MBL/WHOI
THE NAUTILUS
Volume 103
1989-1990
AUTHOR INDEX
Al'FFENBERC, K.
Bond, \' N.
Bol't.HET, P.
BURNHAM, B n
Castacna, M
Deaton, L. E,
Dillon, R. T., Jr.
Kmebson, W. K.
Fairbanks, H L
Fell. P, E
Fuller, S. C.
Harasewych, M. G.
Hastie, L. C.
Hartfield, p. D.
Hu, Y. P.
Itano, D.
Jirka. K J
Kabat, .\ R
Karuson, R H
Kempler, K. D.
Leal, J. H.
LuTZ, R. A.
Manzi, J J
143
McMahon, R. 1
99
Miller, A. C.
1
Mitchell, M
109
Nevf^, R. J
24
Payne, B. S.
42
Petit, R. E.
73
Petuch, E J
131
Pip, E.
20
Portell, R. W
109
Prezant, R. S.
24
{^UINN, J F.. Jb
70, 83
Render. H. A.
99
Reid. D. G.
78
Rex, M. a.
24
Sage, W.
99
Saunders. W. I
136
SCAVIA, E.
113
Sullivan, J A
85
Tan Tiu, A.
42
Vale, F. K,
1
Vermeij, G, J.
24
X'OKES. E. n.
73
Way. C. M
78
78. 96
40
136
78,96
83
92
140
143
36
13
117
43
105
131
99
40
85
36
105
89
124
96
NEW TAXA PROPOSED IN VOLUME 103 (1989-1990)
Peasiella lutulenta Reid, 1989. new species (Littorinidae)
Malea peiiti Petuch, 1989, new species (Tonnidae)
Malea springi Petuch, 1989, new species (Tonnidae)
Chicorem (Stratus) carohjnae E. H. V'okes, 1990, new species (Muricidae)
ChicureuH (Stratus) coltrorum E. H. V'okes, 1990, new species (Muricidae)
Odontocymhiola simulatrtx Leal and Bouchet, 1989, new species (Volutidae)
Tractolira tenehrosa Leal and Bouchet, 1989, new species (Volutidae)
Sanonu'lon Leal and Bouchet, 1989, new genus (Volutidae)
Nanomelon vipertnus Leal and Bouchet, 1989, new species (Volutidae)
Cohizca juliae Harasewych, 1989. new species (Turbinellidae)
Pleiopt\ginatidae Quiiui, 1989, new family (Neogastropoda)
Praticolella prtsca Auffenberg and Portell, 1990, new species (Polygyridae)
59
94
94
126
127
2
6
70
13
143
THE NAUTILUS
Volume 103, Number 1
June 28, 1989
ISSN 0028-1344
A quarterly devoted
to malacology.
Marine Biological Laboratory f
LIBRARY \
JUL 1 0 1989
Woods Hole, Mass.
EDITOR-IN-CHIEF
Dr. M. G. Harasewych
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
ASSOCIATE EDITOR
Dr. R. Tucker Abbott
American Malacologists, Inc.
P.O. Bo.x 2255
Melbourne, PL 32902
CONSULTING EDITORS
Dr. RiJdiger Bieler
Department of Malacoiog)
Delaware Museum of
Natural History
P.O. Box 3937'
Wilmington, DE 19807
Dr. Robert T. Dillon, Jr.
Department of Biology
College of Charleston
Charleston, SC 29424
Dr. William K. Emerson
Department of Living Invertebrates
The American Museum of Natural
History
New York, NY 10024
Mr. Samuel L. H. Fuller
1053 Mapleton Avenue
Suffield, CT 06078
Dr. Robert Hershler
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
Dr. Richard S. Houbrick
Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560
Mr. Richard I. Johnson
Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Aurele La Rocque
Department of Geology
The Ohio State University
Columbus, OH 43210
Dr. James H. McLean
Department of Malacology
Los .Angeles County Museum of
Natural History
900 E.xposition Boulevard
Los Angeles, CA 90007
Dr. Arthur S. Merrill
% Department of Mollusks
Museum of Comparative Zoolog\
Harvard University
Cambridge, MA 02138
Ms. Paula M. Mikkelsen
Harbor Branch Oceanographic
Institution, Inc.
Ft. Pierce, FL 33450
Dr. Donald R. Moore
Division of Marine Geology
and Geophysics
Rosenstiel School of Marine and
Atmospheric Science
University of Miami
4600 Rickenbacker Causeway
Miami, FL 33149
Mr. Richard E. Petit
P.O Box 30
North Myrtle Beach, SC 29582
Dr. Edward J. Petuch
Department of Geology
Florida Atlantic L'niversit\
Boca Raton, FL 33431
Dr. G. Alan Solem
Department of Invertebrates
Field Museum of Natural History
Chicago, IL 60605
Dr. David H. Stansbery
Museum of Zoology
The Ohio State University
Columbus, OH 43210
Dr. Ruth D. Turner
Department of Mollusks
Museum of Comparative Zoology
Harvard University
Cambridge, MA 02138
Dr. Geerat J. Vermeij
Department of Geology
University of California at Davis
Davis, CA 95616
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TH E€7NAUTI LUS
CONTENTS
Volume 103, Number 1
June 28, 1989
ISSN 0028-1344
Jose H. Leal
Philippe Bouehet
New deep-water Volutidae from off southeastern Brazil
(Mollusca: Gastropoda) 1
Pleioptygmatidae, a new family of mitriform gastropods
(Prosobranchia: Neogastropoda) 13
The reproductive anatomy and taxonomic status of
Philomycus venustus Hubricht, 1953 and Philomycus
bisdosus Branson, 1968 (Pulmonata; Philomycidae) 20
Shell and pallet morphology in early developmental stages
of Teredo navalis Linne (Bivalvia; Teredinidae) 24
Shell tubules in Corbicula fluminea (Bivalvia:
Heterodonta): Functional morphology and microstructure 36
Reoccurrence of Cyclonaias tuberculata in the Huron
River, Michigan 40
Occurrence of the ribbed mussel, Geukensia demissa, on
the book gills of a horseshoe crab, Limulus polyphemus 42
James F. Quinn. Jr.
H. Lee Fairbanks
S. Cynthia Fuller
Ya-Ping Hu
Richard A. Lutz
Michael Castagna
Antonieto Tan Tiu
Robert S. Prezant
Ellen Scavia
Mark Mitchell
Lewis E. Deaton
Karen D. Kempler
Marme Biological Laboratory
UBRARY
JUL 10 1989
V^oods Hole, Mass.
THE NAUTILUS 103(1):1-12, 1989
Page 1
New Deep- Water Volutidae from off Southeastern Brazil
(Mollusca: Gastropoda)
Jose H. Leal
Biolog) and Living Resources
Rosenstiel St-hool of Marine and
Atmospheric Science
4600 Rickenbacker Causeway
Miami, FL 33149, USA
Philippe Bouehet
Museum National d'Histoire Naturelle
55 rue Buffon
Paris 75005, France
ABSTRACT
One new genus and three new deep-u ater species of the N'ohiti-
dae are described from the southeastern Brazihan coast. Ac-
cording to accepted supraspeciSc classification of the \'ohitidae,
anatomical characters, such as accessory salivary glands tightly
wound around the primar\ salivary glands, allocate Odonto-
cymbiola simulatrix new species, from the upper bath\al zone,
to the subfamiK Odontocymbiolinae. The rachidian teeth have
characteristic "fang-like" cusps present in the genus Odonto-
cymbiola. Tractolira tenebrosa new species, an upper abvssal
species of which only dead shells are known, is conchologicallv
related to the Antarctic species T. germonae. differing chiefly
in protoconch morpholog)'. The famiK Zidoninae is repre-
sented b\ the bath\al 'Sanomelon viperinus new genus, new-
species, which displays the loose association between the two
types of salivary glands and opened sperm groove (running to
the tip of the penis), diagnostic of the Zidoninae. However, its
small, elongated, heavy shell, with strongly clathrate sculpture,
the broad rachidian teeth with elongated, straight-edged cusps
interlocking with the subsequent teeth, and a large rectal gland
that branches anteriorly from a posterior duct are among the
characters that differ considerabK from those of other genera
in the subfamiK Zidoninae. .\n examination of the taxonomic
literature on the X'olutidae suggests the convergent or primitive
nature of characters traditionally considered as diagnostic of
the subfamilies Odontocvmbiolinae and Zidoninae
INTRODUCTION
Deep-water operations carried on by the research ship
Marion-Dufresne in May, 1987, off the southeastern coast
of Brazil, yielded 5 vohitid species of the subfamilies
Odontocymbiolinae and Zidoninae. Dead shells and liv-
ing specimens were dredged or trawled in depths ranging
from 200 to 3,270 m, in a transect conducted perpen-
dicularly to the Brazilian coast, between 21° and 24°S.
This material includes deep-water species already known
to occur in this sector of the Atlantic, and three unde-
scribed volutid species belonging to two subfamilies. Ex-
cept for two tropical, shallow-water species of the
subfamilies Volutinae {Valuta ehraea Linnaeus, 1758)
and Lyriinae [[?]Enaeta guildingii (Sowerby, 1844)], all
known volutids in the southwestern Atlantic belong to
subfamilies Odontocymbiolinae and Zidoninae, includ-
ing species living in the bathyal zone (Rios, 1985).
The subfamily Odontocymbiolinae was described to
encompass volutids with "three prong or fang-like den-
ticles" on the rachidian teeth and accessory salivary glands
w rapped tightly around the salivary glands (Clench &
Turner, 1964). Subsequent workers have provided ad-
ditional information on radular and anatomical char-
acters of other genera and species in this subfamily (Cas-
tellanos, 1970; Weaver & Dupont, 1970; Stuardo &
Villarroel, 1974; Klappenbach, 1979; Harasewych, 1987),
Based on the Recent distribution and almost complete
restriction of the Odontocymbiolinae to the Southern
Hemisphere, Harasewych (1987) has suggested that the
subfamily evolved in the Weddelian Province after the
separation of New Zealand at the end of the Early Pa-
leocene. Three species from three genera of the Odon-
tocymbiolinae were collected during the MD-55 Cruise:
Minicymbiola corderoi (Carcelles, 1953), Odontocym-
biola simulatrix new species, a species probably closely
related to two temperate South Atlantic species [O. mag-
ellanica (Gmelin, 1791) and O. subnodosa (Leach, 1814)].
The third species collected during our survey, Tractolira
tenebrosa new species, is an abyssal species probably
related to T. germonae Harasewych, 1987 from the South
Sandwich Islands, Scotia Sea, and from which it differs
b\- conchological characters (only dead shells are known
of the new species), particularly protoconch morphology.
The Zidoninae includes 9 living genera, three of which
occur in the southwestern Atlantic Ocean. Clench and
Turner (1964) characterized the Zidoninae as having uni-
serial radulae, rachidian teeth with three pointed cusps
in one plane, two equal lobes at the base of the siphon,
and tubular salivary glands loosely wound around mod-
erateK compacted salivarv glands (see also Ponder, 1970;
Page 2
THE NAUTILUS, Vol. 103, \o. 1
Novelii & N'ovelli, 1982). The subfamily is represented
in the deep-water material of the MD-55 cruise by Ade-
lomelon riosi Clench & Turner, 1964, and by a new
species that differs considerabK' in conchological, ana-
tomical and radular characters from other genera in the
subfamily. Consequently, a new genus, Nanomelon new-
genus, is here erected to accommodate it.
MATERIAL AND METHODS
Unless otherwise indicated, all material mentioned was
obtained during the MD-55 cruise of the Marion-Dufres-
ne b\ P. Bouchet, J. H. Leal, and B. Meti\ier. in May.
1987. Shells were fractured using a table vise, cleaned
in full strength commercial bleach (Clorox) for 30 sec,
rinsed in distilled water, and sonicated for the observa-
tion of shell ultrastructure. The following abbreviations
are used: MNHN, Museum National d'Histoire Natu-
relle, Paris, France; MNRJ, Museu Nacional, Rio de Ja-
neiro, Brazil; MORG, Museu Oceanografico de Rio
Grande, Brazil.
DESCRIPTION
Family Volutidae Rafinesque, 1815
SubfamiK Odonlocymbiolinae Clench & Turner, 1964
Genus Odontocymbiola Clench & Turner, 1964
Odontocymbiola simulatrix new species
(figures 1. 2, 5, 6, 14, 19-21, 26, 27)
Shell (figures 1. 2, 5, 6, 14): Fusiform, reaching 111
mm in length and 41 mm in width. Spire elevated, spire
angle about 43° Earlier teleoconch whorls moderately
solid, last whorl thinner and delicate. Shell surface rough
and opaque. Periostracum grayish to dark-brown. Shell
color cream to pale-orange with straight-edged spiral
bands of interrupted, mahogany-brown spots. Three spi-
ral bands in last whorl, one each at base, mid-whorl, and
abapical to suture. Preceding whorls with only two bands,
mid-w horl band partialK occluded b\ subsequent whorl.
Transition protoconch/ teleoconch (figure 14) poorly de-
fined. Protoconch (figure 14) smooth, with very fine mi-
croscopic spiral striae, with about 2.5 cream-colored
whorls. Calcarella small, of slightly darker color. Teleo-
conch with about 7 whorls, all but last shouldered; con-
cave sutural slope present in shouldered whorls. Suture
moderately impressed. A.xial ribs triangular in cross-sec-
tion, stronger and fewer on earlier whorls (about 15 on
the first, 23 on the third), changing into a pattern ot ill-
defined, numerous a.xial wrinkles (about 70) on last w horl.
Spiral sculpture of 140-150 fine raised lines on last \\ horl.
30-40 on preceding whorls. Spiral lines of same width
over teleoconch surface. Aperture elliptical (length width
about 3). Outer lip simple, thin, thinner in larger shells.
Parietal region glazed. Glazed surface in larger individ-
uals a thin spiral band adapical to suture of last whorl,
as result of shell grow th. Columella arched, with siphonal
fold and 2-3 columellar plaits. Anterior canal wide,
weakly defined. Inner shell surface smooth (see table 1
for shell measurements and whorl counts).
External morphology (figure 26): Living animal with
overall light-orange to salmon coloration, foot sole cream-
white. Coloration faded in preser\ed specimens. Head
broad, flattened, with large central lobe and two large,
semicircular lateral lobes posterior to tentacles. Tentacles
tapered, pointed distally. Eyes present. Foot wide (length/
width = 1.3, preser\ed holot\pe), pointed posteriorly,
bilobed anteriorly. .Aperture of mucous gland ver\ large,
extending slightK' around lateral edges of foot. Mantle
edge moderately thick. Siphon (si) large, muscular. Two
siphonal appendages (Isa, rsa) of equal length, each about
half length of siphon (preserved holot\pe). Siphonal ap-
pendages slightly flattened at distal extremity. Ctenidi-
um (ct) leaf-shaped. Osphradium (os) bipectinate with
pointed extremities (length/width = 3), % as long as
ctenidium.
Anterior part of the alimentary system (figure 27):
Proboscis pleurembolic. Salivary glands (sg) opaque-
white, moderately compacted. Ducts of salivary glands
(dg) long, opening into posterior part of buccal mass
(bm). Accessory salivar\ glands (as) \ellowish-white, rib-
bon-like, very narrow and long, tightK wound around
salivary glands. Ducts of accessory salivar\' glands open-
Table 1 . Odontocymbiola simulatrix new species. Linear shell
measurements (mm) and whorl counts for the holotype (HOL)
and paratypes 1-3 (PA 1-3). For localities see text.
( ^iiaraclt-r
HOL
P,\ 1
PA 1
PA3
Total leiietll
111.3
70.2
.55.0
69.8
Shell widtti
43.6
29.1
23.2
28.1
Leiintli last wliorl
84.9
54.8
43.8
56.7
.\piTlure length
62.8
44.9
34.1
44.3
.\perture width
22.2
14.0
11.0
15.2
Prolocoiicli diameter
5.0
4 1
4()
4.0
Teleoconch whorls
7.0
5.0
3.5
0.0
Protoconch whorls
2.5
2.0
2.5
2.5
Length/width
2.55
2.41
2.39
2.48
.Aperture length/length
0.56
0 64
0.62
0 63
Aperture length ap.
width
2.83
3.21
3.09
2.91
Figures 1, 2. Odontocymbiola simulatrix new species. Holotype, 111.3 mm length, 43.6 mm width Figures ;{, 4. .Xdelomelon
riosi Clench & Turner, 1964. 134 mm length, 54 mm width, off Espirito Santo State, 19°36'S, 38°53'VV, 640 m depth Figures 5,
6. Odontoci/ndnola simulatrix new species. Parat\pe 3, 69.8 mm length. 28.1 mm width. Scale bars = 20 mm Figures 7, 8.
Minicijmhiola corderoi (Cartelles, 1953) 24.5 mm length, 11.1 mm width, off Rio de Janeiro State. 23''36'S, 42°02'\\'. 200-217 m
depth Figures 9, 10. S'anomcton vipcrinus new genus, new species. Holot\pe, 44.2 nmi length, 15.3 mm width Figures II-
i:{. Tractolira tenebrosa new species 1 1. 12. Holotype, 38.3 mm length, 13.8 mm width. 13. Parat\pe 2, 41.9 nun length, 15.3
mtii width. Scale bars = 10 mm.
