pacific Pisaniinac and related
buccinid genera. Records of the .\uckland Institute and
Museum 8:137-167.
Emerson, W. K. 1986. On the type species of Metula H. &
\. Adams, 1853: Buccinum clathratum A. .\dams and
Reeve, 1850. The Nautilus 100(1 ):27-:30
Grecchi, G. 1978. Problems connected with the recorded
occurrence of some mollusks of Indo-Pacific affinit\ in the
Pliocene of the Mediterranean area Rivisla Italiana di
Paleonlologia 84:797-812.
Houbrick, R. 1984. A new "Metula" species from the Indo-
West Pacific. Proceedings of the Biological Society of
Washington 97(2):420-424.
Kaicher, S. D. 1987. Card catalogue of world-wide shells.
Pack 48: Buccinidae, part 3 Published by the author, St.
Petersburg.
Killnirn, R N 1975. Taxonomic notes on South African ma-
rine Mollusca (5): including descriptions of new ta\a of
Rissoidae, Cerithiidae, Tonnidae, Cla.ssididae, Buccinidae,
p-asciolariidae, Turbinellidae, Turridae, Architectonici-
dae, Epitoniidae, Limidae, Thraciidae. Annals of the Natal
Museum 22(2):577-622.
Knudsen, J. 1956. Marine prosobranchs of tropical West Af-
rica (Stenoglossa). Atlantide Report 4:7-110.
Marasti, R and S. Raffi 1979 Observations on the paleocli-
matic and biogeographic meaning of the Mediterranean
p. Bouchet 1988
Page 153
Pliocene molluscs. State of the problem. 7th Intenuilinnal
Congress on Mediterranean Neogene, Athens, 8 unnum-
bered pages.
Olsson, A. and F. Bayer. 1972. American Metulas (Gastro-
poda: Buccinidae). Bulletin of Marine Science 22(4):900-
925.
Pelosio, G. 1966. La Malacofauna dello stratotipo del Tabi-
aniano (Pliocene inferiore) di Tabiano Bagni (Parma). Bol-
lettino della Societa Paleontologica Italiana 5(2):l()l-183.
Ponder, W. 1968. Anatomical notes on two species of the
Colubrariidae. Transactions of the Roval Society of New-
Zealand, Zoology 10(24):217-223.
Ponder, W. 1973. The origin and evolution ot the Neogas-
tropoda. Malacologia 12(2):295-338.
Ponder, W. and A Waren 1988 Classification of the Cae-
nogastropoda and I leteroslropha — a li.sl of the family-group
names and higher taxa. Malacological Review, supplement
4:286-32-1.
Ruggieri, G. 1967. The Miocene and later evolution of the
Mediterranean Sea. In: Adams, C. G. and D. V. Ager (eds.).
Aspects of Tethyan biogeography. The Systematics Asso-
ciation, London. P. 283-290.
Sabelli, B. and M. Taviani. 1984. The paleobiogeographic
distribution of the Mediterranean benthic mollusks and
the Messinian salinity crisis or where did the mollusks go?
Annales de Geologic des Pays Helleniques 32:263-269.
THE NAUTILUS 102(4):154-158, 1988
Page 154
A New Species of Intertidal Terebra from Brazil
Kurl Auffenberg
Malacolog> Division
Florida Museum of Natural History
University of Florida
Gainesville, FL 32611. USA
Harry G. Lee
709 Uomax Street
Jacksonville, FL 32204, USA
ABSTRACT
Terebra imitatrix new species is described from northern Brazil
and compared to morphologically similar species of Hastnla.
The three known terebrid feeding types are briefl\ reviewed.
Dissection of the foregut of this new species revealed characters
that overlap two of the major feeding types.
Key words: Gastropoda; Terebridae; Terebra; anatom\ : Bra-
INTRODUCTION
While compiling distributional records of the West At-
lantic Hastula cinerea species group, the junior author
located an unidentified lot of 23 specimens from Brazil
in the Academy of Natural Sciences of Philadelphia
(ANSP 299957). These specimens were compared to the
known terebrids from the Atlantic and were found to
belong to a distinct undescribed species. An additional
lot of the same species was subsequently found in the
American Museum of Natural History (AMNH 129280).
Matthews et al. (1975:99, fig. 31) in their treatment of
Hastula ciiterea describe and illustrate a protoconch con-
sistent with this new species. The Brazilian specimens of
Hastula salleana (Deshayes, 1859) figured by Rios (1970:
123, pi. 47, 1975:127, pi. 38, fig. 560, 1985:131, pi. 45)
are probably referable to this new species, but the figured
specimen was unavailable for examination. This paper
describes this new species and compares it with mor-
phologically similar species of Hastula. A description of
the foregut anatomy reveals that it does not conform to
any of the three known feeding types, which are re-
viewed herein.
MATERIALS AND METHODS
Only shells possessing 10 or more teleoconch whorls and
with intact protoconchs and apertures were measured
with Vernier calipers. All dissections were made under
a Wild M-5 dissecting micro.scope and line drawings
made with the aid of a camera lucida. Description and
discussion of the anatomy is limited to the foregut due
to poor preservation in the upper whorls. Anatomical
and protoconch measurements were made with an ocular
micrometer and converted to millimeters. Terminology
follows that of Miller (1970, 1971). Two preserved but
completeK retracted adult specimens of the new species
(ANSP 299957), Hastula maryleeae Burch (UF 113539)
and Hastula salleana (Deshayes) (UF 48197, 113540),
were dissected from each lot. Two dried-in specimens of
Hastula maryleeae Burch (T. Bratcher collection) were
rehydrated in a weak solution of potassium h\dro.\ide,
transferred into water, and dissected. The radular sac
was extracted and dissolved in a weak solution of potas-
sium hydroxide. Radular teeth were individual!} mount-
ed on scanning electron microscope specimen stubs cov-
ered with double-sided tape. Micrographs were made
with a Hitachi 5-415.^ scanning electron microscope.