J. H. Leal and P. Bouchet, 1989
Pages
Page 4
THE NAUTILUS, \ol. 103, No. 1
16
18
J. H. Leal and P. Bouchet, 1989
Page 5
iiig into anteriormost region of buccal mass. Gland of
Leiblein (gl) long, convoluted, opening into oesophagus
just posteriorly to circumoesophageal nerve ring and valve
of Leiblein (vl). Stomach (st) with anterior tubular region
(ts).
Radula (figures 19-21): Radular ribbon (figure 19)
uniseriai, relati\el\ short {length = 19.6 mm, length
width = 24.5, holotype), with about 45 tricuspid rachidi-
an teeth (figure 20), each 0.8 mm wide. Basal plate flat-
tened, chevron-shaped, excavated posteriorly in surface
in contact with ribbon. Central cusp at least 1.5 times
longer than and 2 times w ider than lateral cusps. Central
and lateral cusps deepK' curved, "fang-like". Central
cusp rising from anterior edge of basal plate, pointing
posteriorly. Central cusp with very sharp, thin lateral
edges and with two, deep lateral longitudinal grooves,
giving origin to long, dorsal rib narrower than central
cusp. Lateral cusps deeply grooved ventrally (figure 21).
Holotype: MORG 25467, 111.3 mm length, 43.6 mm
width, MD-55 station CP-11 (Beam trawl), ofl^ Rio de
Janeiro State, Brazil (collected alive).
Type locality: Continental slope off the coast of Rio de
Janeiro State, Brazil, 21°35'S, 40°06'VV, at 248 m depth,
muddy bottom,
Paratypes: Paratype 1, MNHN, 70 mm length, 29 mm
width; Paratvpe 2, MNHN, 55 mm length, 23 mm width,
MD-55 station CB-104 (Blake trawl), 23°42'S, 42°07'W,
430-450 m depth, muddy bottom; Paratype 3, MORG
15910, R/ V Mestre Jeronimo, off Ilha de Santa Catarina,
28°03'S, 48°11"W, 113 m depth, 07/1971.
Other material examined: One juvenile shell + 1 bro-
ken shell, station CB-104 (Blake trawl) (same localities
as Paratypes 1 and 2).
Remarks: One dead shell of O. simulatrix (Paratype 3,
MORG 15910) had been collected off southern Brazil in
1971, and had been recognized as an unidentified volutid.
Odontocymbiola simulatrix differs from the other five
known species of the subfamily in size, shell sculpture,
coloration, and radular structure. It can be set apart from
Minicijmbiola corderoi (Carcelles, 1953) by its color pat-
tern of spirally arranged brown spots, larger size {cor-
deroi reaches 28 mm, according to Rios, 1985), less prom-
inent sculpture and larger, slender shell with higher spire
(compare figures 1, 2, 5, and 6 with 7 and 8). The con-
generic Odontocymbiola americana (Reeve, 1856), is
smaller, has a shorter spire, a smoother shell surface,
smaller number of axial ribs, more pronounced shoulders,
thicker outer lip, lacks a calcarella at the protoconch
ape.x, and has a more elaborate, delicate color pattern.
Odontocymbiola simulatrix differs from O. subnodosa
(Leach, 1814), O. magellanica (Gmelin, 1791), and O.
pescalia Clench & Turner, 1964, by its more elongated
shape, narrower aperture, presence of a well-defined spi-
ral color pattern, smaller parietal region, stronger axial
and spiral sculpture, and w eaker columellar folds. Odon-
tocymbiola simulatrix has a radula characteristic of the
genus, with "fang-like", deeply curved cusps in the ra-
chidian. However, the rachidian in the new species (fig-
ures 20, 21) lacks the extremely elongated central cusp
and the blunt extremities of the lateral cusps as in O.
americana. The lateral edges of the central and lateral
cusps are not as sinuated as those of O. pescalia, and the
basal plate lacks the rounded lateral expansions present
in O. magellanica (see Clench & Turner, 1964; Weaver
& Dupont, 1970; Kaiser, 1977 for descriptions and illus-
trations of the mentioned species).
Although the soft parts of only one female individual
of O. simulatrix were examined, the few characters de-
rived from external anatomy (figure 26), alimentary sys-
tem (figure 27), and radula (figures 19-21) are sufficient
to allow subfamilial and generic placement. The long,
equal siphonal appendages, very elongated accessory sal-
ivary glands, tightly wound around rather compacted
salivary glands, a stomach with an elongated anterior
section, and the curved, elongated, and basally broadly
separated, "prong or fang-like" cusps of the radular teeth
are usually accepted as derived characters defining
Odontocymbiolinae (see Clench & Turner, 1964; Stuardo
& V'illarroel, 1974; Harasewych, 1987). The extremely
narrow ("fang-like"), elongated, and intricately sculp-
tured cusps "which extend at a right angle from the basal
plate and then curve downwards" (Clench & Turner,
1964) are here conditionally considered as the character
set that defines the genus Odontocymbiola, as opposed
to the "prong-like" cusps present in the genus Miomelon
and Tractolira (Harasewych, 1987). Klappenbach (1979),
using only shell dimensions and sculpture, has erected
the monotypic genus Minicymbiola for Marginella cor-
deroi Carcelles, 1953. The radular morphology illustrat-
ed in the original description of Minicymbiola indicates
that M. corderoi complies with the definition of the genus
Odontocymbiola by Clench and Turner (1964). Not-
withstanding, we conditionally consider Minicymbiola
to be a valid genus, though only a thorough study of
other anatomical characters in the species could corrob-
orate the validity of the extremely divergent shell char-
acters (see figures 7, 8) in the foundation of the genus.
As observed in the SEM micrographs of the rachidian in
O. simulatrix, and in camera lucida drawings for the
Figure 14. Odontocymbiola simulatrix new species. Protoconch. Figures 15-18. Xanomelon viperinus new species. 15.
Protoconch. 16-18. Shell ultrastructure 16. Collabral cross-section of last half of last whorl: a. external crossed-lamellar layer; b,
middle crossed-lamellar la>er; c. third crossed-lamellar la\er; d. internal prismatic layer 17. Cross-section of third teleoconch whorl
at 4.5° to shell axis; change of orientation of first order lamellae (middle crossed-lamellar layer, b) conforms to whorl curvature.
Dashed line indicates that external crossed-lamellar laver (a) is absent. 18. Cross-section of third teleoconch whorl, approximately
perpendicular to shell axis; notice perpendicular relationship between first order lamellae of middle (b) and third (c) crossed-lamellar
layers, and very thin internal prismatic layer (d). Scale bars, protoconchs = 1 mm. ultrastructure = 0.50 mm.
Page 6
THE NAUTILUS, Vol. 103, No. 1
Figures 19-21. Odontocymhiola sintulatrix new species. Radular teeth. 19. Segment ot radular ribbon showing relati\e position
of rachidian teeth in non-protracted condition. 20. Dorsal view of rachidian; notice lateral grooves and longitudinal rib in central
cusp. 21. Ventral view of rachidian tooth showing attachment surface; arrow indicates groove in lateral cusp. Figure 22. Ade-
lomelon riosi. Radular teeth. Figures 23-25. \anornelon viperinus new genus, new species. Radular teeth. 23. Radular ribbon,
showing relative positions of rachidian teeth in protracted position and "nesting" of the cusp tips in succeeding tooth 24-. Dorsal
view of rachidian tooth. 2.5. Neutral view of rachidian tooth show ing attachment surface. Scale bars = 0..50 mm.
remaining species (Clench & Turner, 1964; Castellanos,
1970; Weaver & Dupont, 1970), all representatives of
Odontocymhiola exhibit very thin lateral edges and a
longitudinal, dorsal rib in the central cusp of the ra-
chidian. The combination of very thin lateral edges and
a longitudinal, reinforcing rib probably results in larger
cutting effect without weakening of the cusp.
Genus Tractolira Dall, 1896
Tractolira tenebrosa new species
(figures 11-13)
Shell (figures 11-13): Elongate, fusiform, reaching
about 51 mm length and 17 mm width. Spire elevated,
spire angle about 25°. Whole shell surface chalky, dirty
white. Shell thin, fragile. Periostracum unknown. Pro-
toconch dome-shaped, with about 5 mm maximum di-
ameter, 2.5 whorls. Teleoconcli with up to 4.0 convex
w horls. First three teleoconch w horls with about 20 axial
ribs. Remainder of teleoconch with axial sculpture of
fine growth lines. Spiral sculpture of fine cords, about
20-25 in third teleoconch w liorl, 80-100 in the last w horl.
Suture excavated. .-Vperture elongate-elliptical (length/
width = 3.5). Outer lip with narrow varix. Parietal region
well delimited, with indication of glazed surface (even
in badlv eroded type-material). Columella arched, with
2-3 columellar plaits (see table 2 for shell measurements
ami whorl counts).
Holotype: MORG 25468, 38.3 mm length, 13.8 mm
width, MD-55 station CP-01 (Beam trawl), off Rio de
Janeiro State, Brazil.
J. H. Leal and P. Bouchet, 1989
Page 7
Figures 26, 27. Odontocymbiola simulatrix new species. Holotype. 26. Diagram showing the relative positions of some of the
organs in the mantle cavity. 27. Anterior part of alimentary system. Scale bars = 20 mm.
as, accessory salivary gland; bm, buccal mass; ct, ctenidium; dg, duct of salivary gland; gl, gland of Leiblein; Isa, left siphonal
appendage; ma, mantle; mo, mouth; os, osphradium; pe, posterior oesophagus; rsa, right siphonal appendage; sg, salivary gland; si,
siphon; st, stomach; ts, tubular extension of stomach; vl, valve of Leiblein.
Type locality: Lower continental slope off the coast of
Rio de Janeiro State, Brazil, 23°04'S, 40°20'W, at 2,370-
2,380 m depth, muddy bottom.
Paratypes: Paratype 1, MNHN, 4L9 mm length, 15.3
mm width, MD-55 station CP-01 (type locality); Para-
type 2, MNHN, 51.3 mm length, broken aperture, MD-
55 station CP-17, 2r08'S, 38°25'W, 3,250-3,270 m depth,
muddy bottom.
Remarks: Tractolira tenebrosa is known only from dead
shells collected at the upper abyssal zone (between 2,370
and 3,270 m depth). Allocation to subfamilial and generic
level is here based solely on shell morphology and pre-
vious records of the other two congeneric species, T.
sparta Dall, 1896 (Gulf of Panama) and T. germonae
Harasewych, 1987 (South Sandwich Islands, Scotia Sea),
from equivalent depths and bottom conditions. Tracto-
lira tenebrosa differs conchologically from T. sparta and
T. germonae by having a dome-shaped protoconch in-
stead of the pointed, calcarella-bearing protoconch of
the latter two species. Also, the first teleoconch whorls
are more crowded, with smaller incremental angles in
T. tenebrosa. Association of these two conditions pro-
duces a blunter, less conical profile in the early teleoconch
whorls. The new species is shorter than T. sparta, has a
proportionally larger aperture, and weaker a.xial ribs re-
stricted to the two first teleoconch whorls. Axial ribs are
absent in T. germonae; spiral threads are more numerous
in T. tenebrosa (80-100, last whorl) than in T. germonae
(60-80, last whorl), where they are also broader. The
new species lacks the flared outer lip and the anteriorly
twisted columella and anterior canal, observed in T. ger-
monae.
Subfamily Zidoninae H. & A. Adams, 1853
Genus Nanomelon new genus
Shell: Small [37.6 ± 6.5 mm (n = 11) in the type
species]. Spire angle small. Shell surface opaque. Proto-
conch with about 2.5 whorls. Spiral sculpture of about
same intensity as axial sculpture, giving clathrate aspect
to shell. Spiral sculpture at sutural ramp of cordlets finer
and more closely set than spiral cords on rest of whorl.
Pages
THE NAUTILUS, Vol. 103, No. 1
Table 2. Traclolira tenebrosa new species. Linear shell mea-
surements (mm) and whorl counts for the holotype (HOD and
paratypes 1-2 (P.A 1-2). For localities see text.
( Miaracter
liOl.
l'\ 1
P.\ 2
Total length
.38.3
41,9
51,3
Shell width
13.8
15.3
—
Length last u horl
28.4
32.8
37.2
Aperture length
21.0
24.8
27.8
Aperture width
6.3
6.6
8.4
Protoconch diameter
4.5
5.3
5.0
Teleoconch whorls
3.50
3.00
4.00
Protoconch whorls
2.25
2.75
2.50
Length width
2.78
2.74
—
.Aperture length length
0.55
0.59
0 54
Aperture length ap width
3 33
3 7fi
3 31
A.\ial sculpture absent along the base. Parietal region
well defined. Columellar plicae poorly defined, Radula
\\ ith wide, tricuspid rachidian teeth. Cusps elongated,
e.xtremities of cusps nested in small, deep depressions of
subsequent teeth. Outer margins of lateral cusps straight.
Type species: Nanomelon viperinus new species.
Nanomelon viperinus new species
(figures 9, 10, 15-18, 23-25, 28-33)
Shell (figures 9, 10, 1 5): Fusiform (length/ width about
2.7), imperforate, reaching 45 mm length, 16 mm width.
Spire angle about 30°. Shell surface opaque, dirty-white
to light-gray. Periostracum very thin, grayish-brown. Shell
sometimes eroded where periostracum absent. Transition
from protoconch to teleoconch poorly defined (figure 15).
Protoconch (figure 15) white, cylindrical, about 2.5 whorls,
first 1.5 whorls smooth, remaining whorls sometimes with
fine spiral striae. Last protoconch whorl with same di-
ameter as preceding whorl. Teleoconch with up to 4.5
convex whorls, wider at periphery. Suture impressed,
sutural ramp slightly concave. Spiral sculpture of about
4 wavy cordlets on sutural ramp and 5 larger spiral cords
on remainder of whorl. Interspaces between cords at least
three times larger than between narrower, adapical spiral
cordlets. Rase with about 12 spiral ribs, more crowded
abapically. Aperture elongate (length/ v\idth about 4),
Outer lip simple, thin. Interior of aperture glazed. Pa-
rietal region smooth, well defined, and sometimes delim-
ited by narrow glazed band. Columella arched, with
siphonal fold and 3-4 columellar plaits. Anterior canal
wide (see table 3 for shell measurements and whorl
counts).
Shell ultrastruclure (figures 16-18): Shell with three
crossed-lamellar ((JSLj and one internal prismatic layer.
External CSL (figures 16, a, 18) about 100 nm thick (in
collabral cross-section), observed only in last half of last
« horl, horizontal axis of first order lamellae perpendic-
ular to collabral section. Middle CSL (figures 16-18, b)
about 550 ^m thick, horizontal axis of first order lamellae
parallel to collabral section. Middle CSL comprises spiral
and axial shell ribs. Third CSL (figures 16-18, c) about
270 ^m thick, horizontal axis of first order lamellae per-
pendicular to collabral section. In middle and third CSL's,
second order lamellae in adjacent first order lamellae
form an angle of 80° with each other. Innermost pris-
matic layer (figures 16-18, d) simple, 20-50 ^m thick
External morphology (figures 28-31): Living animal
dull gra\ ish-w hite. Head broad and flattened, with two
semicircular lateral lobes (figure 28, lo) posterior to ten-
tacles. Tentacles (figures 28, 29, te) small and short. Eyes
present. Foot (figure 28, f) narrow (length/width = 2,
preserved animal), tapered posteriorly, .\perture of mu-
cous gland (figure 28, mg) situated in anterior, broad
extremity of foot. Mantle edge thin. Two siphonal ap-
pendages (figure 29, Isa, rsa) of equal length, each about
'/3 of length of siphon (figure 29, si); right siphonal ap-
pendage with tapered distal extremity, left with blunt,
flat distal extremity. Ctenidium (figure 29, ct) leaf-shaped,
filaments (figure 30) triangular and elongated laterally,
with lateral cilia (figure 30, Ic) distributed in well-de-
fined, elongated area at ventral half of filament. Os-
phradium (figures 29, os, 31) bipectinate, with rounded
extremities (length/ width = 3), % as long as ctenidium.