Repositories of examined specimens are indicated by
the following abbreviations:
AMNH American Museum of Natural Histor\'
ANSP Academy of Natural Sciences of Philadelphia
UF Florida Museum of Natural History
SYSTEMATICS
Family Terebridae Morch, 1852
Genus Terebra Bruguiere, 1789
Terebra imitatrix new species
(figures 1-6, 8, 9, table 1)
Description: Shell (figures 1-3, 6) medium in size,
broadening anteriorK ; color variable, ranging from
banded, or cream, to purplish-brown; teleoconch whorls
10-12; sides flat to slightly convex. Protoconch whorls
1.5-2.0; glassy, transparent (figure 6). .\xial sculpture of
close-set, recurved ribs of \ariable strength, generally
distinct near the suture, becoming obsolete anteriorly;
major axial ribs 34-57 (x = 41.6) on the penultimate
whorl. Spiral sculpture of very faint microscopic incised
lines most distinct in the intercostal spaces on the upper
whorls, obsolete on later whorls, rarely cro.ssing the axial
ribs; spiral rows of pits absent. Last whorl with obsolete
axial ribs and spiral incised lines; color pattern variable,
typicalK of five diffuse color bands: one white pre-sutural
band usiialK w ith distinct brown spots that become ob-
solete toward apertural lip, one broad bluish-black zone
at shoulder, one pinkish band at periphery, one purplish-
K. Auffeiiberg and H. G. Lee 1988
Page 155
4
Figures 1-3. Terebra imilatrix new species. 1. Holotype, ANSP 299957 (27.5 mm shell length) 2, 3. Paratypes, ANSP 369293,
all from Rio Grande do Norte, Brazil, -sand island at mouth of the inlet at Areia Branca.
brown band below periphery, and one white basal zone.
Columella brown, centrally concave and slightly re-
curved; rounded rib present on anterior edge. Parietal
callus thin, transparent to light brown. Fasciole white to
bluish-gray with distinct white rib. Anterior siphonal
notch moderately broad, straight. Aperture light to dark
brown with white band.
External anatomy: Animal cream-colored with no ap-
parent pattern in alcohol-preserved specimens. Oper-
culum corneous, small, thin. Eyes on short, broad eye-
stalks. Labial tube large, spoon-shaped (figure 4). Anterior
end of labial tube terminating in thick muscular lips
bounding mouth slit; well-developed sphincter lacking.
Foregut anatomy: Labial cavity large, dominated by
massive (3.7 mm), extendable, muscular organ (accessory
feeding apparatus; Miller, 1970, 1971) that tapers an-
teriorly; muscular organ with two rows of papillae on
ventral side (figure 4); attached posteriorly to the left
side of the cephalic hemocoel by connective tissue; re-
tractor muscle large, originating in foot below and slight-
ly posterior to anterior siphon, passing through cephalic
hemocoel. Buccal tube muscular, short (0.9 mm), taper-
ing anteriorly. Buccal cavity small (0.6 mm), rounded.
Pre-ganglionic esophagus enters buccal mass posteriorly
(figure 4). Salivary glands not located. Radular sac blade-
shaped, small (0.8 mm). Radular caecum (0.4 mm) with
well-developed groove running posteriorly, two distinct
bulbs anteriorly (figure 5). Radular organs attached to
right side of anterior portion of buccal cavity by very
short duct (0.1 mm). Two rows of radular teeth situated
obliquely in radular sac and caecum (figure 5). Radular
teeth about 30; 0.1-0.2 mm in length, slightly curved,
not barbed (figures 8, 9). Poison gland long (6.2 mm),
extremely convoluted, entering right side of buccal cav-
ity slightly behind radular sac (figure 5). Poison bulb
small (0.9 mm), weak, seemingly vestigial, lying at ven-
tro-posterior end of cephalic hemocoel.
Etymology: From the Latin feminine noun meaning
one who imitates, in reference to the deceptively close
resemblance of the shell to that of Hastula cinerea (Born,
1778), with which it occurred in the type lot.
Table I. Measurements (mm) of shell characters of Terebra
imitatrix new species and Hasttila rnaryleeae Burch
Species
Character
X
Range
SD
T. imitatrix
Shell length
27.3
24.1-29.7
1.9
Shell width
6.4
5.7-7.4
0.5
Protoconch
0.50
0.46-0.52
0.03
width
H. rnaryleeae
Shell length
18.2
15.0-24.1
2.7
Shell width
4.6
3.7-5.5
0.6
Protoconch
40
0.38-0.42
02
width
Page 156
THE NAUTILUS, Vol. 102, No. 4
•■s be
Figures 4-6. Terebra irnitatrix new species. 4. Diagrammatic dorsal view nf the organs of the foregut 5. Right side of the buccal
tulie showing radular organs and insertion of the poison gland. 6. Protoconch. 7. Hastula manjleeae Burch, protoconch (.-WINH
191815) Scale bars = 1 mm. af, accessory feeding apparatus; be, buccal cavity; bt, buccal tube; It, labial tube; pb, poison bulb; pe,
pre-ganglionic esophagus; pg, poison gland; re, radular caecum; rm, retractor muscle; rs, radular sac.
Type locality: Brazil, Rio Grande do Norte, sand island
at mouth of the inlet at Areia Branca, 04°57'S, 37°08"W,
G. & M. Kline et al., 14 December 1963.
Holotype: ANSP 299957, shell length 27.5 mm, width
6.7 mm.
Paratypes: Paratypes 1-20, ANSP 369293, from the
tvpc locality (15 dry, 5 in alcohol). Paratypes 21-22, UF
115180, from type locality. Paratypes 23-24, AMNH
129280, Brazil, Ceara, Acaraii, 02°53'S, 40°07'W.
Distribution: PresentK know n onl\- from northern Bra-
zil.
Ecology: Based on the locality label of the type lot, this
species is found in sand near or at inlets much like some
populations of Hastula. One specimen of Hastula ci-
nerea (Born, 1778) was found in the type lot and another
(AMNH 129269) from the same locality as the AMNH-
paratypes. Although these two species may occur micro-
s\ mpatrically, we assume there is no trophic competition
due to the strikingly divergent feeding organs (see dis-
cussion).
Comparative remarks: Terelrra irnitatrix is similar in
shell morpholog\- to the West Atlantic Hastula cincrea
group. It can be easily separated from Hastula cinerea
and Hastula salleana by the lack of spiral rows of pits
and the generally more numerous, but less prominent,
axial ribs. .Although similar in size, T. irnitatrix is usually
broader anteriorly . Hastula cinerea and H. salleana have
3.5-4.0 protoconch whorls, while T. irnitatrix has 1.5-
2.0. The typical color pattern of T. irnitatrix is more
distinctK banded with larger and more distinct brown
spots al the suture and a broader white subsutural band.
Terelna irnitatrix and Hastula manjleeae are more dif-
ficult to separate, particularly southern Caribbean pop-
K. Auffenberg and H. G. Lee 1988
Page 157
Illations of H. niarijleeae described as Terehra toba-
goensis Nowell-l'slicke, 1969, now placed in the
s\non\ my of H. manjleeae by Bratcher and Cernohorsky
(1987). Examination of the lectotype (AMNH 195453
designated bv Bratcher and Cernohorsky, 1987:194, pi.
60, fig. 235c)' and paralectotypes (AMNH 191819) of T.
tobagoensis. as well as several other lots from Tobago,
typical specimens of H. manjleeae from Te.xas, and spec-
imens of a weakly ribbed form of H. maryleeae from
the Dominican Republic (see Bratcher and Cernohorsky,
1987, for discussion) revealed consistent shell characters
b\ which the two species may be separated. Anatomical
features are discussed below. Typical H. maryleeae is
easily separated from T. imitatrix by the distinctive shell
shape caused by the enlarged nodes of the axial ribs at
the suture and by the slight crenulations at the sutures
of the upper whorls. Hastula manjleeae has a much
smaller shell than T. imitatrix at the same whorl count
(table 1). The shell of T. imitatrix broadens more an-
teriorK- and has a proportionately slightly larger aper-
ture. The protoconchs of both species have 1.5-2.0 whorls,
however, the protoconch of T. imitatrix is more bulbous
(figures 6, 7; table 1). Typical T. imitatrix also resembles
the West African Hastula aciculina (Lamarck, 1822),
particularK- in color pattern. However, T. imitatrix lacks
the supra-sutural groove and callosity found in H. acic-
ulina (Bouchet, 1982; Bratcher and Cernohorsky, 1987).
DISCUSSION
Miller (1970) proposed a division of the Terebridae into
three major groups based on the anatomy of the foregut
(feeding type) and later published a series of papers
(Miller, 1971, 1975, 1979) on this subject supported by
in-depth life history studies. These data are reviewed
and the species assigned to each group are listed in
Bratcher and Cernohorsky (1987). Type I species have
a long labial tube and a short buccal tube. They do not
possess a radular apparatus or poison organs. This group
is further divided into two subgroups, lA and IB, based
primarily on habitat and pre%' (see Bratcher and Cer-
nohorsky, 1987). Type II encompasses species exhibiting
typical toxoglossan feeding characters. The labial tube
is long and eversible and the buccal cavit\ is relatively
large. The buccal tube is long and retractile. They have
a poison gland and bulb as well as a radular sac containing
two rows of harpoon-like radular teeth. This feeding type
is further divided into two subgroups, types IIA and IIB,
based primarily on habitat and behavior. Type IIA in-
cludes several Indo-Pacific species, as well as the West
Atlantic Hastula discussed in this paper (see Bratcher
and Cernohorsky, 1987). Type III species possess an ac-
cessory feeding organ which grasps prey and pulls it into
the labial cavity. The\' lack a radular apparatus and some
have lost the buccal tube and salivary glands. This group
presentK contains no .\tlantic species, but is represented
b\' several Indo-Pacific taxa (see Bratcher and Cerno-
horsky, 1987).
Terebra imitatrix has very different foregut anatomy
Figures 8, 9. Radular teeth of paratype (ANSP 369293) of
Terehra imitatrix. new species. 8. Whole tooth, 60 x . 9. Tip
of lootli, 200 X.
from the West Atlantic Hastula. We have dissected H.
salleana and H. manjleeae for comparison, and both
possess a Type IIA polyembolic proboscis (Miller, 1970,
1971). Terebra imitatrix has a large spoon-shaped labial
tube, presumably ineversible, a labial cavity dominated
by the accessory feeding apparatus, an extremely short
buccal tube incapable of extending outside the mouth
and the buccal cavity, and the associated radular organs
are minute in comparison to those of Hastula. The ves-
tigial poison gland and bulb are barely recognizable as
such and are considered homologous to the massive poi-
son gland and bulb of Hastula only by the similarity of
position and the site of its entrance into the buccal cavity.
The radular teeth are similar in size and shape to those
of Hastula maryleeae (0.15 mm). The teeth of H. sal-
leana and H. cinerea are larger (0.5 mm) and barbed.
The presence of an accessory feeding apparatus and the
small size of other foregut organs place T. imitatrix near
the group possessing a Type III polyembolic proboscis.
However, the presence of radular and poison organs, such
as occur in T. imitatrix, has not been reported for this
proboscis type. Also, the shell of T. imitatrix is very
different from the species in this group, all of which have
shells with deeply impressed sutures, a sub-sutural groove
and strong to moderate axial sculpture.
The feeding behavior of Hastula cinerea and of H.
inconstans (Hinds, 1844) have been well documented
by Marcus and Marcus (1960) and Miller (1979), re-
spectively. The long eversible labial tube forages for prey
items. A single radular tooth is passed through the labial
tube, held in the tip, and inserted into the prey to fa-
cilitate penetration of the venom. The prey is then in-
gested via the labial tube. Miller (1970) suggests that
species with a Type III proboscis forage by utilizing the
accessory feeding apparatus, and that food items are
passed into the opening of the interior buccal tube. We
do not know if the radular and poison organs are func-
tional in Terebra imitatrix. If so, the radular tooth is
either transferred from the buccal tube to the accessory
feeding apparatus and inserted into prey outside the
body, or once it reaches the buccal tube.
Page 158
THE NAUTILUS, Vol. 102, No. 4
The presence in Terebra imitatrix of the accessory
feeding apparatus recorded in a few Indo-Pacific Tere-
bra and the poison anil radiilar organs of Hastula make
this species unique. Terelna imitatrix may be closely
related to the West African species, Hastula aciculina,
however, this species must be studied anatomically be-
fore this assignment can be verified Generic limits within
the Terebridae are presently p<}ori\ understood with much
overlap in shell and anatomical characters. Additional
anatomical studies are needed to clarify the taxonomic
and e\oiutionar\ significance of the foregut in the Tere-
bridae.