Hypobranchial gland consisting of delicate, whitish la-
mellae occupying large area at right side of the mantle
cavity. Secretion of hypobranchial gland dark-purple in
preserved animals.
Alimentary system (figure 32): Proboscis pleurembolic.
Salivary (sg) and accessory salivary glands (as) partially
cover circumoesophageal nerve ring (en) and valve of
Leiblein (vl). Salivary glands grayish-white, well com-
pacted. Ducts of salivary glands (dg) opening into an-
terior oesophagus, close to buccal mass (bm). Accessory
salivary glands white, loosely wound, situated anteriorly
to salivary glands. V'alve of Leiblein slightly anterior to
circumoesophageal ner\e ring. Gland of Leiblein (gl)
Figures 28-33. Nanomelon viperinus new genus, new species Male specimen, 28. Anterior part of animal, 29. Diagram showing
the relative positions of some organs of the mantle cavit\, 30. Ctenidial filament in frontal \iew 31. Pair of ospliradial lamellae
in frontal view, 32. Alimentary ssstem, 33. Anterior part of male reproductive svstem; arrow indicates prostate opened ventralK .
Scale bars = 5 mm, except for 26, 27, bar = 1 mm, and 29. bar = 2 mm
an, anus; as, accessory salivary gland; bm, buccal mass; en, circumoesophageal ner\ e ring; ct. ctenidium; dd, duct of digestive gland;
dg, duct of salivary gland; do, dorsal groove of prostate; dr, duct of rectal gland; e, e\c; f, foot; gl, gland of Leiblein; in, intestine;
lb, nerve of gland of Leiblein; Ic, lateral cilia of ctenidium; lo, lateral cephalic lobe; Ip. lateral glandular lobes of prostate; Isa, left
siphonal appendage; ma, mantle; mo. mouth; mw, mantle wall; pe, posterior oesophagus; pn. penis; pr, prostate; os. osphradium;
re, rectum; rg, rectal gland; rs, radular sac; rsa, right siphonal appendage; sg, salivar\ gland; si, siphon; sn, snout; sp, sperm groove;
st, stomach; te, tentacle; vd, vas deferens; vl, valve of Leiblein; vo, ventral opening of prostate.
J. H. Leal and P. Bouchet, 1989
Page 9
mo
Page 10
THE NAUTILUS, Vol. 103. No. 1
Table 3. Sanomelon viperinus new genus, new species. Lin-
ear shell measurements (mm) and whorl counts. ,-\ll specimens
from the t\ pe localit\ . off Rio de Janeiro State, 2-3°-47'S. -12°10'\\'.
610 m (ieptli III = 11, inchKlinu; liolotspe and paratypes 1-7).
Character
Range
X
SD
Total length
27.3-4.5.4
37.6
6.5
Shell width
11.2-16.6
13.6
1.7
Length last whorl
19.2-31.9
25.9
4.2
Aperture length
14.7-24.3
19.6
3,5
Aperture width
4.1-5.8
4.9
0,6
Protoconch diameter
3.3-3.9
3.5
0.2
Teleoconch whorls
5.75-7.00
6.59
0.51
Protoconch whorls
2.25-3.00
2.70
0.19
.A.xial ribs last whorl
15-24
19
3
Length width
2.37-3.02
2.73
0.19
.Aperture length length
0.49-0.54
0.52
0.02
.Aperture length ap u idtli
3.38-4.42
3.97
0.31
relatively short, tightly wound, surrounded by connec-
tive tissue, innervated, through its anterior extremit\ , by
nerve of gland of Leiblein (lb). Ner\e of gland of Leiblein
originating at left buccal ganglion in circumoesophageal
nerve ring. Posterior oesophagus (pe) very long. Stomach
(st) short, embedded in digestive gland and showing as
slight swelling of posterior oesophagus. Intestine (in) rel-
ativelv long. Rectum (re) slightly swollen in preserved
specimens. Rectal gland (rg) brownish-gray, elongated,
with many lateral, short and blunt branches. .\ short,
posterior duct (dr) connects rectal gland to dorsal surface
of rectum. Anus (an) with weak anal papilla.
Radula (figures 23-25): Radular ribbon (figure 23)
uniserial. Rachidian teeth (figures 24, 25) 0.14 mm wide,
tricuspid, with crescent-shaped basal plate. Cusps situ-
ated in appro.ximately same plane as basal plate, growing
from its posterior margin. Central and lateral cusps
cur\ed, all about same length, extremit\ of central cusp
slightK more posterior than extremities of lateral cusps.
Dorsal surface of rachidian deeply impressed by cusps
of preceding teeth. When teeth are in same plane (parts
of radula not in protracted condition), extremities of
cusps (figure 23) interlock w ith base of cusps of adjacent
tooth.
Male reproductive system (figure 33): Testis grayish-
white, situated along adapical part of digestive gland.
Seminal vesicle extremely convoluted, anterior to kidney.
Vas deferens (vd) straight, joins prostate. Prostate (pr)
cream-white, ventrally opened (vo), internally and dor-
sally grooved (do), laterally bilobed (Ip). Prostate and
rectum loosely joined by connective tissue. Sperm groove
(sp) becomes open posterior to anus. Open sperm groove
runs to distal extremity of penis. Penis (pe) small, without
papilla, situated immediately behind right lateral head
lobe, not folded back inside mantle cavity.
Holotype: MORG 25469, 44.2 mm length, 15.3 width,
MD-55 station CB-105 (Blake trawl), off Rio de Janeiro
State, Brazil (collected alive).
Type locality: Continental slope off the coast of Rio de
Janeiro State, Brazil, 23°47'S, 42°I0'\V, 610 m depth,
mudd\ bottom.
Paratypes: Paratypes 1-5, MNHN, MD-55 station CB-
105 (Blake trawl) (type locality); Paratypes 6, 7, MN'RJ
5762. MD-55 station CB-104 (Blake' trawl). 23''42'S,
42°07'W. 430-450 m depth, mudd\ bottom.
Other material examined: MNHN, 7 juvenile speci-
mens. lOjuvenileor damaged shells; MN'RJ 5763, 3 shells,
10 juvenile or damaged shells; MORG 25470, 3 speci-
mens, 8 juvenile specimens, 3 juvenile shells + 2 dissected
animals, MD-.55 station CB-105 (Blake trawl) (type lo-
cality); MORG 25471, 1 juvenile shell, MD-55 station
CB-i06 (Blake trawl), 23°54'S, 42°ir\V, 830 m depth,
mudd\' bottom; MORG 25472, 10 juvenile or damaged
shells, MD-55 station CB-104 (Blake trawl) (same locality
as Paratypes 6, 7).
Remarks: Nanomelon differs in shell and radular mor-
pholog)' from all known zidonine genera; shell sculpture
is similar to Minicyrnbiola corderoi and Miomelon alar-
coni Stuardo & X'illarroel, 1974, having a similar, clath-
rate and chalky white surface. Clathrate or reticulate
sculpture is also found in some species of the operculate
volutid genus Fiisivoluta (Calliotectinae), from deep-
water off South .Africa (\N'eaver & Dupont, 1970; Liltved
& Millard, 1986). Nanomelon viperinus has a very elon-
gated shell for a Zidoninae (length/width = 2.73 ± 0.19
mm, n = 10): see table 3 for other measurements and
Weaver and Dupont (1970) for shell dimensions in other
genera. Within the subfamily, general similarity is re-
stricted to the New Zealand species comprising the genus
AlcUhoelflerningi Dell, 1978, lutea (Watson, 1882), wil-
sonae (Powell, 1933); see Dell. 1978]. S'anomelon vi-
perinus is easily separated from the Alcithoe species group
by its smaller total shell length and relative smaller aper-
tural size, general shell proportions (table 3), fainter col-
umellar plicae, clathrate sculpture, characteristic set of
crowded spiral cordlets in the sutural shelf, and larger
number of protoconch whorls. The shell ultrastructure
agrees with the basic arrangement described by Hara-
sewych (1987) for Tractolira germonae. the most re-
markable difference being the presence of a thin, internal
lining of prismatic crystals (figures 16-18, d). B0ggild
(1930), examining thin sections of the shell of "Voluta
sp.", described three layers: the most external layer finely
prismatic, and the second and third la\ers crossed-la-
mellar. The third layer was portrayed by B0ggild as
composed of two sub-layers. The new genus also has
distinct radular characters. Cusps of the rachidian teeth
are very elongated for a zidonine, the basal plate is wide
but not much curved (compared to those of Alcithoe),
the lateral cusps have straight outer margins, and the
tips of the cusps produce characteristic, relatively deep
depressions in subsequent teeth.
The alimentar\ system of Nanomelon is characterized
b\ accessor)' salivary glands looseK connected to the
primary salivary glands, not tightly wound around them,
and the stomach lacks an anterior tubular extension.
J. H. Leal and P. Bouchet, 1989
Page 11
Clench and Turner (1964) have considered the small
degree of physical association between the two types ol
salivary glands as one of the characters distinguishing
Zidoninae from Odontocymbiolinae, in which the ac-
cessory salivary glands are tightly wound around the
principal salivary glands.
Nanomelon viperinus has a relativeK' large rectal gland
that opens in to the rectum through a duct situated in its
posterior part, as opposed to that in Alcithoe arabica
(Gmelin, 1791) (Zidoninae, Ponder, 1970) and Tractolira
germonae (Odontocymbiolinae. Harasewych, 1987), in
which the rectal gland branches posteriorly from an an-
terior duct situated immediately behind the anus. The
new species has a male reproductive s\stem typical of
the Zidoninae; Novelli and Novelli (1982) have noted
that the presence of a ventralK' opened prostate, and an
open sperm groove running to the distal extremity of the
penis are characters unique to the subfamily. Their con-
clusions were based on their own work on Adelomelon
ancilla (Lightfoot, 1786), A. beckii. A. brasiliana. Zidona
dufresnei, and Provocator corderoi and on data from
Woodward (1900), Clench and Turner (1964), and Pon-
der (1970). The Odontocymbiolinae have a closed sperm
duct along the mantle cavity floor and penis (Clench &
Turner, 1964; Harasewych, 1987).
DISCUSSION
In the process of assigning the above species and genera
to the different subfamilies, it became evident that some
traditionally used characters are convergent, or primitive
at their respective levels of utilization. For instance, shell
size, general outline, number and shape of a.xial ribs in
early teleoconch whorls, and shape and internal color-
ation of the aperture render O. aimulatrix superficially
similar to subadults of Adelomelon riosi Clench & Turn-
er, 1964. The latter zidonine volute has accessory salivary
glands loosely wound around moderately compacted sal-
ivary glands, and rachidian teeth with the cusps and basal
plate roughly in the same plane (figures 3, 4, 22). The
two species were found microsympatrically during the
MD-55 cruise, Blake trawl CB-104, at 430-450 m depth.
The new species is also conchologically similar to the
Fulgorariinae species Nipponomelon prevostiana (Crosse,
1978), iV. magna (Kuroda & Habe, 1950), Musashia hi-
rasei (Sowerby, 1912), and M. cancellata Kuroda & Habe,
1950. all from the western Pacific (see descriptions and
illustrations in Kuroda & Habe, 1950; Shikama, 1967;
Weaver & Dupont, 1970; Moore, 1984; Okutani et al.
1988).
Convergence in shell shape between representatives
of the volutid subfamilies Odontocymbiolinae and Zi-
doninae has been observed previousK'. The conchological
mi.xing of Odontocymbiola magellanica and Adelomel-
on ancilla (Lightfoot, 1786) by Pilsbry and Olsson (1954)
and the consequent taxonomic implications at supraspe-
cific levels were noted b\ Clench and Turner (1964) in
the original description of Odontoc\mbiolinae. The su-
perficial conchological convergence of Odontocymbiola
simulatrix with certain species of Fulgorariinae as well
as w ith A. riosi, provides further evidence of the unre-
liability of shell characters in the supraspecific volutid
taxonomy.
Some anatomical characters used in subfamilial tax-
onomy ma\ also be convergent. The "loosely wound"
condition of the accessory salivary glands, considered to
be diagnostic of the Zidoninae, is found not only in all
known alimentary systems in the subfamily (this paper;
Clench & Turner, 1964, Ponder, 1970; Novelli & Novelli,
1982), but also in the odontocymbioline Miomelon alar-
coni Stuardo & Villarroel, 1974, and probably in Trac-
tolira germonae Harasewych, 1987 (the illustration given
in the original description of this species depicts a rela-
tionship between the two types of salivary glands more
likely to be found in a zidonine species).
Radular and male reproductive system characters are
apparently more adequate to define the above subfam-
ilies, although much variation in radular morphology is
found in the Zidoninae (see Weaver & Dupont, 1970).
This latter family can also show convergence in radular
morphology with the Fulgorariinae (Stuardo & Villar-
roel, 1974). The presence of a ventrally open prostate,
and an open sperm groove are characteristic of the Zi-
doninae, as opposed to the sperm duct closed from the
mantle floor to the penis of the Odontocymbiolinae; one
of the two conditions is most probably primitive at the
subfamilial level. The suitability of the above characters
to formulate a higher classification of the V'olutidae will
be decided only after careful phylogenetic analysis of
the family on a world-wide basis, a task which is beyond
the scope of this regionally based work.
ACKNOWLEDGEMENTS
We are indebted to Alain Guille, Museum National
d'Histoire Naturelle (MNHN), Paris and to J. M. Ramos,
Universidade Santa Ursula, Rio de Janeiro, for their ef-
forts as Chief Scientists during the MD-55 cruise con-
ducted for Terres Australes et .Antarctiques Franfaises.
A. C. S. Coelho, Museu Nacional, Rio de Janeiro, pro-
vided the opportunity for Jose H. Leal to participate in
the cruise and helped with logistic support in Rio de
Janeiro. M. G. Harasewych, National Museum of Natural
History, Washington (NMNH), offered criticisms of the
manuscript and information on the Volutidae. R. S.
Houbrick made possible the study of material in the
NMNH and reviewed the manuscript. E. C. Rios, Museu
Oceanografico, Rio Grande, Brazil, gave information on
the Brazilian volutids and kindly loaned material for this
study. B. Metivier, MNHN, collaborated in all stages of
this work, especially aboard the Marion-Dufresne. P.
Lozouet, MNHN, prepared the photographs in figures
1-13. Prof. C. Levi made it possible for Jose H. Leal to
work temporarily as a short-term Associate Assistant in
MNHN. We thank P. Blackwelder, Electron Microscopy
Laboratory, Rosenstiel School of Marine and Atmospher-
ic Science, Miami, for information on shell ultrastructure
and for use of the scanning electron microscope under
Page 12
THE NALTILLS, Vol. 103, No. 1
her charge. This work was supported in part by a Doc-
toral Scholarship from Conselho Nacional de Desenvol-
vimento Cientifico e Tecnologico, Brazil, to Jose H. Leal,
and by travel funds from the same agency. .Additional
funding was pro\ided in part b\ the Bader Memorial
Student Research Fund, USA.
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monograph of the Recent \'olutidae of the world. Dela-
ware Museum of Natural History. Greenville, .w -I- 375 p.
Woodward. M. F. 1900 Note on the anatomy of Valuta
ancilla (Sol.), Nctuneopsis gilchristi (Sby.) and Volutilith-
es ahyssicola (Ad and Rve). Proceedings of the Malaco-
logical Society of London 41 17-125.
THE NAUTILUS 103(1):13-19, 1989
Page 13
Pleioptygmatidae, a New Family of Mitriform Gastropods
(Prosobranchia: Neogastropoda)
James F. Quinn, Jr.
Florida Marine Research Institute
Department of Natural Resources
100 Eighth Avenue S.E.
St. Petersburg, FL 33701, I'SA
ABSTRACT
A new familv' of mitriform gastropods is proposed, comprising
only the genus Pleioptygma Conrad, 1863, based on exami-
nation of the foregut of P. helcnae (Radwin & Bibbe\, 1972).
Pleioplygma helenae is the onK known Recent species of this
genus, known otherwise only from species from the Neogene
of the southeastern United States. The unique configuration of
the foregut separates this group from the Mitridae, Costellarii-
dae, N'olutidae, and Turridae. Features characterizing Pleiop-
tygmatidae include a proboscis introvert not connected to the
head, presence of a proboscis bulb, buccal mass divorced from
the proboscis, and h> pertrophied rhachidian tooth.
Key words: Pleiopt\gmatidae; systematics; Pleioptygma hel-
enae; anatomy.