ACKNOWLEDGEMENTS
We wish to thank several people who have assisted us
during this study. Special thanks go to Robert Robertson,
George M. Davis, and Mar) \. (iarback of the .\cademy
of Natural Sciences of Philadelphia for allowing us to
borrow and dissect the specimens on w hich this descrip-
tion is based. William K. Emerson, Harold Feinberg, and
Walter Sage at the .American Museum of Natural Histor\
allowed us to visit their department and borrow several
lots of \ery pertinent material. Twila Bratcher of Hol-
lywood, California kindly provided comparative mate-
rial, as did Eliezar C. Rios (Funda^ao Universidade do
Rio Grande do Sul, Brazil), the late Joseph Rosewater of
the U.S. National Museum, and Thomas Pulley and Con-
stance Boone of the Houston Museum of Natural Science.
Marcos Guianaldo and Armando Garcia, Florida Mu-
seum of Natural History, prepared figures 4-7. The mi-
crographs were made by the senior author with the scan-
ning electron microscope housed in the Department of
Zoology, University of Florida. In addition, we thank
Richard S. Houbrick, William K. Emerson, and Fred G.
Thompson for reviewing earlier drafts of the manuscript.
LITERATURE CITED
Bouchet. P. 1982. Les Terebridae (Mollusca, Gastropoda) de
r.\tlantique Oriental. Bollentino Malacologico, Milano
18(9-12):18.5-2I6.
Bratcher, T. and \V. O. Cernohorsky 1987 Living terebras
of the world .\ monograph of the Recent Terebridae of
the world .Xmerican Malacologists, Inc.. Melbourne, FL,
240 p.
Marcus, E. and E. Marcus 1960. On Hastula cinerea. Bo-
leiitim Faculdade de Filosofia Ciencias e Letras. Univer-
sidade de Sao Paulo, no. 260, Zool. no. 23:25-54, pis. 1-5.
Matthews. H. R., \. C. Dos Santos Coelho, P. De Sa Cardoso,
and M. Kempf. 1975. Notas sobre la familia Terebridae
no Brasil (Mollusca, Gastropoda). .-Vrquivos do Museu Na-
cional. Rio de Janiero 5585-104.
Miller. B .A 1970 Studies on the biologN of Indo-Pacific
Terebridae. Ph D. dissertation. I'niversity of New Hamp-
sliirc. Durham. 213 p.
Miller, B. .\. 1971. Feeding mechanisms in the family Tere-
bridae. .Annual Report of the American Malacological
I'nion for 1970. 1970:72-74.
Miller. B. .A. 1975. The biolog\ of Tereiwa gouWi Deshayes,
1859, and a discussion of the life history similarities among
other terebrids of similar proboscis tvpe Pacific Science
29:227-241.
Miller, B. .\. 1979. The biology of Hastula inconstans (Hinds,
1844) and a discussion of life histor\ similarities among
other Hastula of similar proboscis type. Pacific Science
33(3):289-306.
Nowell-l'sticke, G. W. 1969 .A supplementary listing of new
shells (illustrated). Privately published. St. Croix, Virgin
Islands, 31 p., 4 pis.
Rios, E. C. 1970, Coastal Brazilian shells Funda(,ao Cidade
do Rio Grande, Museu Oceanografico de Rio Grande, Bra-
zil, 255 p.. 60 pis.
Rios, E. C. 1975. Brazilian marine mollusks iconograph\.
Fundagao Universidade do Rio Grande, Centro de Cien-
cias do Mar, Museu Oceanografico, Brazil, 331 p., 91 pis.
Rios, E. C. 1985. Seashells of Brazil Funda^ao Cidade do
Rio Grande, Fundagao Universidade do Rio Grande, Mu-
seu Oceanographico, Brazil, 329 p , 1421 pis.
THE NAUTILUS 102(4):159-163, 1988
Page 159
Notes on the Biology and Morphology of Margaritifera hembeli
(Conrad, 1838) (Unionacea: Margaritiferidae)
Douglas G. Smith
Museum of Zoology
Universit\ of Massachusetts
Amherst, MA 01003-0027, USA
ABSTRACT
The freshwater pearl mussel Margaritifera hembeli is found
onK in the Red River basin and a few nearby drainages in
Louisiana. Though of concern to conservationists because of its
declining numbers, M. hembeli remains virtually unknown with
respect to its anatomy and biology. The species contains all the
anatomical characters that typify margaritiferid species. The
sexes are separate and the gonads show a definite seasonality
in activity. Gametogenesis is pronounced in specimens collected
in the fall, followed by degeneration of reproductive tissues in
the late winter through to late spring. It is concluded, on the
basis of observed gonadal activity, that oviposition and spawn-
ing take place between late November and late January Char-
acters are evident in the morphology of the visceral nervous
svstem and the stomach of M. hembeli that clearly distinguish
M. hembeli from M. marrianae and other eastern North Amer-
ican margaritiferid species. A distinct relationship between M.
hembeli and M. marrianae, however, is suggested by the mu-
tual occurrence of lateral hinge teeth and a corrugated surface
of the posterior portion of the shell. Due to the lack of knowl-
edge of the anatomy and biology of other margaritiferid species,
especially those living in Asia, it is premature to suggest rela-
tionships lietween M. hembeli and other described margari-
tiferid species, particularly those with lateral hinge teeth.
Key uords: Margaritifera: anatomv, reproduction. North
.America
INTRODUCTION
The North American freshwater mussel Margaritifera
hembeli (Conrad, 1838) was once believed to comprise
two geographically discontinuous populations ta.xonom-
ically linked b\ vague similarities of the shell. Johnson
(1983) separated the two populations taxonomically by
describing the Alabama group as M. marrianae, thus
restricting the M. hembeli group to Louisiana. His de-
scription included characters of the shell only, principally
the degree of sculpturing on the shell surface and the
shape of the ventral shell margin. Additional concho-
logical differences between the Louisiana and Alabama
populations were noted by Smith (1983) who pointed out
dissimilarities in the mantle-shell attachment scars on the
inner nacreous surface.
Margaritifera hembeli probably had a more extensive
range in the Red River drainage as indicated by museum
records, particularly a specimen in the American Mu-
seum of Natural History (AMNH 193786) from the Red
River in Arkansas. During the present century, however,
the range has contracted considerably due to deterio-
rating environmental conditions. The present range is
limited to the Bayou Teche drainage (Vidrine, 1985) and
a single stream in the Red River drainage. The drastic
reduction in range has elicited concern from the federal
government, which has provided protection for the re-
maining populations (Stewart, 1988).
Despite increased concern for M. hembeli, very little
is known about this species other than the characteristics
of its shell. Ortmann (1912) provided a brief description
of the anatomy of M. "hembeli, ' but the specimens upon
which he based his description came from Alabama and,
therefore, are appropriately referred to M. marrianae.
Hence the anatomy of M. hembeli remains undescribed,
and nothing is known about its biology. The present study
provides some information on gonadal activity and de-
tails of the anatomy of the stomach and nervous system.
Comparisons are made with other North American mar-
garitiferid species, including M. marrianae, as studied
by Smith (1979a, 1980, 1986, and unpublished).
MATERIALS AND METHODS
A total of 43 partially or completely relaxed, preserved,
specimens were studied. All were collected on various
dates from 1973 to 1986 from Brown Creek, Gardner,
Rapides Parish, Louisiana. Specimens had been fixed in
10% formalin, washed in water, and stored in 50% iso-
propyl alcohol. Five specimens lacked information on
date of collection and were utilized for dissection pur-
poses only. The remaining lots were used for histological
investigations of gonadal activity and sexual character-
istics, as well as for anatomical studies. The collection
dates and numbers of specimens used in the study of
gonadal activity were as follows: October 1, 1973 (1
specimen); October 5, 1974 (4 specimens); February 22,
1975 (8 specimens); March 28, 1975 (4 specimens); April
25, 1975 (3 specimens); June 21, 1975 (2 specimens);
March 30, 1986 (5 of 16 specimens).
Page 160
THE NAUTILUS, Vol. 102, No. 4
Figure 1. Photomicrograph of a transverse section through
the gill of Margaritifera hcmheli showing the interlamellar
junction (ILJ), 30 x
Portions of the viscera of each specimen were infil-
trated with paraffin, sectioned at seven micrometer thick-
ness, and stained with Ehrlich's hematoxylin and eosin.
Some sections were stained in a picro- ponceau connective
tissue stain following the method described in Humason
(1979:147). At least five slides were prepared for each
specimen. The barren gills of two specimens and a por-
tion of the posterior mantle lobe of one specimen were
also sectioned in a similar manner and stained with picro-
ponceau connective tissue stain.
Dissections were undertaken on the stomach and vis-
ceral nervous system of eight specimens. Three speci-
mens were investigated for gro.ss morphology of the gills,
nervous system, and excretory system. The method of
dissection and exposition of specific internal organs, and
the terminologv used to describe various organ compo-
nents, follows Smith (1980, 1986).
.\11 material relevant to this study has been cataloged
in the invertebrate collections of the Museum of Zoology
(Nos. MO. 1643-1645), University of Massachusetts, Am-
herst, Massachusetts.
RESULTS
Gross Anatomy
Anterior and posterior adductor muscles subequal, foot
musculature and associated pedal and retractor muscles
well developed Cerebral and pleural ganglia fused, kid-
ney with both glandular and non-glandular chambers,
renal pore and gonopore closely set but clearly separate.
Labial palps falcate and large, gills or demibranchs la-
mellar, inner gill larger than outer gill, both inner and
outer gills free from mantle posterior to pallial line. Both
EX
^.^:
^d.r't-'^r'
Figure 2. Photomicrograph of a transverse section through
the posterior portion of the mantle of Margaritifera hembeli
showing the partial!) contracted diaphragmatic septum (DS),
which separates the exhalent (EX) and inhalent (IN) chambers,
SO X.
lamellae of each gill held together b> solid, separate
interlamellar junctions (figure 1, ILJ), lined with squa-
mosal epithelium and composed of loose connective tis-
sue and fine fibers that are more muscular appearing
than collagenous (in M. margaritifera. see Smith, 1979a).
Interlamellar junctions for the most part arranged in
oblique rows in typical margaritiferine fashion (Ort-
mann, 1912; Smith and Wall. 1983: fig. lb), similar to
gills of A/. niarr!ana<'(Ortmann, 1912:235). Gill junctions
are somewhat patternless along lower margin and an-
terior and posterior extremities of each gill plate.
Mantle lobes free all around, no indication of division
of exhalent region of mantle margins into .separate anal
and supra-anal apertures. Inhalent and exhalent cham-
bers separated posteriorly by diaphragmatic septa (figure
2, DS), and though not observed in the living animal,
are presumed to function similar to M. margaritifera
(Smith. 1980) Inhalent margin of mantle with densely
pigmented papillae, exhalent region pigmented, with
crenulate margin.
Gonadal Activity and Sexuality
.All animals examined histologicalK were sexually ma-
ture, including several small specimens ranging in shell
length from 49 to fj9 mm (believed to be from 6 to 9
D. G. Smith 1988
Page 161
'f
^
^i V
>*.. s'
Figures 3-6. Photomicrographs of histological sections through male and female gonads of Margaritifera hembeli. 3. Gonad of
a male specimen of M. hembeli collected in Februar\, following spawning of gametes, 65 x. 4. Gonad of a female specimen
collected in February following spawning, 65 x. 5. Gonad of a male specimen of M. hembeli collected in October and showing
male gametes which have filled the entire gonadal stroma, 100 x . 6. Gonad of a female specimen of M. hembeli collected in
October and containing fully developed ova, 100 x .
years old on the basis of shell annuli). This would suggest
that M. hembeli matures at an earlier age than North
American M. margaritifera (Smith, 1979b). No evidence
of hermaphroditism was observed.
Although no gravid females were among the available
specimens, a specific reproductive cycle was indicated
by the gonads of sectioned specimens. Animals collected
in Februarv', March, and April showed characteristic post-
spawning features (figures 3, 4) including partial or com-
plete occlusion of gonadal acini by granules, presumed
pycnotic cells, and unspawned gametes in various stages
of development or cytolysis. Animals collected in June
showed little change from April specimens indicating
that complete resorption of reproductive tissues, corre-
sponding to an undifferentiated stage of non-reproduc-
tive activity, apparently does not occur. By early Octo-
ber, gonadal activity is resumed and sex cells, including
the latter stages of spermatogenesis and oogenesis, are
abundant within all observed acini (figures 5, 6). It is
therefore hypothesized that the oviposition of eggs into
marsupial demibranchs and the spawning of male ga-
metes takes place sometime between late November and
late December with release of larvae occurring in late
December or January. Based on examined specimens,
there is no evidence that production of glochidial larvae
takes place at any other part of the year.