INTRODUCTION
The genus Pleioptygma Conrad, 1863 (type species Va-
luta carolinensis Conrad, 1840), comprises a small group
of species with medium to large (attaining 150 mm in
length) mitriform shells bearing conspicuous spiral cords,
columellar plicae, and siphonal notch. Three nominal
species and one subspecies [P. carolinense. P. lineolatum
lineolatum (Heilprin, 1887), P. lineolatum saginatum
(Tucker & Wilson, 1933), and P. prodroma (Gardner,
1937)] and possibly two additional unnamed species are
represented in the Tertiary formations of the southeast-
ern United States (W. G. Lyons, personal communica-
tion). Only one Recent species, P. helenae (Radwin &
Bibbey, 1972), is known. Numerous specimens of P. hel-
enae have been collected from the continental shelf off
northern Honduras (Caribbean), an area in which several
endemic, relict genera have been found (Petuch, 1981,
1982a,b; Houbrick, 1986).
MATERIALS AND METHODS
Prior to the summer of 1983, all know n specimens of P.
helenae were shells that had been carried into lobster
pots by hermit crabs. However, through the efforts of
Mr. Louis Kotora, a St. Petersburg shell dealer, a live-
collected specimen of P. helenae (male, shell length 69
mm; figure 1) was obtained from lobster fishermen in
July-August, 1983. The specimen, collected near Hon-
duras, was preserved in commercial rubbing alcohol (iso-
propanol) and brought to W. G. Lyons at the Florida
Marine Research Institute, Department of Natural Re-
sources, for examination. The animal was extracted from
the shell and given to me by Lyons for dissection. A
preliminary report on that specimen was presented at
the 1984 American Malacological Union Meeting (Quinn
& Lyons, 1984). Subsequently, Mr. Kotora provided
another live-collected specimen (female, shell length 107
mm) preserved in rum and collected from the same area
as the first specimen. Because the shells were to be main-
tained intact, the animals were extracted by hooking
them with a dental probe and pulling with a twisting
motion, with consequent loss of most of the posterior
portions of the two animals. This report describes the
anatomy of the anterior portions, especially the foregut
morphology, and proposes the new family Pleioptyg-
matidae, based on differences from the foregut mor-
phologies of the Mitridae, Costellariidae, Volutidae, and
Turridae.
SYSTEMATICS
Superfamily Muricoidea Rafinesque, 1815
{sensu Ponder & Waren, 1988)
Family Pleioptygmatidae Quinn new family
(type genus Pleioptygma Conrad, 1863, Miocene-Re-
cent)
Genus Pleioptygma Conrad, 1863
(type species Voluta carolinensis Conrad, 1840, Plio-
cene)
Superfamilial placement follows Ponder and Waren
(1988). Foregut anatomy of Pleioptygma helenae (Rad-
win & Bibbey, 1972) (discussed below) suggests a taxo-
nomic ranking equal to that of the Mitridae and Cos-
Page 14
THE NAUTILUS, Vol. 103, No. 1
Figure I. Pleiuplyginu ItiUnat: lUadwiii 6i Bibbe\, 1972).
Ventral and dorsal views of shell of 69 mm male specimen.
tellariidae. Pleioptygma is derived from the Greek pleios,
full, and ptijgma, a fold. Contrary to common usage, the
genus name is neuter, not feminine. Species names in
this account are emended accordingly.
Pleioptygma helenae (Radwin and Bibbey, 1972)
Type locality: Gorda Bank, Honduras (here emended).
The type material of P. helenae was reported to be from
Cay Sal Bank, Bahamas (Radwin & Bibbey, 1972), ob-
tained from lobster pots by a commercial lobster fish-
erman. Subsequent attempts to collect this species from
Cay Sal have failed. Instead, numerous specimens have
been collected from the banks off Honduras. I believe
the original types were mislocalized and here propose
that the type locality be emended to the Gorda Bank,
Honduras.
ANATOMY
External features: The remains of the animals after
extraction comprise about one whorl each (figure 2); the
posterior portion of the mantle cavity and all organs
posterior to that were lost. The foot is large, very mus-
cular, uniformly tan to cream in the larger specimen;
the smaller is dark chocolate-brown on the sole, fading
to grayish buff above. The head is small relative to the
foot, with conical snout; the tentacles are short, tapering
to rather blunt tips, with conspicuous black eyes located
at about midlength. The penis is broad, dorsoventrally
flattened, folded to right, and located to the right of the
head. The mantle is very thin except for a narrow (1-2
mm wide), muscular band along the edge; the posterior
portion and right side are badly torn in both specimens.
The siphon is large, thick-walled, muscular, and lacks
siphonal appendages. The remains of the ctenidium and
osphradium are broad but otherwise unremarkable. The
remains of the h\pobranchial gland indicate that it is
large and rather thick The columellar muscle is broad
and thick.
Proboscis complex: Situated anteriorly is a large, broad
introvert (figures 3, 4, 7). The base of the introvert at-
taches to the right anterior face of the proboscis bulb.
The walls of the introvert are thick near the base but
rapidK become thin; they are composed principalK of
circular muscle forming an outer tube or "sheath and
measuring 11-13 mm long when contracted. DistalK',
the introvert invaginates, runs back through the outer
tube sheath, and enters the proboscis bulb (figure 3). In
the larger specimen (figures 4, 7), the introvert turns
sharply, runs to the left side of the bulb lumen, then
continues diagonally to the right posterior and attaches
to a muscular mound on the floor of the bulb. The total
length of the contracted introvert is approximately 45
mm. In the smaller specimen, the inner tube of the in-
trovert enters the proboscis bulb, where it is packed in
a convoluted mass, exits through the right postero\entral
wall of the proboscis bulb, and finalK joins the buccal
mass. The total length of the introvert is approximately
65 mm.
The proboscis bulb is a large, conspicuous, roughly
rectangular structure attached to the floor of the cephalic
cavity close to the anterior bod\ wall (figures 3, 4, 7).
The bulb of the larger specimen measures 17.0 x 18.5
mm, and that of the smaller 3.5 x 7.5 mm; the greater
length of both is perpendicular to the longitudinal axis
of the head-foot. The bulb is attached to the cephalic
cavit\ floor by a broad, rather thick band of muscle
originating in the foot and inserting along the antero-
dorsal edge of the bulb. The bulb walls are very thick
and are composed of several la\ers of muscle oriented
at right angles to each other. In the smaller specimen,
there are four distinct layers oriented transversely-lon-
gitudinally-transversely-longitudinally (outer to inner
layers, respectively, direction relative to axis of head-
foot). The muscle la}ers of the bulb in the larger spec-
imen are less distinctly demarcated, especiall) the mid-
dle two, because of the addition of much oblique muscle.
Buccal mass: The buccal mass is a muscular, pear-shaped
organ lying, in situ, on its left side just to the right of
the proboscis bulb, its longitudinal axis parallel to the
head-foot longitudinal axis and its morphological ante-
rior end topologicalK posterior (figures 5, 6). The mor-
phological anterior end is separated from the proboscis
bulb b\ a 1.5-2.0 mm length of intro\ert in the smaller
specimen. However, in the larger specimen, the anterior
part of the buccal mass extends through the wall of the
proboscis bulb and protrudes into the bulb lumen as a
prominent mound to « Inch the introvert attaches (figure
7). In the left center oi the mound is a narrow . elongate
opening leading from the introvert to a small pouch into
which the true mouth protrudes. A narrow, thick-walled
tube leads from the mouth to the buccal cavity, a distance
of about 9 mm. of which the first 6 mm run to the right
through the posteroventral wall of the proboscis bulb.
J. F. Quinn, Jr., 1989
Page 15
Figures 2, 3. Pleioptygma helenae. 2. Diagrammatic sketch of animal of 69 mm specimen removed from sfiell, right lateral and
dorsal views. 3. Diagrammatic sketch of foregut of 69 mm specimen. Structures not in life positions, bm, buccal mass; as, esophagus;
f, foot; hg, h\ pobranchial gland; in, intro\ert; m, muscular mantle edge; pb, proboscis bulb; rs, radula sac; s, siphon; ug, unpaired
foregut gland.
The buccal mass walls are very thick, ranging from about
3 mm at the anterior end to about 1.5 mm near the
opening of the esophagus (figure 8). The buccal cavit\'
is spacious, and its floor has a prominent transverse ridge
just posterior to the oral tube opening. The opening of
the radula sac is subrectangular and is located in the
center of the buccal cavity floor. The radula sac is broad,
comprising the posteroventral % of the buccal mass. Just
behind and above the posterior edge of the radula sac
opening is a small (2 mm long, 1 mm wide) caecum.
Directly above the caecum is the opening of the anterior
esophagus.
Radula: The triserial ribbon is broad and short (length :
width = 1.3-1.6:1), about 5% of shell length, with about
70-100 transverse rows of teeth (figures 9-11). The pos-
terior 60% of the rows are above the bending plane, the
anterior 40% below. The rhachidian teeth of the larger
specimen are 1.05 mm wide and the lateral teeth are
1.30 mm wide at the bending plane. The rhachidian and
lateral teeth are similar in structure, each having a broad,
narrow basal plate and numerous equal, sharp, narrowly
triangular cusps (the rhachidian tooth has about 25 cusps,
the lateral teeth about 35).
Esophagus: Upon exiting the buccal cavity, the esoph-
agus curves slightly to left, makes a 180° ventral turn
and runs underneath itself for about 5 mm, continues
through the nerve ring and turns upward, forms a con-
voluted mass of half-loops, and finally runs posteriori)
and diagonalK' to the left beneath the proboscis bulb
(figures 5, 6). A valve of Leiblein was not found.
Salivary glands: These glands are elongate, roughK- lan-
ceolate organs 1\ ing along either side of the radula sac
and are fused at the broad ends (figures 4. 5). The salivar\
ducts are short and do not pass through the nerve ring;
the right duct enters the right side of the radula sac just
posterior and ventral to the external separation of the
esophagus from the buccal mass; the left duct enters on
the left side slightly more ventrally than does the right
duct. No accessory salivary glands were observed.
Unpaired foregut gland: The duct is long (about 25
mm), and coiled into a ball to the right of the buccal
mass (figure 3). The duct terminates in a muscular, lan-
ceolate bulb 4 mm long and 1.8 mm in greatest width.
This organ was found only in the smaller specimen, and
the connection of the duct with the foregut was not
found.
DISCUSSION
Familial placement of Pleioptygma has always been con-
troversial. Conrad (1840) first described the type species
of Pleioptygma as Vohita but soon (Conrad, 1842) trans-
ferred the species to Mitra, a placement followed by
Tuomey and Holmes (1856). However, Conrad (1863)
seemingly reversed himself by assigning his new genus
Pleioptygma to Volutidae. Heilprin (1887) ignored
Pleioptygma and maintained the species group in Mitra.
Dall (1890) noted Conrad's placement of Pleioptygma
in Volutidae but treated it as a subgenus of Mitra, a
classification followed bv Cossmann (1899), Tucker and
Wilson (1933), Gardner (1937), Mansfield (1937), Olsson
and Harbison (1953), and Dubar (1958, 1962). Cerno-
horsky (1970:61) stated unequivocally that Pleioptygma
was not referable to Mitridae but instead possessed many
features "all consistent with the Volutidae. An assign-
ment to the Scaphellinae might be appropriate." Radwin
and Bibbey (1972:95-96) rejected that placement, stating
that characters Cernohorsky (1970) used to exclude
Pleioptygma do occur in some Mitridae, and concluded:
"We are thus tentativeK placing the subgenus Pleiop-
tygma in the Mitridae, pending examination of the rad-
ular dentition of M. (P.) helenae." Cernohorsky (1976:
282) ignored Radwin and Bibbey and maintained that
Page 16
THE NAUTILUS, Vol. 103, No. 1
__. ,^es
5 6
Figures 1—8. Flcioptygma helenae. Foregiit morpholog\ of 107 mm specimen, l-. Diagrammatic sketch ot proboscis complex in
relation to head, cephalic cavity opened dorsaily. 5, 6. Diagrammatic sketches of buccal mass and associated organs. 5. Dorsal
(morphological right side) view. 6. Ventral (morphological left side) view. 7. Dissection of proboscis bulb and introvert, opened
dorsalK 8. Dissection of buccal mass, opened dorsaily. be, buccal cavity; bm, buccal mass; bmi, inner opening to buccal mass;
bmo. outer opening to buccal mass; bw, body wall; c, caecum of buccal mass; dvm, dorsoventrally oriented muscle mass; es,
esophagus; h, head; in, proboscis introvert; ini, inner tube of introvert; ino, outer tube of introvert; m\\ , muscular wall of proboscis
bulb; pb, proboscis bulb; pg, pedal ganglia; r, radula; rs. radula sac; rsg, right salivar> gland; s. siphon.
"shell features of Pleioptygma are consistent with the
\'(iluti(lae, and the genus should be referred to the Volu-
tid subfamily Scaphellinae." Weaver and duPont (1970)
omitted Pleioptygma from their monograph of world-
wide Voiutidae. In all cases, judgments of affinities of
Pleioptygma were based entirely on shell characters. Al-
tliougli bv no means complete, tiie anatomical data pre-
sented above provide an opportunits to re-evaluate the
relationships of Pleioptygma. The following discussion
compares the anatoin\ of Pleioptygma helenae with that
of the Mitridae, Costellariidae, and \'olutidae, the fam-
ilies to which Pleioptygma has traditionalK been assigned,
and the Turridae, a group that has some superficial sim-
ilarities of foregut anatom\ to that of Pleioptygma. Table
1 summarizes the comparison of characters of Pleiop-
tygma and the four families cited above.
J. F. Quinn, Jr., 1989
Page 17
Figures 9-11. Pleioptygma helenae. SEM micrographs of radula. 9. View of portion of intact ribbon, x 25.
teeth. X 100. 11. Lateral teeth, x 100.
Ponder (1972) investigated the anatomies of several
mitriform gastropods from the southwestern Pacific. In
that stud>', he found that the Mitridae, as then perceived,
actually comprised two famihes, the Mitridae s«'n.s(j stric-
to and the Costellariidae (as \ e.xiUidae). I am here fol-
lowing that decision and will compare Pleioptijgnia to
each group separately. Terminology of anatomical struc-
tures follows Ponders (1973) review of the comparative
anatom\' of the Neogastropoda. The shells of the Mitridae
and Pleioptygma are rather similar in shape and sculp-
ture, with spiral ridges predominating; however, the shells
of Pleioptygma are larger and lighter. The radulae of
the two groups are also quite similar, both having comb-
like lateral teeth and multicuspid rhachidian teeth (e.\-
cept in Pterygia, which lacks lateral teeth). Pleioptygma,
however, has hypertrophied rhachidian teeth that are
only slightK less broad than the lateral teeth. Mitrids, on
the other hand, have rhachidian teeth that are noticeably
narrower than the lateral teeth. The radula of Mitra
sigillata Azuma, 1965 (Cernohorsky, 1970, fig. 26), is
most similar to that of P. helenae in that the rhachidian
has about 20 cusps and the laterals have about 22. How-
ever, the rhachidian is only about 60% of the width of
the laterals, compared to 80% in P. helenae. The soft
parts of Pleioptygma offer a number of important char-
acters that differ from the Mitridae. The foot of Pleiop-
tygma is relatively large, whereas that of mitrids is small.
The head of Pleioptygma has a distinct, relatively large
snout; that of the Mitridae has no such structure. An
epiproboscis, one of the hallmarks of the Mitridae, is
missing in Pleioptygma. The proboscis mouth of mitrids
has a strong, muscular sphincter, the "peristomal rim"
of Ponder (1972). In Pleioptygma the proboscis "mouth"
has no such rim, the opening of the snout having assumed
that function. In fact, the true mouth of Pleioptygma
has been divorced from the extensible portion of the
proboscis. Mitrids have a proboscis sac, but it is a very
thin-walled structure, in contrast with the extremely thick-
walled, muscular proboscis bulb of Pleioptygma, how-
ever, it is not \ et possible to determine whether the two
structures are homologous. The prominent unpaired
foregut gland of Pleioptygma is totally absent in the
Mitridae.
Members of the Costellariidae differ from Pleioptyg-
ma in the following characters: 1) axial, rather than spi-
ral, shell sculpture is predominant; 2) lateral teeth of the
radula are simple and sickle-shaped; 3) the rhachidian
is much narrower; 4) an introvert and proboscis bulb are
absent; 5) a pair of accessory salivary glands is present;
and 6) an unpaired foregut gland is present, but it com-
prises only the Gland of Leiblein (see Ponder, 1972).
The volutids are distinguished from Pleioptygma by
the following characters: 1) the dominant shell sculpture,
when present, is axial; 2) the radula is usualK' uniserial
with a uni-, tri-, or multicuspid rhachidian that is com-
paratively much smaller; 3) siphonal appendages are
present; 4) an introvert is absent; 5) there is no proboscis
bulb; 6) paired accessory salivary glands are present; and
7) an unpaired foregut gland is present but comprises
only the Gland of Leiblein.