Visceral Nerve Anatomy
In M. hembeli, the first pallial bifurcation (figure 7, BI)
of the posterior nerve is well anterior of the mantle nerve
separation, usually arising from the visceral ganglion it-
self at a point near the origin of the posterior nerve cord
(figure 7, PNC). Some variation exists in the position of
this bifurcation relative to the visceral ganglion. The
overall pattern differs from that observed in other eastern
North .American margaritiferids including A/, marrian-
ae. in which the first bifurcation is normally posterior of
the visceral ganglion, but similar to that observed in
Cumberlandia monodonta (Smith, 1980, unpublished
data). Moreover, in both M. hembeli and M. marrianae,
the accessory nerve, which is typically present in M.
Page 162
THE NAUTILUS, Vol. 102, No. 4
VG
PNC
OES
Figures 7-9. .\natoniy of the visceral nervous system and
stomacli of Margarilifcra hcmbeli. 7. Visceral ganglion (VCJ)
of M. hemheli. 20 x. 8. Morphology of the stomach roof of
M. hembeli, 5 x . 9. Morphology of the stomach floor of M.
hemheli. 5 x . APR, anterior protractor muscle; ASA, anterior
sorting area; .ASR, anterior sorting area of roof; BI, first pallial
bifurcation; BN, branchial nerve; C, commissure to cerebro-
pleural ganglion; MN, mantle nerve; PNC, posterior nerve cord;
OES, esophagus; PSf\, posterior sorting area; HD, right duct;
RSA, right side sorting area; T, major typhlosole.
margaritifera and C. monodonta (Smith, 1980), is almost
aKva> s absent, being observed only once in M. marrianae
(Smith, unpublished data).
Stomach Anatomy
In addition to the posterior sorting area, which is typical
of other eastern North American margaritiferids, the
stomach roof of M. hembeli contains a well developed
anterior sorting area and a continuation of the posterior
sorting area extending along the right side of the stomach
roof and separated from the anterior sorting area by a
ridge (figure 8).
The floor of the stomach interior (figure 9) is char-
acterized by an anterior sorting area that is somewhat
similar to M. marrianae (Smith, 1986; fig. 4b); however,
the right side sorting area of M. hembeli is completely
unlike ,V/. marrianae in that a distinct platform is absent
in A/, hembeli and the sorting ridges run primariK lat-
eralK , rather than in the anterior-posterior pattern seen
in M. marrianae. A groove sets off the right side sorting
area from the anterior sorting area in A/, hembeli and
the morphologN of the stomach Door sorting areas of M.
hembeli can be considered more similar to M. marga-
ritifera in overall appearance than to other eastern North
.\merican species.
DISCUSSION
As demonstrated by anatomical investigations of the vis-
ceral nervous system and the stomach (this studv), and
conchological differences discussed elsewhere (Johnson,
1983; Smith, 1983), A/, hembeli and A/, marrianae clear-
1\ represent distinct lineages within the genus Marga-
ritifera. Although both species have lateral teeth, these
teeth are also present in M. auricularia (southern Europe)
and A/, laosensis (southeast .\sia). Lateral teeth may
therefore represent structures that have arisen separately
in different "stocks of margaritiferid species, or ma> be
symplesiomorphies indicative of an ancestral relation-
ship.
The presence of a corrugated surface of the posterior
slope of the shell (weakK expressed in A/, hembeli) and
the close geographical proximit\ of these two species are
the strongest lines of evidence indicating a relationship
between them. Ne\ ertheless, the pattern of \isceral nerve
bifurcation and stomach morphologv in A/, hembeli and
M. marrianae show interspecific variation as great as
that observed between either of these two species and
M. margaritifera or C. monodonta. Biochemical data of
the sort de\eloped for M. margaritifera. C. monodonta,
and A^. hembeli b> Davis and Fuller (1981) is not avail-
able for M. marrianae, precluding a comparison of ge-
netic distances among the four species. However, the
available biochemical data i Davis and Fuller, 1981),
combined u ith anatomical information presented in this
stud) and elsewhere (Smith, 1980, 1986), clearK supports
the concept that the North .American species of the Mar-
garitiferidae have been isolated from one another for a
considerable period of time (Smith, 1976).
ACKNOWLEDGEMENTS
I thank Malcolm F. Vidrine, Mark E. Gordon, and James
E. Williams for gracioiisK- supplying me with the pre-
served specimens used in this studv. Mark E. Gordon
kiiulK read an early draft of the paper.
D. G. Smith 1988
Page 163
LITERATURE CITED
Conrad, T. A. 1838. Monography of the family Unionidae,
or naiades of Lamarck (fresh water bivalve shells) of North
America, Numbers 10, 11. Philadelphia, PA, p. 81-102
Davis, G. M. and S. L. H. Fuller. 1981. Genetic relationships
among Recent Unionacea (Bivalvia) of North ,\merica.
Malacologia 20:217-2.53.
Humason, G. L. 1979. .Animal tissue techniques, 4th ed W .
H. Freeman & Co., San Francisco, 661 p.
Johnson, R. I. 1983. Margaritifera marrianae, a new species
of Unionacea (Bivalvia: Margaritiferidae) from the Mo-
bile-.Alabama-Coosa and Escambia River systems, Ala-
bama. Occasional Papers on Mollusks (Harvard Universitv)
4:299-304.
Ortmann, .\. E. 1912. Notes upon the families and genera of
the najades. .Annals of the Carnegie Museum 8:222-.365.
Smith, D. G. 1976. The distribution of the Margaritiferidae:
a re\iew and a new synthesis Bulletin of the .American
Malacological Union 1976:42 (abstract).
Smith, D. G. 1979a. Marsupial anatomy of the demibranch
of Margaritifera margaritifera (Lin. ) in northeastern North
America (Pelecypoda: Unionacea). Journal of Molluscan
Studies 4.5:39-44.
Smith, D. G. 1979b Se.xual characteristics of Margaritifera
margaritifera (Linnaeus) populations in central New En-
gland. Veliger 21:381-383.
Smith, 1) (; 1980 Anatomical studies on Margari/i/era mar-
garitifcra and Cumherlandia monodonia (Mollusca: Pe-
lec> poda: Margaritileridae). Zoological Journal of the Lin-
nean Society (i9:257-270.
Smith, D. G. 1983. On tlie so-called mantle muscle scars on
shells of the Margaritiferidae (Mollusca: Pelecypoda). with
observations on mantle-shell attachment in the Unionoida
and Trigonioida. Zoologica Scripta 12:67-71.