The proboscis complex of Pleioptygma is superficially
similar to that of some turrids, particularly to some mem-
bers of the subfamily Daphnellinae (Smith, 1967). In the
Turridae, the buccal mass has also been divorced from
the proboscis tip; there is an introvertible proboscis, at
least in Philbertia (Smith, 1967); and the poison gland
resembles the unpaired foregut gland of Pleioptygma.
However, the proboscis bulb of Pleioptygma and the
hypertrophied rhachidian teeth with comb-like laterals
are not similar to any described turrid, and most of the
similarities in other structures are analogous, not ho-
mologous.
Considering the anatomical differences separating
Pleioptygma from other similar neogastropod families,
it is evident that Pleioptygma cannot be accommodated
in any defined family. Therefore, I propose that Pleiop-
tygma represents a family separate from other mitriform
gastropods, defined principalK' by the autapomorphous
proboscis complex, the position of the buccal mass, and
the hypertrophied rhachidian tooth. The morphologies
Page 18
THE NAUTILUS, Vol. 103, No. 1
Table 1. Comparison of shell and anatomical characters of Pleioptygmatidae, Mitridae. Costeliariidae, V'olutidae, and Turridae.
Characters of Mitridae and Costeliariidae from Ponder (1972. 1973); X'olutidae from C^iench and Turner (1964) and Ponder (1973);
and Turridae from Smith (1967).
(. Iiaraeter
Plei(ipt\ gmatidae
Mitridae
Costeliariidae
\ olutidae
Turridae
Shell size
Predominant sculp-
ture
Radula
Lateral teeth
Rhachidian teeth
Foot
Medium to large Medium to large Small to large
Spiral Smooth or spiral .\xial
Multicuspid
Small to large
Smooth or axial
Small to large
Smooth, a.vial or spiral
Multicuspid or ab- Simple, curved UsualK absent Usually absent, mul-
sent ticuspid if present
Wide multicuspid Usually relatively Relatively large. Small to large. Usually absent, rudi-
small, multicus- 3 or more uni-, tri-, or mentary if present
pid cusps multicuspid
Large relative to Small relative to Moderate relative Large relati\e to Small relative to shell
shell shell to shell shell
Siphonal append-
Absent
Absent
Absent
Present
Absent
ages
Alimentary canal
Proboscis
Introvert
Pleurembolic
Pleurembolic
Pleurembolic
PoKembolic or in-
traembolic
Proboscis bulb
Present
.■\bsent
Absent
Absent
Absent
Epiproboscis
Absent
Present
Absent
Absent
Absent
Mouth
At base of probos-
M tip of probos-
.-^t tip of probos-
.\t tip of probos-
At base of proboscis.
cis complex, no
cis, with peris-
cis, no peristo-
cis
with peristomal
peristomal rim
tomal rim
mal rim
rim
Accessory sali-
Ab.sent?
Absent
Present, paired
Present, paired
Usuall) ab.sent. paired
vary glands
if present
Unpaired fore-
Present
.•\bsent
Present
Present
Present, poison gland
gut gland
Valve of Leib-
.absent
Small or absent
Large
Large
Absent
lein
of radula and shell suggest that the Pleioptygmatidae
may be derived from an ancestor in common with the
Mitridae.
The function of the proboscis complex of Pleioptygma
can only be surmised. In both specimens dissected for
this study, the anterior portion of the introvert was lying
free in the cephalic cavity, wedged between the anterior
face of the proboscis bulb and the body wall. Evidence
was lacking of any previous attachment to the walls of
the snout. This position may have been the result ot
extreme retraction in response to contact with the pre-
servative. In life, the introvert probably lies partially
within the snout and is everted by a combination of
contractions of the intrinsic circular muscle of the intro-
vert and muscular contraction of the proboscis bulb. The
length of the everted introvert must be quite extensive,
as the wrinkled and folded appearance of the contracted
organ indicates. The length of the introvert, combined
with the dissociation of the radula from the proboscis
complex, suggests that Pleioptygma helenae feeds on
soft-bodied prey, probably tubicolous and crevice-dwell-
ing worms such as sipuiiculans and poKchaetes. The
position of the radula suggests that it acts principally in
triturating prey and transferring food into the esophagus.
Two major differences were observed between the tw o
specimens. The presence of an unpaired foregut gland
in the smaller specimen, but not in the larger, cannot be
explained. Differences in the length and morphological
relationship of the introvert and proboscis bulb, as well
as the morphological relationship of the proboscis bulb
and buccal mass, may be explained by ontogenetic
changes. ApparentK', the total length of the introvert
develops relatively early in ontogeny , at least b\ the time
the shell reaches a length of about 70 mm. With growth,
the length remains about the same, but the width in-
creases. Increasing the volume of the lumen of the pro-
boscis bulb also allows the enclosed introvert to become
less convoluted. In a similar manner, grow th ot the pro-
boscis bulb, especially the thickening of the bulb w alls,
encroaches upon and eventually envelops the morpho-
logical anterior end of the buccal mass, creating the
mound on the Door of the proboscis bulb.
ACKNOWLEDGEMENTS
I thank Mr. Louis Kotora, w ho provided the two speci-
mens of Pleioptygma helenae, and William G. Lyons,
who allowed me to perform the dissections, provided the
J. F. Quinn, Jr., 1989
Page 19
shell photographs, and made valuable comments on the
manuscript Thomas H. Perkins, David K. Camp, Dan
C. Marelli, and two anon\ mous re\ iewers also provided
critical comments on the manuscript
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Houbrick. R. S. 1986. Discover) of a new living Cerithioclava
species in the Caribbean (Mollusca: Prosobranchia: Ceri-
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\lanshcld. W. C. 1937. Mollusks of the Tampa and Suwannee
Limestones of Florida. The Florida Geological Survey,
Geological Bulletin 15:1-334.
Olsson, A. .A. and A. Harbison. 1953 Pliocene Mollusca of
southern Florida « ith special reference to those from North
Saint Petersburg « ith special chapters on Turridae, Vitri-
nellidae and fresh-water mollusks. The .Academy of Nat-
ural Sciences ot Philadelphia, Monographs 8:v -I- 457 p.
Petuch, E. J. 1981. A volutid species radiation from northern
Honduras, with notes on the Honduran Caloosahatchian
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Society of Washington 94(4):1 110-1)30
Petuch, E. J. 1982a. Geographical heterochronv: contempo-
raneous coexistence of Neogene and Recent molluscan
faunas in the .Americas Palaeogeography, Palaeoclima-
tology, Palaeoecologv' 37:277-312.
Petuch, E. J. 1982b. Paraprovincialism: remnants of paleo-
provincial boundaries in Recent marine molluscan prov-
inces. Proceedings of the Biological Society of Washington
95(4):774-780.
Ponder, W. F. 1972. The morphology of some mitriform
gastropods with special reference to their alimentary and
reproductive systems. Malacologia 1 1(2):295-342.
Ponder, W. F. 1973. The origin and evolution of the Neo-
gastropoda. Malacologia 12(2):295-338.
Ponder, W. F. and .A. Waren. 1988. .Appendix. Classification
of the Caenogastropoda and Heterostropha — a list of the
family-group names and higher taxa In: Ponder, W. F.
(ed). Prosobranch ph\ logeny Proceedings of a symposium
held at the 9th International Malacological Congress, Edin-
burgh. Scotland, .August 31-September 6. 1986. Malaco-
logical Review, Supplement No. 4:288-317.
Quinn, J. F., Jr and W. G. Lyons. 1984. .A partial description
of the anatom\ of Pleioptygma helenae (Gastropoda: Neo-
gastropoda). .American Malacological Bulletin 3(l):97-98
(abstract).
Radwin, G. E. and L J Bibbey. 1972. .A new mitrid from
the western .Atlantic. Transactions of the San Diego Societ\
of Natural History 17(7):95-99.
Smith, E. H. 1967. The proboscis and oesophagus of some
British turrids. Transactions of the Ro\al Society of Edin-
burgh 67(1): 1-22.
Tucker, H. I. and D. Wilson. 1933. .A second contribution to
the Neogene paleontolog)' of South Florida. Bulletins of
-American Paleontology 18(66):1-21.
Tuomey, M. and F. S. Holmes. 1855-1857. Pleiocene fossils
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ware Museum of Natural History, Monograph Series No.
l:xv + 375 p.
THE NAUTILUS 103(1 ):20-23, 1989
Page 20
The Reproductive Anatomy and Taxonomic Status of
Philomycus venustus Hubricht, 1953 and
Philomijcus bisdosus Branson, 1968
(Pulmonata: Philomycidae)
H. Lee Fairbanks
The Penns\ Kania State Universit\
Beaver Campus
Monaea, PA 15061, USA
ABSTRACT
Specimens of Philomycus venustus and P. Insdosus were col-
Ifcted from their type localities. Reproductive-system mor-
phology of these two species is described. C^omparisons of ex-
ternal characteristics re\ eal clear, consistent differences in mantle
pattern, length, and foot-edge color. Comparisons of the re-
productive s\stems demonstrate specie.s-specific differences in
penial anatomy. These data support the recognition of Philo-
mycus hisdostis as a distinct species
INTRODUCTION
Thirty years ago it was not unusual to describe terrestrial
gastropods using only external characters. One such de-
scription was that of Philomycus venuslus Hubricht, 1953.
Philomycus hisdosits Branson, 196S was described on the
basis of external and selected reproductive-system char-
acteristics. Hubricht (1974:33) placed P. bisdosus in the
synonymy of P. venustus noting that the latter species
was "... a rather variable species and P. bisdosus rep-
resents one extreme of that variation." Because no com-
parative data vN'ere presented in support of the synony-
mization, the present study was initiated. Specimens of
P. venustus and P. bisdosus were collected, and com-
parisons involving their external and reproductive-sys-
tem morphologies were made. The goals of this study
were: (1) to describe the reproductive-system anatomy
of Philomycus venustus and of P. bisdosus and (2) to
obtain comparative data in support of the above noted
synonymization.
MATERIALS AND METHODS
Specimens of Philomycus venustus were collected on
May 22, 1986 and May 21, 1987 at the type locality for
this species (Comer's Rock, Jefferson National Poorest,
Grayson County, Virginia) under the bark of dow n dead
trees (elevation ca. 1,220 meters). Specimens of Philo-
mycus bisdosus were collected from the t\pe locality
(Breaks Interstate Park, Dickensen County, X'irginia) on
May 23, 1986 and May 22, 1987. The specimens were
found approximately 0.3 kilometers northwest of Cold
Spring, crawling on trees (elevation ca. 540 meters).
External characteristics were compared with species
descriptions to ensure correct identification (i.e., length,
mantle color and pattern, and color of foot edge). .\11
specimens were drowned in distilled water and then
dissected. Complete reproductive svstems of each species
were stained and mounted using Gregg's (1959) proce-
dure. Additional reproductive systems of each species
were dissected so that the internal structure of the penis
could be described. The Student's ^-test was used to test
for statistical differences between species. .All material
was preserved in 70% ethanol subsequent to dissection.
Collection and dissection dates were essentially the same
in each \ear to remove those differences in anatomy
attributable to differences in the phase of the life cycle.
Voucher specimens have been deposited in the Na-
tional Museum of Natural Histor\, Smithsonian Insti-
tution (USNM 860412 and USNM 860413 for two spec-
imens of Philomycus venustus; USNM 806414 and USNM
860415 for two specimens of Philomycus bisdosus).
RESULTS
.\ total of eight specimens of Philomycus venustus (two
in 1986, six in 1987) and seven specimens of P. bisdosus
(one in 1986, six in 1987) were collected. The mantle
colors and patterns of these specimens agreed with the
type descriptions for each species (figures 1-4). The total
length of each specimen was measured, the results are
shown in table 1. Measurements were taken from active,
but not crawling, slugs. .\ Student's ^test indicated a
significant difference between the lengths of these species.
.•\11 of the P. bisdosus had gray foot margins, all P. ve-
nustus had white foot margins.
H. L. Fairbanks, 1989
Page 21
Table 1. Measurementsof slug length and results of a Student's
Mest comparing the means of the measurements. Length in
niilHmeters; standard deviation in parentheses.
flulonii/cus
bisdosus
Philunujcus
venuslus
N
7
8
Range
40-55
50-65
Mean
46.9(6.31)
( = 3.46 p=<001'
56.8 (4.46)
Degrees of freedom = 13.
Of the specimens collected, the reproductive systems
of four Philomycus bisdosus and si.x P. veniistus were
stained and mounted on glass slides. The reproductive
SNStems of three P. bisdosus and two P. venustus were
prepared such that the internal anatomy of the penises
could be observed.
Genitalia of Philomycus venustus Hibricht, 1953
(FIGURES 6, 8)
Atrium approximately 80% penial length, glandular on
outer surface. Vagina approximately 10% penial length.
Spermathecal duct diameter greater than that of oviduct,
length slightly less than that of oviduct, slight taper to-
ward spermatheca. Spermatheca round. Dart sac and
dart smaller than spermatheca diameter. Penis thick,
straight, slightly tapered distalK ; penial sheath reaching
junction of vas deferens and penis. Internally, penis with
chamber at distal end, chamber containing several pa-
pillose ridges; chamber length equal to approximately
40% of penial length. \'as deferens encircles distal end
of penis, entering penis along outer edge. Vas deferens
approximately 'IVi times length of spermathecal duct,
approximately same diameter throughout its length. Pe-
nial retractor as wide as distal end of penis, length ap-
proximately 80% penial length. Accessory retractor pres-
ent, small compared to accessory retractor of P. bisdosus.
3
Figures 1-4. Mantle patterns. I, 2. Philomycus venustus. 3,
4-. Philomycus bisdosus. Scale bar ecjuals 5 mm.
Genitalia of Philomycus bisdosus Branson, 1968
(figures 5, 7)
Atrium length approximately equal to penial length,
glandular on outer surface. Vagina approximately 15%
penial length. Spermathecal duct diameter about same
as that of oviduct, length approximately IV2 times that
of oviduct, slight taper toward spermatheca. Sperma-
Table 2. Reproductive system measurements and results of Student's (-tests comparing the means of the measurements. Mea-
surements in millimeters obtained via an ocular micrometer. Standard deviation in parentheses.
Philomycus bisdosus
Philomycus
venustus
(N
= 3)
(N =
4)
(
Range
Mean
Range
Mean
Probability'
Spermatheca diameter
2.4-3.9
3.2 (0.76)
3.8-5.5
4.7(0.75)
2.60
<0.052
Length of spermathecal duct
9 1-10.3
9.8 (0.64)
8.6-13.8
10.5 (2.26)
0.59
>0.50
Maximum diameter of spermathecal duct
1.0-1.2
1.1 (0.12)
1.2-1.5
1.4(0.13)
3.16
< 0.052
Length of oviduct
6.9-8.1
7.6 (0.62)
10.9-16.0
13.2 (2.25)
4.74
<0.0P
Maximum diameter of oviduct
0.7-1.0
0.9(0.15)
0.8-1.2
1.0(0.17)
0.82
>0.40
Length of vas deferens
17.9-19.8
19.1 (1.02)
18.8-22.4
21.3(1.66)
2.16
>0.05
Maximum diameter of vas deferens
0.4-0.7
0.6(0.15)
0.5-0.7
0.6(0.10)
—
—
Length of penial retractor
4.3-7.2
5.4(1.57)
2.2-8.1
4.9(2.55)
0.32
>0.50
Maximum width of penial retractor
0.5-0.8
0.7(0.15)
0.9-1.7
1,4(0.36)
3.50
< 0.022
' Degrees of freedom = 5
- Significant difference at 5Tc level.
Page 22
THE NAUTILUS, Vol. 103, No. 1
Figures 5, 6. Genitalia. 5. Philomycus bisdosm. 6. Philo-
mycus ueniistus. Scale bar equals 10 mm. A, atrium; AG, al-
bumen gland; DS, dart sac; G. gonad; GP, genital pore; HD,
hermaphroditic duct; P, penis; PR, penial retractor; S, sper-
matheca; SD. spermathecal duct; UV, free oviduct; V. vagina;
\ D, vas deferens.
theca round. Dart sac and dart smaller than spermatheca
diameter. Penis diameter at proximal end approximately
5 times distal diameter, with distal diameter slightly less
than diameter of vas deferens. Penial sheath reaches
junction of penis and vas deferens. InternalK , penis with
apparent continuation of the vas deferens reaching prox-
imal end of penis. Proximal end of penis projected into
upper atrium. Vas deferens enters distal end of penis at
its center. Vas deferens approximately I'/s times length
of spermathecal duct, somewhat swollen at penial end,
but approximately same diameter throughout its length.