Smith, D. G. 1986. The stomach anatomy of some eastern
North .American Margaritiferidae (Unionoida: L'niona-
cea). .American Malacological Bulletin 4:13-19.
Smith, D. G. and W. P. Wall. 1983. The Margaritiferidae
reinstated: a reply to Davis and Fuller (1981), "Genetic
relationships among Recent Unionacea (Bivalvia) of North
America." Occasional Papers on Mollusks (Harvard Uni-
versity) 4:321-330.
Stewart, J. H. 1988. Endangered and threatened wildlife and
plants; final endangered status for the Louisiana pearl shell
"Margaritifera hembeli." Federal Register 53:3567-3570.
\'idrine. M. F. 1985. Fresh-water mussels (Unionacea) of
Louisiana; a zoogeographical checklist of post-1890 rec-
ords. The Louisiana Environmental Professional 2:50-59.
THE NAUTILUS 102(4): 164-166, 1988
Page 164
The Two Printings of J. F. Gmelin's Systema Naturae,
13th Edition (1788-96)
Alan R. Rabat
Deparlmenl of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138, USA
Richard E. Petit
Research Associate
Department of Invertebrate Zoology
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560, USA
One of the more important 18th century reference works
in systematics is J. F. Gmelin s l.'3th edition of Linnaeus'
Systema Naturae. This work represented a considerable
updating of and expansion upon the 12th edition of the
Stjstema Saturae (Linnaeus, 1767). Gmelin not only pro-
\ ided additional bibliographic references for the species
described by Linnaeus, but he also described numerous
new species, using the same format as had Linnaeus.
Gmelin's magnum opus was published in three volumes,
comprising 10 parts altogether: Regntim Animalium (7
parts); Regnum VegetaMe (2 parts); and Regnum La-
pideuni (1 part). For comments on the structure and
content of Gmelin's work, see Dodge (1958:157-158).
Few systematists, however, have realized that Gme-
lin's work actually underwent two separate printings.
The initial German printing was by Georg. Emanuel.
Beer of Lipsiae [= Leipzig]; the later French printing
was by J. B. DelamoUiere (subsequently as Bernuset,
Delamolliere, Falque et Soc. ) of Lugduni [= Lyon]. Both,
of course, were written in Latin.
Hopkinson (1907) provided a valuable collation of the
Lipsiae printing, based on an analysis of various German
literature abstracts and review publications. He noted
that only the first part in the two multi-part volumes had
a date on the title page and that this date "did not apply
to the whole of the parts in that \olume " (Hopkinson,
1907:1036), since the later parts were published in fol-
lowing years. Hopkinson did not refer to the Lugduni
printing. Agassiz (1852:69) listed this work as "Leipz.
1788-1793,. . . Lugduni, 1789-1796." Engelmann (1846:
103) provided the same dates as had Agassiz, but erro-
neously gave the place of publication as "Lugd. Batav."
which is actually Leiden [= Lugduni Batavorum]. Sher-
born (1902:.\.\xv) gave the same dates, made the same
geographical transposition as had Engelmann, and spec-
ified that the second printing was a reprint of the first.
Hulth (1907:13) and Soulsby (1933:16, 50) provided a
partial collation ol these two printings; however, their
dates are based solely on those stated on the title page
for the first part of each of the two multi-part volumes.
Stafleu and Cowan (1976:956) briefly mentioned the two
printings, and noted for the Lugduni printing: type
reset, no copy seen . Frequently , librarx copies will have
the same date stamped on the binding of each part or
hand-written on the first page of each part; this is in-
correct. The important facts are that the Lugduni print-
ing was published after the Lipsiae printing, and was
printed from reset type w hich resulted in a number of
differences due to printer s error. Therefore, the date of
publication of each part of the Lugduni printing must
post-date that of the relevant part of the Lipsiae printing.
L'nfortunately , we have not been able to determine
the date of publication for the succeeding parts of the
Lugduni printing (only for the first part in the multi-
part volumes). However, we present this comparison here
in order to alert s\ stematists to the existence of these two
printings. Should the reader know of further bibliograph-
ical sources bearing on this matter, we would greatly
appreciate hearing from them.
Herein we present as complete a collation as possible
of the two printings of Gmelin's Systema Saturae. based
on Hopkinson (1907) and on our examination of complete
sets of this work in the libraries of the Museum of Com-
parative Zoology , the British Museum (Natural History),
and the Museum National d Histoire Naturelle, Paris.
Gmelin, jo. Prid. [Joliann P^iedrich]. 1788-1793. Ca-
roli a Linne, Systema Naturae per Regna Tria Naturae,
Secundum Classes, Ordines, Genera, Species, cum cha-
ractcribus, diftcrentiis, synonymis, locis. Editio Decima
Tertia, Aucta, Reformata. Georg. Emanuel. Beer, Lip-
siae; [Second printing, 1789-1796], J. B. Delamolliere,
Lugduni.
Tomus 1, REGNUM ANIMALIUM
1(1) Mammalia to Aves Picae, pp. xii -I- 1-500
Lipsiae, 1788; Lugduni, 1789.
1 (2) Aves Anseres — A\es Passeres. pp. 501-1032
Lipsiae, 1789; Lugduni (post 1789?).
I (3) Amphibia — Pisces, pp. 1033-1516
Lipsiae, 1789; Lugduni (post 1789?).
A. R. Kabat and R. E. Petit 1988
Page 165
I (4) Insecta General, Coieoptera — Hemiptera, pp. 1517-
2224
Lipsiae, 1790; Lugduni (post 1790?).
I (5) Insecta Lepidoptera — Aptera, pp. 2225-3020
Lipsiae, 1790; Lugduni (post 1790?).
1(6) Vermes, pp. 3021-3910
Lipsiae, 1791; Lugduni (post 1791?).
1(7) Inde.x 1-3, pp. 3911-4120
Lipsiae, 1792; Lugduni (post 1792?),
Tomus II, REGNUM VEGETABILE
II (1) Monandria to Polyandria, pp. .\1 + 1-884
Lipsiae, 1791; Lugduni, 1796.
II (,2) Didynamia to Cryptogamia, pp. 885-1662
Lipsiae, 1792; Lugduni (1796 or later?).
Tomus 111, REGNUM LAPIDEUM
111, pp. 1-476, plates 1-3 (of crystals)
Lipsiae, 1793; Lugduni. 1796.