Penial retractor width approximateK' equal to \as def-
erens diameter, length equal to length of penis. Accessory
retractor present, two times accessory retractor width of
P. venustus.
Nine measurements of various organs of the repro-
ductive systems were obtained (table 2). Student's f-tests
indicate significant differences (probability < 0.05) in
four of the measurements: spermatheca diameter; max-
imum diameter of spermathecal duct; length of oviduct;
maximum width of penial retractor.
DISCUSSION
In their discussion of terrestrial slugs, Chichester and
Getz (1968:159) stated that "In almost all species the
distal genitalia are specifically diagnostic." Fairbanks
Figures 7-9. Longitudinal sections of the penises. 7. Philo-
mycus Imdosus. 8. Philomycus venustus. 9. Philomycus lo-
gatiis (9 from Fairbanks, 1986). Scale bars equal 10 mm. \.
atrium; AR, accessory retractor; DS, dart sac; P, penis; PR,
penial retractor; PS, penial sheath; SD, spermathecal duct; UV,
free oviduct; V, vagina; YD, vas deferens.
(1986) has shown species-specific differences in the distal
genitalia (penial anatomy) for two s\mpatric species of
Philomycus. P. togatits Gould, 1841 and P. carolinianus
Bosc, 1802. Comparisons of the reproductive sNstem
anatomy of P. bisdosus and P. venustus (this study) dem-
onstrated consistent comparable differences in penial
anatomy (figures 7, 8). These two species have not been
shown to be sympatric, howe\er each is s\ mpatric with
P. togatus and P. carolinianus. Four other reproducti\e-
system characteristics (table 2) provided additional sup-
port for species-specific differences between P. bisdosus
and P. venustus.
Examination of external characters of Philomycus
venustus and P. bisdosus also revealed diagnostic dif-
ferences between these two species. The mantle patterns
were clearly different (figures 1-4), so the specimens
were alwa\s easiK' separable. P. venusttis was a signifi-
cantly larger slug than P. bisdosus (table 1). A consistent
difference in foot margin color provided a third discrim-
inating character.
Hubricht 1 1974 I did not specif) which characters were
used to provide the basis for the synon\ mization of Phi-
lomycus bisdosus with P. venustus. Because no definitive
H. L. Fairbanks, 1989
Page 23
data were available for comparisons of reproductive-
system anatomy, one might conclude that the mantle
pattern u as the primar\ criterion. However, in the pres-
ent study, P. venustus always had transverse oblique
bands, either solid or broken into spots. P. bisdosus had
no transverse bands. This, coupled with the size differ-
ence, foot margin color difference and the differences in
penial anatomy, lead to the conclusion that the specimens
examined for this study were representatives of two dif-
ferent species. .Accordingly, Philomycus bisdosus Bran-
son, 1968 should be recognized as a distinct species.
Based upon e.xternal appearance and size, Philomycus
bisdosus was most similar to P. togatus (see Fairbanks,
1986). The mantle patterns of these two species were
similar, and the reproductive systems appeared similar
in gross morphology. However, the foot margin of P.
bisdosus was gray, whereas that of P. togatus was orange.
In addition, the penial anatom\- was different (figures 7,
9) and P. bisdosus had a dart sac and dart that was
appro.ximately 65% the size of the dart sac and dart of
P. togatus.
ACKNOWLEDGEMENTS
Financial support for the field trips associated with this
stud\ was provided b\' grants from the Faculty Schol-
arship Support Fund of The Pennsylvania State Univer-
sitv.
LITERATURE CITED
Bosc, L. A. G. 1802. Histoire naturelle des coquilles, contenant
leur description, et leiirs moeurs, Vol. I. Paris, 343 p., 1
pi.
Branson, B. .\. 1968. Two new slugs (Pulmonata; Philomy-
cidae: Philomycus) from Kentiicicv and Virginia. The
Nautilus 81(4):127-133.
(Chichester, L. F. and L. L. Getz. 1968. Terrestrial slugs. The
Biologist 50(3-4):148-166.
P'airbanlcs. H. L. 1986. The ta.vonomic status of Philomycus
togatus (Pulmonata: Philomycidae): a morphological and
electrophoretic comparison with Philnmycuscarohnianiis.
Malacologia 27(21:271-280,
Gould, A. .\. 1841. Report on the invertebrates of Ma.ssacliu-
setts, comprising the Mollusca, Crustacea, Annelida and
Radiata. Cambridge, \iii -I- 373 p., 15 pis
Gregg, W. O. 19.59, A technique for preparing in (ok; mounts
of molluscan anatomical dissections. The .American Mala-
cological Union Annual Report for 1958 25:39.
Hubricht, L. 1953. Three new species of Philomycidae. The
Nautilus 66(3):78-80.
Hubricht, L. 1974. A review of some land snails of the eastern
United States. Malacological Review 7:33-34.
THE NAUTILUS 103(1 ):24-35, 1989
Page 24
Shell and Pallet Morphology in Early Developmental Stages
of Teredo navalis Linne (Bivalvia: Teredinidae)
S. Cynlhia Fuller
Ya-Ping Hu
Rirhard A. Lutz
Institute of Marine and Coastal
Sciences
Rutgers University
New Brunswick, NJ 08903, USA
Michael Castagna
N'irginia Institute of Marine Science
C^ollege of William and Mary
Wachapreague, \'A 23480, USA
ABSTRACT
Dimensions of the shell and provinculum distinguish Teredo
nacalis larvae from the larvae of other bivalve mollusks. In the
present scanning electron microscopic study of shell and pallet
morpholog) during early ontogenetic stages of this species, the
characteristic teredinid pr()\ inculum. with two interlocking pairs
of small teeth and a wide central tooth and socket, was well-
de\eloped in shells 90 tim long. Provinculum length ranged
from 44 to 51 ^lu\ during the larval period. Average lengths of
provincular teeth of the left valve were 9.6 fim for the anterior
tooth and 8.2 ^m for the posterior tooth; in the right valve,
lengths were 7.2 ^lm for the anterior tooth, 16.4 nm for the
central tooth, and 6.6 ^m for the posterior tooth. Larval shell
height prior to metamorphosis was just under 230 fim.
Formation of a ligament pit preceded secretion of the dis-
soconch and development of the apophsses, cond\ les, and ex-
terior denticulated ridges. Shell loss along the posterior margin
of the right valve accompanied a shift in the axis of articulation
from the dorsal to the posterior plane; subsequent secretion of
the dissoconch restored bilateral symmetry. The initial trian-
gular shape of the pallet blade became increasingly rectangular
and finally elongate with growth along the distal margin.
Key words: Larvae; post-larvae; shell morpholog) ; pallets; Te-
redo navalis; teredinid; shipworm
INTRODUCTION
The common shipworm Teredo navalis Linne, 1758, is
distributed worldwide in temperate waters (Turner, 1966,
1971; Abbott, 1974). This larviparous bivalve releases
veligers at the straight-hinge stage of development (Si-
gerfoos, 1908; Grave, 1928; j0rgensen, 1946; Sullivan,
1948; Loosanoff & Davis, 1963; Loosanoff et a/., 1966;
Turner, 1966, 1971; Scheltenia, 1971; Turner & Johnson,
197 1 ). EarK morphology of T. navalis lias been described
with optical photomicrographs and gross shell dimen-
sions at various developmental stages (j0rgensen, 1946;
Sullivan, 1948; Invdi ct at.. 1950; Loosanoff & Davis, 1963;
Loosanoff et al., 1966; Chanley & Andrews, 1971; Cul-
liney, 1975). Jorgensen (1946) characterized the teredi-
nid provinculum as having three teeth on the right valve
and two teeth on the left \al\e; }iowe\er, dimensions of
the lar\ al hinge teetii are not documented for this species.
Growth rates of T. navalis during larval and post-larval
stages were tabulated by Imai et al. (1950). Morpholog)
of the shell and pallets of T. navalis during post-larval
stages has not been described to date.
The present scanning electron microscopic study pro-
vides a comprehensive description of the morphological
features of the shell and pallets of T. navalis during earK-
developmental stages. Provinculum length and dimen-
sions of provincular teeth of T. navalis lar\ae are com-
pared with the same measurements of pre\iousl\ de-
scribed teredinid larvae to facilitate identification of
specimens isolated from plankton samples. Scanning
electron microscopic methods for reproducible and con-
sistent orientation of post-larval specimens are described.
Details of the rapid changes in shell morphology during
metamorphosis are elucidated with micrographs of se-
ciuential developmental stages.
MATERIALS AND METHODS
.Adult specimens of Teredo navalis were collected from
a subticial stake of sweet gum {Liquidauihar styracifltia
Linne) located in a coastal ba\ near W achapreague Inlet,
X'irginia. Larvae (initial shell stages) were removed from
three of these adult shipworms and were reared in fil-
tered (50 nm mesh) ba\\\ater (salinity range = 29.5 to
'34.5%c; temperature range = 22 to 28 °C) using standard
culture techniques (Loosanoff & Davis, 1963). During
metamorphosis, animals colonized several pieces of con-
ditioned wood that were floated in the culture tank.
Larval and post-lar\al samples were treated with a
5.25% sodium Inpochlorite solution for 10 minutes to
remove soft tissues (after Rees, 1950); disarticulated valves
and pallets were rinsed several times with distilled water
and were stored in 95rc ethanol. Specimens were mount-
ed on siKer tape, were coated with approximateK' 600
A of gold-palladium, and were examined using an ETEC
Autoscan scanning electron microscope (SEM). Consis-
S. C. Fuller et al., 1989
Page 25
Figure 1 . Scanning electron micrographs of disarticulated valves of Teredo navalis larvae. Numbers indicate greatest shell dimension
in jum.
tent orientation for documentation of shape was obtained
by positioning larval shells with points of the shell margin
aligned in a plane normal to the electron beam of the
SEM. [For further details of SEM methods for larval
shells, see Fuller et ai. (19S9).] Similar orientation of post-
larval valves was impossible because points along the
post-larval shell margin do not lie in a single plane.
Throughout the post-larval developmental period, how-
ever, points along the dorsoventral margin of the anterior
slope (except those at the extreme ventral region) com-
prise a plane (figure 3). Thus, consistent orientation of
post-larval shells was achieved by positioning specimens
such that this plane was parallel to the electron optical
axis. Additional adjustments were made so that dorsal
and ventral condyles were at an equal working distance.
Shells mounted for documentation of external shell mor-
Page 26
THE NAUTILUS, Vol. 103, No. 1
Figure 2. Scanning electron micrographs of the hinge of dis-
articulated valves of Teredo navalis larvae seen in figure 1.
Numbers indicate greatest shell dimension in ^m.
phology were placed uith the condyles and posterior
slope resting on a specimen mount. v\hich was tilted
slightly for ma.ximal visibility of the external surface.
Shell height is defined as the greatest dorsoventral
dimension. Shell length is defined as the greatest antero-
posterior dimension rougliK parallel to the hinge line in
larval and early post-larval specimens; measurements of
this dimension include the anterior and posterior slopes
as the\ developed in late post-larvae. Shell nomenclature
is taken from Turner (1966, 1971).
Outer surface morphology of the pallets was docu-
mented with the blade positioned approximately per-
pendicular to the electron optical axis. Pallet length is
the distance between proximal and distal ends (see Tur-
ner, 1971:26, for pallet terminology).
RESULTS
Scanning electron micrographs of disarticulated valves
and hinges of sequential ontogenetic stages of Teredo
nucalix larvae are shown in figures 1 and 2. Straight-
hinge larval shells ranged from 77 to 87 fim long (x ±
SD = 81.5 ± 3.3 /urn; n = 30) and from 66 to 70 urn
high (X ± SD = 66.6 ± 1.5 Mm; n = 30). Larvae 90 ^m
long had a well-developed provinculum, with two in-
terlocking pairs of small teeth and a wide, central tooth
and socket. Provinculum length ranged from 44 to 51
txm (x ± SD = 47 7 ± 1.7 ^m; n = 21 ) during the larval
period. Measurements of provincular teeth are summa-
rized in table 1. A low umbo formed in shells approxi-
matel) 120 nm long. During mid-larval stages, the long-
est valve dimension shifted from an anteroposterior to a
dorsoventral axis; larvae were equidimensional at ap-
proximateK 150 ^ni. Length of larv ae just prior to meta-
morphosis ranged from 195 to 210 nm (x ± SD = 202.0
± 4.8 ^m; n = 30); height was just under 230 ^m at this
stage.
The first morphological evidence of metamorphosis
was formation of a ligament pit, which was observed
when valves were approximateK 230 fim high (figures
3, 4). ImmediateK' following formation of the ligament
pit, dramatic morphological changes occurred in the shell
and hinge. An apophysis grew from a base beneath the
anterior tooth and socket; this base extended below the
entire hinge area and expanded at the posterior end to
form an earK dorsal condvle (figures 3, 4, height 240
fim, DC). Formation of the ventral cond>le began with
an inward protrusion of the ventral margin (figure 3,
height 240 //m. XC) Shell loss along the posterior margin
of the right valve began at this stage (figures 3. 5. height
240 nm). An initial denticulated ridge formed on the
anterior margin of the shell exterior (figure 5, height 240
Mm).
A shift in the axis of articulation from an anteropos-
terior to a dorsoventral orientation occurred when shell
height was between 230 and 240 ixm (figure 6). Loss of
approximately 25 to 40 ^m of the shell along the posterior
margin of the right valve accommodated this shift (fig-
ures 5, 7). New ridges were added on the external surface
of the anterior margin (figure 5, height 250 ^m).
Figure 3. Scanning electron micrographs of disarticulated valves of Teredo navalis post-larvae. Numbers above the shells indicate
shell height (greatest dorsoventral dimension); numbers beside the shells indicate shell length (greatest anteroposterior dimension).
Dimensions are in ^m and are accurate to within .5 Mm. .Vrrows designate the dorso\entral margin of the anterior slope; points on
this margin were aligned in a plane for consistent orientation of valves. A, apoplnsis; I)C:, dorsal condyle; \C, ventral condyle; R,
umbonal-ventral ridge.
S. C. Fuller et a/., 1989
Page 27
Page 28
THE NAUTILUS, Vol. 103, No. 1
Figure 4. Scanning electron micrographs of the hinge of dis-
articulated valves of Teredo navalis post-larvae seen in figure
3. Numbers indicate shell height (greatest dorsoventral dimen-
sion) in lim. LP, ligament pit.
in shells approximately 255 to 275 jxvn liigli, the ventral
condyle had developed into a prominent knob, and the
proviiiculurn was no longer recognizable (figures 3, 4).
Rapid growth along the ventral margin increased the
depth of the valves and caused articulated shells to be-
come nearly spherical. Protrusion of ventral condyles
further separated right and left valves. Five to 12 ridges
covered the external surface of the anterior slope (figure
5, height 265 275 nm). Demarcation of the posterior
slope began in valves 300 nm high (figures 3, 5). Posterior
margins of right and left valves were s\ mmetrical; un-
eijual secretion of the dissoconch along this margin re-
stored earlier as\mnietr\ of the \aKes (figure 8).
External grow th lines marked the pronounced ventral
extension of the valves (figure 5, height 330/295 iim).
.\s development continued, the posterior slope became
more prominent (figure 3, height 565 615 tim). The um-
boiial-vcntral ridge appeared on the interior surface of
the shell (figure 3, height 565, 615, 1,750 nm). The dorsal
condyle developed (figure 4, height 565/615 ^lm) from
the earlv expanded base of the apophysis (figure 4, height
240, 270/255, 300 320 //m) and was positioned posterior
to the apophysis (figure 9). ,At a shell height of approx-
imateK 1 mm, the posterior gape was well-developed
(figure 10).
The external surface of the anterior slope of both \al\es
was sculptured w ith closeK -spaced, denticulated ridges
in early post-larvae. Subsequent ridges were more widely-
spaced, so that the relationship of shell length to number
of ridges over the entire post-lar\al period w as logarith-
mic (figures 5, 11). In contrast to earl\ post-lar\al ridges
w ith uniform denticles, late post-larval ridges had fine
denticles in the anterior section, while large, coarse den-
ticles marked posterior ridges (figure 5, height 610 490.
855 /im). In addition, these late post-lar\al ridges formed
a sharp angle at the ventral junction of the anterior and
posterior sections of the anterior slope.
The earliest pallets were removed from animals with
a shell length of approximateK 240 /um. The pallet blade
at this stage was triangular, w ith the margin of the inner
face extending further distalK than that of the outer face
(figure 12, 75 fxm). The shape of the blade became more
rectangular with additional growth of the inner margin
(figure 12, 125/110 ^m). Inner and outer margins of the
pallet blade elongated, and the margin of the outer face
became considerably more concave than that of the inner
face (figure 12, 200/195, 285/265, 485/470 Mm).
Throughout development of the pallets, each pair had
a symmetrical shape. Close examination of an indi\idual
pallet, however, revealed that the ventral surface was
longer than the dorsal surface of the blade. The resulting
asymmetrv of the blade w as increasingK' e\ident as \en-
tral and dorsal surfaces rounded and the dorsal distal tip
extended further than the ventral distal tip (figure 12,
715/795, 1,025/1,035 nm). Further growth was greatest
in a distal direction, resulting in an elongate shape of
pallets approximateK 1,500 ^im long (figure 12),
Pallet length increased more rapidly than shell length
during initial post-larval stages, .\fter anterior and pos-
terior slopes were well-developed, pallet length and shell
length increased at approximately equal rates (figure 13).
Figure 5. Scanning electron micrographs of the exterior surface of disarticulated valves of Teredo navalis post-larvae. Numbers
above the shells are shell height (greatest dorsoventral dimension); numbers beside the shells are shell length (greatest anteroposterior
dimension). Dimensions are in m'" and are accurate to within 5 Mm .Arrows mark late post-larval ridges, which are comprised of
both fine and coarse denticles. .-XS, anterior slope; PS, posterior slope.
S. C. Fuller et al., 1989
Page 29
Page 30
THE NAUTILUS, Vol. 103, No. 1
Figures 6-8. Scanning electron nncrographs ot early post-larval shell specimens ot Tercdu navalis. 6. Dorsal view alter the shift
in axis of articulation to a dorsoventral orientation. Scale bar = .50 nm. 7. Posterior view after the shift in axis of articulation to a
dorsoventral orientation. Scale bar = .iO ^ni K. Posterior view when symmetry of right ami left posterior margins is nearK complete.
Scale bar = .50 /jm
Figures 9. 10. 9. Scaiming electron micrographs of a left valve ot a post-larval specimen of Teredo navalis. Stereo pair was taken
uith an S° angular displacement between exposures. 10. Scanning electron micrograph of an articulated post-larval shell specimen
of Teredo navalis: posterior view. Scale bar = 200 ftm.
DISCUSSION
Larval Development
Previously reported shell dimensions of Teredo navalis
larvae are summarized in table 2. Dimensions of straight-
hinge larvae in the present study were consistent with
those in previous descriptions. Mean provinculuni length
of T. navalis in the present study was 47.7 fim, close to
the 51.3 nm mean given by Culliney (1975) and within
the 45 to 50 nm range given by Chanley and Andrews
(1971). Umbones appeared when lar\ae were about 120
|tm long, similar to the 123.3 ^m mean length reported
by Culliney (1975) and within the 110 to 150 Mm length
range given by Sullivan (1948), but larger than the 95
to 100 ixm length range found by Chanley and .Vndrews
(1971). Shell length and shell height were equal when
larvae of this species were approximately 150 jum, which
was the size given by Imai et al. (1950) but was larger
than the 123 and 122 ^m sizes of equidimensional larvae
found b> Culline\ (1975) and Mann and Gallager (19S5),
respectively . Size at metamorphosis w as smaller than that
reported by Sullivan (1948), Imai et a/. (1950), and Mann
and Gallager (1985) and approximateK the same as that
reported by Loosanoff and Davis (1963), Loosanoff et al.
(1966), Chanley and Andrews (1971), and Culliney (1975).
Difficulty in achiex ing consistent shell orientation due to
great convexity of the valves during late larval stages
may account for the discrepancies among measurements.
Measurements of disarticulated valves, aligned with the
shell margin in a plane, would improve accuracy'.
Scheltema ( 1 97 1 ) summarized previous descriptions of
teredinid larvae from the North .Atlantic. He emphasized
that features included in these early descriptions, such
as larval size ranges and color, are not always sufficient
for positi\e identihcation of unknown species. On the
other hand, differences in external shell surface sculpture
revealed by scanning electron microscopy distinguished
S. C. Fuller et al., 1989
Page 31
2.0
E
£
1.0
r
O)
c
0)
Ti
0.5
xz
W
0.2 -
25
0 5 10 15 20
Number of ridges
Figure 1 1. Relationship between shell length (greatest antero-
posterior dimension) and number of (early and late post-larval)
ridges on the e.xterior surface of shells of Teredo navalis. Regres-
sion equation is y = -0.28 -I- O.OSx; r- = 0.93.
several species of the Teredinidae (Scheltema, 1971;
Turner & Boyle, 1975; Calloway & Turner, 1979). Also,
Turner and Johnson (1971) illustrated the distinct nature
of the velum in two teredinids and suggested use of this
character as an aid in species identification.
The value of hinge structures for identification of lar-
val bivalves is well-documented (Jorgensen, 1946; Sul-
livan, 1948; Rees, 1950; Loosanoff et al, 1966; Chanley
& Andrews, 1971; Le Pennec, 1980; Lutz et al. 1982;
Jablonski & Lutz, 1983). In teredinids, provinculum length
is constant throughout the larval period (Rees, 1950);
therefore, identification of teredinid species based on
provinculum length can be made without regard to larval
size or stage of development. In specimens of T. navalis
larvae examined in the present study, dimensions of
provincular teeth showed individual variation but \\ ere
independent of developmental stage or size.
Provinculum length of Teredora malleolus (Turton,
1822) (= Teredo thomsonii Tryon; Turner, 1966; Schel-
tema, 1971) is approximately 70 ^m (Rancurel, 1965: fig.
1) and of T. navalis averages 47.7 ^m (present study);
therefore, larval specimens of T. navalis and T. malleolus
are easily distinguishable on the basis of provinculum
length. Furthermore, measurements and shapes of in-
dividual teeth of T. malleolus reported by Rancurel (1965)
differ from shapes and dimensions of teeth of T. navalis
in the present study.
Scanning electron micrographs of a disarticulated right
valve of Lyrodus pedicellatus (Quatrefages, 1849),
another larviparous teredinid, were presented by Turner
and Johnson (1970: figs. 1, 2). Although measurements
of teeth from these micrographs are approximate, mea-
surements of the length of the central tooth relative to
the anterior and posterior teeth indicate distinct differ-
ences in the provinculum of the right valve of T. navalis
and L. pedicellatus.
Dimensions of pro\incular teeth of T. navalis also
differ from those of Bankia martensi (Stempell, 1899)
(table 1 ). The most striking difference in the provinculum
of these two species, however, is the shape of the anterior
tooth of the right valve, which is J-shaped in B. nmrtensi
and rectangular in T. navalis (B. Campos, personal com-
munication). A J-shaped anterior tooth is also noted in
the provinculum of the right valve of Bankia setacea
(Tryon, 1863) (Quayle, 1951: fig. 8).
Met.\morphosis
Lutz and Hidu (1979) suggested that development of the
ligament pit was among the earliest morphological
changes that occur in planktonic bivaKes during meta-
morphosis. Further evidence of the early formation of
the ligament pit is seen during metamorphosis of Teredo
navalis. In this species, the ligament pit formed ventral
to the hinge structure in post-larval shells approximately
230 ^m high. Secretion of the dissoconch and formation
of apophyses, dorsal and ventral condy les, and external
ridges occurred subsequently, when shipworms were ap-
proximately 240 ;um high.
Table 1. Lengths (distance across center of tooth and parallel to hinge line) in ^m of provincular teeth of Teredo navalis and
Bankia martensi.'
Teredo navalis
Position
Range x ± SD
Left valve (n = 9)
Anterior
8.8-10.2 9.6 ± 0.5
Posterior
7.7-8.6 8.2 ± 0.3
Right valve (n = 12)
Anterior
6.3-8.3 7.2 ± 0.6
Central
13.7-18.2 16.4 ± 1.1
Posterior
6.0-7.0 6.6 ± 0.3
Bankia martensi
Range
X ± SD
Left valve (n = 10)
8.5-12.8 9.1 ± 1.0
4.2-8.5 6.4 ± 1.0
Right \alve (n = 9)
4.2-6.4 5.2 ± 1.2
14.9-17.0 16.4 ± 0.8
4.2-8.5 5.9 ± 1.0
' Data on Bankia niartensi were provided b) B. Campos, Universidad de X'alparaiso. \"ina del Mar, Chile. Specimens were sampled
from X'alparaiso Ba) , Chile.
Page 32
THE NAUTILUS, Vol. 103, No. 1
Figure 12. Scanning I'Ifctron micriintaplis ol llic (lutt-r face of pallets of Teredo navalia post-larvae. Micrographs of siinilarh-
sized right and left pallets are not neces.sarily from the same individual. Numbers indicate distance between distal and pro.\imal
ends in nm. I, inner margin; O, outer margin; D, dorsal surface; V, ventral surface.
Ill T. navalis, major modification.s of tlie shell enable
post-larvae to penetrate wood. Preparation for these
changes begins in mid-larval stages with increased growth
along the ventral shell margin, so that the new axis of
articulation is oriented along the longest shell dimension.
The shift in axis of articulation from the dorsal to the
posterior plane is accomplished by shell loss along the
posterior margin of the right valve. Quayle (1959) de-
scribed similar shell loss along the posterior margin of
the right \aKe of Bankia setacca following settlement
and suggested that the loss was due to shell erosion. The
shift occurs rapidK' in T. navalis; shells approximately
2.'30 nm high articulated along an anteroposterior axis,
and shells approximateK 240 jum high articulated along
a dorsoventral axis. Subsequent adjustments for this rear-
rangement, however, occur more gradually with growth
S. C. Fuller et ai, 1989
Page 33
of the animal. These include further separation of the
valves by an inward protrusion of the condyles until shells
articulate against the cond\ les rather than against dorsal
and \entral shell margins .Also, depth of the shell \alves
increases until articulated shells are nearly spherical. Sep-
aration of the valves allows extension of the foot and
siphons.
The sudden shift in the axis of articulation is followed
by an abrupt change in the orientation of growth axes.
Growth axes of the anterior section of the anterior slope
are roughly similar to larval growth axes. Growth at the
extreme ventral margin of the dissoconch, however, is
roughly perpendicular to the larval shell margin (figure
9).
Figure 13. Relationship between shell length (greatest antero-
posterior dimension) and pallet length (distance between prox-
imal and distal ends). Each point represents an individual an-
imal; when right and left values differed, the average was used.
Regression equation is y = 0.23 + 0.84x; r- = 0.99.
0.0 0.5 1.0 1.5 2.0
Pallet length (mm)
Table 2. Summar\ of literature reports on larval shell dimensions (in nm] of Teredo navalis at: (1) time of release from parent;
(2) appearance of umbones; (3) equidimensional stage; and (4) metamorphosis. For comparative purposes, 1 = anteroposterior
dimension; h = dorso\entral dimension; p = pro\ inculum length.
Equi-
Source
Release
Umbones
dimensional
Metamorphosis
Grave (1928)
1 = 85-90
h = 70-75
—
—
—
Jorgensen (1946)
1 = 80 (x)
—
—
h == 250
Sullivan (1948)
h X 1 « 80 X 95
1= 110-1.50
—
h X 1 =: 250 X 220
Imai et al (1950)
1 = 85 (x)
h = 72 (x)
—
*150
1 = 215 (x)
h = 235 (x)
Loosanoff and
h X 1 = 70 X 80 (min)
—
—
1 > 200
Davis (1963)
1 = 90 (max)
1 = 85-95 (average)
1= 190 (min)
h = 206 (min)
1 = 200 (max)
h = 231 (max)
Loosanoff et al.
h X 1 = 70 X 80 (min)
—
—
1 > 200
(1966)
1 = 90 (max)
Chanley and
1 = 70-90 (min)
1 = 95-100
130-140
1 = 190- > 200
Andrews (1971)
p = 45-50
Culliney (1975)
1 = 87.7 (x);
1 = 123.3 (x)
*123
'1 = 205.5;
range = 77-95
h = 123.3 (x)
range = 200-216
h = 75.4 (x);
depth = 90
h = 239.2;
range = 66-85
range = 192-260
depth
range = 55-57
p = 51.3 (x);
range = 46-54
depth = 190
Mann and
1 = 89.3 (x)
—
*122
21 = 212.1
Gallager (1985)
h = 76.1 (x)
h = 230.0
Present study
1 = 81.5 (x);
range = 77-87
h = 66.6 (x);
range = 66-70
1 = 120
= 150
1 = 202.0 (x);
range = 195-210
' Pediveliger
- First appearance
of pedi\eliger.
Page 34
THE NAUTILUS, Vol. 103, No. 1
External Shell Morphology
Numerous, closely-spaced denticulated ridges cover the
external surface of the anterior slope of earK post-lar\al
shells of Teredo iiavalis. .\lthougii the number of ridges
found in early stages is variable, observations show a
consistent decrease in total number of ridges after for-
mation of the posterior slope (compare figure 5, right
\al\es, heights 275 and 295 ^m) This decrease suggests
that eari\ post-larval ridges are obliterated as late post-
larval ridges (ridges with both fine and coarse denticles)
are added at the anterior shell margin.
Hill and Kofoid (1927) described the function of ex-
terior ridges of the adult shell of T. navalis. B\ carving
in an anterior direction, shipworms deepen burrows with
the fine serrations on anterior ridges; at the same time,
posterior ridges with coarser teeth increase the diameter
of the burrow. SlightK cur\ed earK ridges have uni-
torinK sized denticles (figure 5, height 330 295 i^m). As
described b\ Sigerfoos (1908) for Bankia gouldi (Bartsch,
1908), a difference in spacing at the dorsal and ventral
ends of the shell of T. navalis gradualK increases the
curvature of a single ridge until a sharp right angle forms
at the junction of the anterior and posterior sections of
the anterior slope. In T. navalis, fine denticles form in
the central portion of the ridge at this stage (figure 5,
height 610 490 ^l^\). Successive ridges are comprised of
only fine denticles on the anterior section and only coarse
denticles on the posterior section of the anterior slope
(figure 5, height 855 ^m).
P.m.let Development
Pallet morphology during early post-larval develop-
iiu'ntal stages was one of the characters used in deter-
mining phylogenetic relationships of species within the
family Teredinidae (Turner, 1966). Unfortunately, de-
velopment of pallets is described for only a few species.
Pallets of Lijrodus pedicellatus were described b\ Isham
and Tierney (1953) from first appearance until an esti-
mated length of 80 ^"1- Rancurel (1955) described de-
velopmental stages of pallets approximateh 400 fim long
and larger for Teredora malleolus. Early pallet blades
of both Teredo navalis in the present study and of L.
pedicellatus described b) Isham and Tiernev (1953) are
initially triangular in shape and then grow distally. Early
pallets differ in the tw o species, however, in that the stalk
of the pallets in L. pedicellatus is thin and transparent,
w hile the stalk of the pallets in T. navalis is calcified and
opaque. Also, in L. pedicellatus the stalk forms before
the blade (Isham & Tierney, 1953: fig. 20), whereas the
earliest pallets examined of 7". navalis had a blade (figure
12, 75 ^im). In T. malleolus, the blade of pallets nearly
600 jum long is circular in cross-section and, with sub-
sequent elongation, develops a medial fissure (Rancurel,
1955: fig. 4; Turner, 1966). In T. navalis, the blade of
pallets approximately 700 ^m long is circular in cross-
section and then becomes elongate, but development of
a fissure does not occur (figure 12).
The degree of individual and environmental variation
in pallet morphology of adult T. navalis was demon-
strated by Miller (1923). Turner (1966) stated that two
pallets removed from an individual shipworm often
showed unequal erosion, breakage, and deformation.
Pallets in the present stud\ were less subjected to effects
of an adver.se environment because of controlled culture
conditions and collection at an earK age. In general,
sampled pallets had similar morphology, and sequential
developmental stages were consistent. .\ striking s\m-
metr\ of opposing pallets was observed throughout the
post-larval developmental period (figure 12).
ACKNOWLEDGEMENTS
We gratefully acknowledge the help and advice of Dr.
R. D. Turner throughout this stud> and thank her es-
pecially for species verification of parent shipw orms and
for critical review oi the manuscript. We thank R Cam-
pos for her insightiul suggestions and comments on the
manuscript and for generously sharing her data on Ban-
kia martensi. Special thanks are extended to Dr. A. Poo-
ley for valuable discussions on shell growth patterns and
for technical advice on stereomicroscopv . D. Christensen
of the U.S. Department of Agriculture Forest Service
identified the wood specimen from which adult ship-
worms were collected. This is New Jersev Agricultural
Experiment Station Publication No. D-32401-1-89. sup-
ported by New Jersey State funds, NSF Grant EAR-84-
17011, and various NOAA Sea Grants to Rutgers Uni-
versity.
LITERATURE CITED
Abbott, R. T. 1974. American seashells, 2nd ed. Van Nostrand
Reinhold Companv . New York, 663 p.