It has been our experience that molluscan taxonomists
who have used Gmelin's work have usually referred sole-
ly to the Lipsiae printing. Among others, these include
Dodge (1958:157-158); and Kohn (1966) who provided
biographical information on Gmeiin and a useful critique
of his work. .\n example of the taxonomic problems that
can arise with respect to the two printings of Gmelin's
\\ ork is that of certain species of Cancellariidae (Gastrop-
oda) reviewed by Petit (1984). For Buccinum piscato-
rium Gmeiin. 1791 (p. 3496), the Lipsiae printing gave
two references: "List. Conch, t. 1024 f. 89"' and "Martin.
Conch. 4. t. 124 f. 1151, 1152". Based on this, Petit chose
the figures from the latter reference (i.e., Chemnitz, J.
H., 1780, Neues Systetnatisches ConchyHen-Cabinet ,
\"ol. 4, Nurnberg) to be representative of this species.
Subsequently, A. Verhecken [in litt.) inquired as to how
the Chemnitz illustration could be used in this manner
since it was not cited by Gmeiin. In later correspondence,
N'erhecken advised that he had found that he was re-
ferring to a cop)- of the Lugduni printing. In that printing
only one reference, the aforementioned "List. Conch, t.
1024 f. 89" (i.e.. Lister, M.. 1692/1770, Historiae sive
Synopsis Methodicae Conchyliorum, London), was giv-
en for the species in question; the Chemnitz reference
being omitted. ObviousK', the omission of the reference
to Chemnitz was a printer's error, as a comparison of the
two printings makes it evident that the type was reset
and errors were made.
Incidentally, the third volume of this work (the Reg-
num Lapideum) was placed on the Official Index of
Rejected and Invalid Works in Zoological Nomenclature
(along with the related sections of Linnaeus and of Tur-
ton), since the "generic names of fossil animals used in
these works actually corresponded to the classes of Recent
animals (ICZN, 1954, Opinion 296).
We conclude that (1) since the Lipsiae printing was
published first, it should serve as the basis for discussion
of Gmelin's species; (.2) the Lugduni printing should be
treated as a reprint (or reissue) in which errors occur,
and not as a "second edition "; (3) systematists who are
studying species described by Gmeiin should compare
the two printings in order to uncover any discrepancies
with respect to species descriptions or references.
ACKNOWLEDGEMENTS
Mr. Andre Verhecken, Mortsel, Belgium, first brought
the discrepancy in the Gmeiin citations to our attention,
and then followed up with information on the existence
of the two printings. His interest and assistance is greatly
appreciated. We thank the librarians of the Museum of
Comparative Zoology, the British Museum (Natural His-
tory), and the Museum National d'Histoire Natureile,
who allowed us to examine the sets of Gmelin's work in
their collections. A number of other hbraries were checked
for this work; most had only partial sets of one or the
other printing which did not yield additional useful in-
formation. Prof. Kenneth J. Boss provided a helpful re-
view of this manuscript.
LITERATURE CITED
Agassiz, L. 1852. Bibliographia Zoologiae et Geologiae, Vol.
3 (of 4). Ray Society, London, 657 p.
Dodge, H. 1958. .\ historical review of the mollusks of Lin-
naeus. Part 6. The genus Trochus of the class Gastropoda.
Bulletin of the American Museum of Natural History
116(2):153-224.
Engelmann, W. 1846. Bibliotheca Historico-Naturalis. Ver-
zeichniss der Biicher iiber Naturgeschichte . . . Erster Band.
Verlag von Wiihelm Engelmann, Leipzig, x -I- 786 p.
Hopkinson, J. 1907. Dates of publication of the separate parts
of Gmelin's edition (13th) of the 'Systema Naturae' of
Linnaeus. Proceedings of the Zoological Society of London
1907(49):1035-1037.
Hulth, J. M. 1907. Bibliographia Linnaeana: materiaux pour
servir a une Bibliographie Linneenne. Kungliga Veten-
skaps Societeten I Upsala, 170 p. -H 11 pis.
International Commission on Zoological Nomenclature, 1954.
Opinion 296. Suppression, under the Plenary Powers, for
nomenclatorial purposes, of volume 3 (Regnum Lapi-
deum) of the Twelfth Edition of the Systema Naturae of
Linnaeus published in 1768 and of the corresponding vol-
ume in the Houttuyn (1785), Gmeiin (1793) and Turton
U806) editions of the above work. Opinions and Decla-
rations rendered by the International Commission on Zoo-
logical Nomenclature 8(13):167-178.
Kohn, A. J. 1966. Type specimens and identity of the de-
scribed species of Conus III. The species described by
Gmeiin and Blumenbach in 1791. The Journal of the Lin-
nean Societv of London, Zoology 46(308):73-102, pis.
1-3.
Linnaeus, C. 1766-67. Systema Naturae, per Regna Tria Na-
turae, . . . Editio Duodecima Reformata. Direct. Laur. Sal-
vii, Holmiae. Regnum Animale, 1328 -t- 36 p.
Petit, R. E. 1984. Some early names in Cancellariidae. Amer-
ican Malacological Bulletin 2:57-61.
Sherborn, C. D. 1902. Index Animalium sive index nominum
quae ab .\.D. MDCCL\'III generibus et speciebus ani-
malium imposita sunt . . Sectio Prima. L niversity Press,
Cambridge, Ix + 1195 p.
Page 166 THE NAUTILUS, Vol. 102, No. 4
Soulsby, B. H. 1933. A catalogue of the works of Linnaeus Stafleu, F. A. and R. S. Cowan. 1976 [in 1976-86]. Taxonomic
(and publications more immediately relating thereto) pre- literature: a selective guide to botanical publications and
served in the libraries of the British Museum (Bloomsbury) collections with dates, commentaries, and types. Second
and the British Museum (Natural History) (South Ken- Edition. Regnum X'egetabile; Bohn. Scheltema & Holke-
sington). British Museum (Natural History), London, 2nd ma, Utrecht, \'ol. 1 (of 6):xl + 1136 p.
ed., p. xii -I- 1-312; 7 pis.
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Reprints may be ordered through the editor.
Manuscripts, corrected proofs and correspondence re-
garding editorial matters should be sent to; Dr. M.G.
Harasewych, Editor, Division of Mollusks, NHB stop 118,
National Museum of Natural History, Smithsonian In-
stitution, Washington, DC 20560, USA.
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