Callovvav. C B. and R D. Turner. 1979. New techniques for
preparing slieils of bi\ alve larvae for exaiiiiiiatioii with the
scanning electron microscope. Bulletin of the .\nierican
Malacological L'nion for 1978, p. 17-24,
Chanley, P. and J. D. .\ndrevvs. 1971. .Aids for identification
of bivalve larvae of Virginia. Malacologia 11:45-119.
Culliney, J. L. 1975. Comparative larval development of the
shipworms Bankia gouldi and Teredo navalis. Marine Bi-
ology 29:245-251.
Fuller, S. C, R. A. Lutz, and A. Pooley. 1989. Procedures for
accurate documentation of shapes and dimensions of larval
bivalve shells with scanning electron micro.scopv. Trans-
actions of the .\merican Microscopical Societv 108:58-63.
Grave, B. H. 1928. Natural history of the shipworm. Teredo
navalis. at Woods Hole, Massachusetts. Biological Bulletin
55:260-282.
Hill, C. L. and C. A. Kofoid (eds.). 1927. Marine borers and
their relation to marine construction on the Pacific Coast.
Final report ot the San Francisco Bav Marine Piling C"om-
mittee I niversitv of California Press, San Franci.sco, .3.57 p
Imai, T., M. Hatanaka, and R Sato. 1950 Breeding of marine
timber-borer. Teredo navalis L . in tanks and its use for
anti-boring test. Tohoku journal of .Agricultural Research
1:199-208.
S. C. Fuller et a/., 1989
Page 35
Isham, L. B. and J, Q. Tierney. 1953. Some aspects of the
larval development and metamorphosis of Teredo (Lij-
rodus) pedicellaUt De Qiiatrefages. Bulletin ot Marine Sci-
ence of the Gult and (Caribbean 2:574-589.
Jablonski, D and R. .V. Lutz. 1983. Larval ecology of marine
benthic invertebrates: paleobiological implications. Bio-
logical Reviews ot the C'ambridge Philosophical Society
58:21-89.
Jergensen, C. B, 1946. Reproduction and larval development
of Danish marine bottom invertebrates, with special ref-
erence to the planktonic larvae in the Sound (0resund),
Lamellibranchia. Meddelelser fra Kommissionen for Dan-
marks Fiskeri-og Havunders0gelser, Serie: Plankton 4:277-
311.
Le Pennec, M. 1980. The larval and post-larval hinge of some
families of bi\alve molluscs. Journal of the Marine Bio-
logical Association of the United Kingdom 60:601-617.
Loosanoff, \". L. and H. C. Davis. 1963. Rearing of bivalve
mollusks. Advances in Marine Biology 1:1-136.
Loosanoff, V. L., H. C. Davis, and P. E. Chanley. 1966. Di-
mensions and shapes of larvae of some marine bivalve
mollusks. Malacologia 4:351-435.
Lutz, R., J. Goodsell, M. Castagna, S. Chapman, C. Newell, H.
Hidu, R. Mann, D. Jablonski, \". Kenned), S Siddall, R
Goldberg, H. Beattie, C. Falmagne, A. Chestnut, and A.
Partridge. 1982. Preliminary observations on the use-
fulness of hinge structures for identification of bivalve
larvae. Journal of Shellfish Research 2:65-70.
Lutz, R. A. and H. Hidu. 1979. Hinge morphogenesis in the
shells of larval and earl\ post-larval mussels {Mytilus edulis
L. and Modiolus modiolus (L.)). Journal of the Marine
Biological .Association of the L'nited Kingdom 59:111-121.
Mann, R. and S. M. Gallager. 1985. Physiological and bio-
chemical energetics of larvae of Teredo navalis L. and
Bankia gouldi (Bartsch) (Bivalvia: Teredinidae). Journal
of E.xperimental Marine Biology and Ecology 85:211-228.
Miller, R. C. 1923. Variations in the pallets of Teredo navalis
in San Francisco Bay. University of California Publications
in Zoology 22:401-414.
Quayle, D. B. 1951. The larvae of Bankia setacea Tryon.
Report, British Columbia Department of Fisheries (1951):
88-91.
Quayle, D. B. 1959. The earl\ de\elopment of Baiifaasc/acea
Tryon. In: Ra\', D. L. (ed.). Marine boring and fouling
organisms. University of Washington Press, Seattle, p. 157-
174.
Rancurel, P. 1955. Teredo thomsoni Tryon et Teredo lie-
herkindi Roch: transformations morphologiques des pa-
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fais d'.'Xfrique Noire 17:1149-1156.
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mellibranch larvae. Hull Bulletins of Marine Ecology 3:
73-104.
Scheltema, R. S. 1971. Dispersal of phytoplanktotrophic ship-
worm larvae (Bivalvia: Teredinidae) over long distance by
ocean currents. Marine Biology 11:5-11.
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Bulletin of the Fisheries Research Board of Canada 77:1-
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Turner, R. D. 1966. A surve\' and illustrated catalogue of the
Teredinidae (.MoUusca: Bivalvia). The Museum of Com-
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Turner, R. D. 1971. 1. Identification of marine wood-boring
molluscs In: Jones. E. B. G and S. K. Eltringham (eds.).
Marine borers, fungi and fouling organisms of wood. Or-
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Turner, R. D. and P. J. Boyle. 1975. Studies of bivalve larvae
using the scanning electron microscope and critical point
drying. Bulletin of the American Malacological Union for
1974, p. 59-65.
Turner, R. D. and A. C. Johnson 1970. Some problems and
techniques in rearing bivalve larvae. Annual Reports for
1969 of The .American Malacological Union, p. 9-13.
Turner, R. D. and .\. C. Johnson. 1971. 13. Biology of marine
wood-boring molluscs. /)!: Jones, E. B. G. and S. K. El-
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THE NAUTILUS 103(1 ):36-39, 1989
Page 36
Shell Tubules in Corbicida fliiminea (Bivalvia: Heterodonta):
Functional Morphology and Microstructure
Anionieto Tan Tiu
Harbor Branch Oceanographic
Institution
5600 Old Dixie Highway
Fort Pierce, FL 34946, USA
Kobert S. PrezanI
Biolog) Department
Indiana University of Pennsylvania
Indiana, PA 15705-1090, USA
INTRODUCTION
Narrow tubules are relatively common structures that
permeate bivalve mollusc shells (Oberling, 1964; Koba-
yashi, 1969; Taylor et ai, 1969; Omori et al, 1976; Rob-
ertson & Coney, 1979; Waller, 1980; Prezant, 1981; \Va-
tabe, 1988). Although well docimiented, the\ remain
poorly understood. .According to Waller (1980) these tu-
bules are more common in epifaunal than infaunal bi-
valves. There have been several hypotheses concerning
the function of these tubules. Waller (1980) summarized
these in a publication on the shell and mantle of arcoid
bivalves. Briefly, these include; w indows for photorecep-
tion (allowing transduction of light to underlying pallial
photoreceptors); anchorage sites for the mantle; aque-
ducts involved with, and as a means of, expanding surface
area for respiratory exchange; protection against boring
organisms; and a site for mobilization of calcium and
carbonate ions to buffer extrapallial fluid during acidi-
fication events as might occur in long periods on adduc-
tion. Additionally, such tubules could be involved with
embryonic nourishment (Rosso, 1954) or monitoring water
conditions (Robertson and Coney, 1979).
Curbicula fluminca (Miiller, 1774), unlike the corbic-
ulids Polymcsoda carolin iana (Bosc, 1901 ) (Tan Tiu, 1987)
and P. (Geloina) erosa (Solander, 1786) (Prezant & Tan
Tiu, 1986), possesses numerous shell tubules. These tu-
bules, which are filled by mantle extensions, could be
involved in lightening juvenile shells thereby decreasing
the drift load during "planktonic dispersal, and help in
anchoring the mantle to the shell during locomotion or
biomineralization.
MATERIALS AND METHODS
Shell microstructure of 30 specimens drawn randomly
from some 250 specimens of Curbicula fluminea col-
lected at different seasons and from various habitats in
southern Mississippi and ranging in size from 7 to 38
mm shell length, were closely scrutinized for possession
of shell tubules. Exact details of collection times and sites
can be found in Prezant and Tan Tiu (1986) and Tan
Tiu (1987). Whole or fractured valves were critical point
dried in a Denton DCP- 1 Critical Point Drier using litiuid
carbon dioxide as a transfer agent from absolute ethanol,
either without prior treatment or following treatment in
50% (v/v) hydrochloric acid or 50% (v/v) commercial
Clorox (sodium hypochlorite). Specimens were coated
with a thin layer of gold in a Polaron SEM Coating Unit
E5100, and examined at accelerating voltages of 30 kV
in an AMR 1000 scanning electron microscope.
RESLLTS
Narrow tubules, circular in cross-section, are a connnon
feature of the early dissoconch shell of Corbicula flu-
minea. The extent of shell that has tubules covers a
concentric band about 2 mm w ide parallel to the curva-
ture of the shell margin. The proximal border is about
700 nm from the hinge. Tubules averaging 6 ^m in
diameter consistently permeate the early dissoconch in
this band, but not the prodissoconch nor the later dis-
soconch shell ill all specimens examined.
Straight or sinuous tubules penetrate "well forined"
cross-lamellar shell microstructures (.figures 1, 2) and are
usually oriented roughly perpendicular to the shell sur-
face (figure 3) up to the \icinity of the periostracum.
Here they are infrequently deflected to become roughly-
parallel to the inner surface of the periostracum. This
occasionally is revealed in specimens treated with acid.
In these the surface shell is eroded and remnant surface
shell has rarely show n a single mantle extension exiting
one tubule in cross-section and entering another along a
plane parallel to the shell surface. Internally, tubules are
filled by extensions of the mantle (figure 4), The leading
end of this finger-like mantle projection is smooth (figure
5). These tissues occasionally show a bifurcating or anas-
tomosing tip (figures 5, 6) toward the shell exterior.
The relatively uniform distribution of mantle exten-
sions into tubules is illustrated in figure 7. This micro-
graph shows the basal regions of mantle extensions on
the shell-facing surface of a reflected portion of the man-
tle. Disregarding shell tubule branchings and anasto-
moses, and a.ssimiing that the tubules were perfect cy 1-
inders, a conservative estimate of \olume occupied by
A. Tan Tiu and R. S. Prezant, 1989
Page 37
Figure 1. Shell fracture showing Dared end of tubule on inner surlace ,li>p ul pluilu; ut shell, and normal cms.sL J-lamellar
microstructure of tubular wall. Horizontal field width (HFW) = 192 ^m. Figure 2. Shell fracture showing normal crossed-lamellar
microstructure of tubular wall (inner surface of shell towards lower right) HFW = 16 nm. Figure 3. Shell fracture showing
tubules traversing the entire calcareous shell component. Shell tubules are oriented roughh perpendicular to the shell surface (inner
surface of shell towards upper right). Portions of periostracum are visible on lower left of photo. HFW = 332 Aim. Figure 4. Shell
fracture showing mantle extension in a tubule (inner surface of shell towards upper left). HFW = 89 fim. Figure 5. External
surface of shell [periostracum and thin layer of shell digested by 50?( (v v) hydrochloric acid for 30 seconds] showing leading edge
of mantle extension. HFW = 16 tim. Figure 6. External surface of shell [periostracum and thin layer of shell digested by 505*
(v/v) hydrochloric acid for 30 seconds] showing bifurcation of tubular extension. HFW = 228 nm.
Page 38
THE NAUTILUS, Vol. 103, No. 1
tubules relative to total volume of shell possessing tubules
ranged from 0.8 to 2.1'^. Figure 8 shows the inner shell
surface with an estimated densit> ot 19() tubule openings
per square millimeter, which is about lour times greater
than the densit\ ol possible tubule openings in the ex-
ternal shell surface (figure 9).
DISCUSSION
We believe this is the first recorded observation of the
distal tips of mantle extensions associated with shell tu-
bule formation. The mantle extensions in Corbicula flu-
minea do not penetrate the periostracum, as is true for
the pisidiids (Robertson & Cone\', 1979) and arcoids
(Waller, 1980). The tubules can detour awa\ from the
periostracum and anastomose lateralK , as viewed from
the shell exterior in a partially decalcified shell. Unlike
shell tubules in arcoids (Ober'ling, 1964; Waller, 1980)
and pisidiids (.Robertson & Cone\ , 1979), which are dis-
tributed throughout the shell medial to the pallial line,
shell tubules in C. jiuminea are restricted to the early
dissoconch. It is uncertain whether the dissolution occurs
only proximalK in early shell deposition or distalK and
radialK along the base and length of the pallial extension.
In fact, without cyto- or histochemical analyses, we can
only speculate that the mantle extensions filling the tu-
bules are responsible for penetration of the shell and
likel)' do so through chemical means. It is possible that
an acid, as found in the mantle of Lithophaga Roding,
1798 (Morton, 1983), or a proteolytic enzyme similar to
those produced by the accessor)' boring organs of muricid
gastropods (Carriker & Williams, 1978), is in\oKed in
shell tubule formation by dissolution. It is also possible,
though unlike!)', that shell biomineralization occurs
around an inhibitory mantle extension.
We propose that supplementar\ anchorage of mantle
to shell is the primar\ I unction of the tubular tissues in
Corbicula fluminea. This is suggested by the orientation
of the mantle extensions that branch or anastomose to-
ward the shell exterior, thus forming a root or anchor
within the shell. During earK stages of juvenile devel-
opment, including small mature clams (7-14 mm shell
length), C. jiuminea is able to disperse through the water
column using ctenidial-produced mucous drouge lines
(Prezant & (>halermwat, 1985). During this drifting pe-
riod the valves remain w ideK abducted. This, as well as
the usualK' active behavior of benthic juxeniles (Krae-
mer, 1986), could place strain on the mantle-shell junc-
ture. Mantle extensions could aid in maintaining this
connection. Moreover, the decrease in shell \olume di-
rectly attributed to the shell tubules could offer the bi-
valve a slightly lighter shell with less mineral content.
As drifting occurs in smaller clams and the tubules are
onK found in the early dissoconch shells of C. fluminea.
there is a possible correlation between drifting and [)os-
session of lighter shell. Beyond the drifting stage, the
tubules possibly become a liability rather than an a.sset
as suggested b\ the differential erosion of the external
shell surface at the umbonal region, as observed b\ Kat
Figure 7. Mantle reflected to show uniform distribution of
mantle e.\teiisions on .surface facing shell tubules (upper right).
HFVV = 707 tint. Figure 8. Opening of tubules on inner
surface of shell (estimated densil\ = 196 tubules per square
millimeter), I mbci towards the top, HFV\' = 453 ^m. Figure
9. Densit) of tubule openings on external surface of untreated
shell is four times lower (estimated density = 45 tubules per
square millimeter) than nn inner surface (see figure 8). HFW
= (itiO ^ini
A. Tan Tiu and R. S. Prezant, 1989
Page 39
(1982), in clams inhabiting acidic waters. The mantle
extensions anchored to the wall \ ia shell tubules could
facilitate biomineralization b\ bringing tiie mantle in
close apposition to the deposition site, i.e., shell. This
could be particularK important in the umbonal region
as, under certain conditions, this is the thinnest part of
the shell (i.e.. in those with eroded umbos). It is also
possible that whenever the mantle e.\tensions of C. flu-
minea are exposed, due to extreme external erosion of
shell, the mantle extensions secrete substances to the outer
surface of the shell, preventing further shell erosion {Fritz
et at.. 1988). Some shell tubules that penetrated the peri-
ostracum (figure 9) and as noted by Fritz et al. (1988)
are likely preservation and erosion artifacts, respectively.
At present, all proposed functions of shell tubule and
associated mantle extensions remain speculative.
ACKNOWLEDGEMENTS
We gratefully thank Dr. M. G. Harasewych and two
anonymous reviewers for their suggestions on the pre-
vious versions of this research note. We also acknowledge
the Department of Biological Sciences, University of
Southern Mississippi for financial aid and use of research
facilities, and Mr. Tom Smoyer for darkroom assistance
(printing negatives). Publication cost was supported by
Harbor Branch Oceanographic Institution, Inc. (HBOI
contribution no. 675).
LITERATURE CITED
Carriker, M. R. and L. G. Williams. 1978. Preliminary char-
acterization of the secretion of the accessory boring organ
of the shell-penetrating muricid gastropod Urosalpinx ci-
nerea. Malacologia 17(1):125-142.
Fritz, VV. L.. L. M. Ragone, and R. A Lutz. 1988. Pores in
the shells of Corbicula fiuminea. National Shellfisheries
Association Meeting Abstracts, p. 208.
Kat, P. W. 1982. Shell dissolution as a significant cause of
mortality for Corbicula fluminea (Bivalvia; Corbiculidae)
inhabiting acidic waters. Malacological Review 15:129-
134.
Kobayashi, I. 1969. Internal microstructure of the shell of
bivalve molluscs. American Zoologist 9:663-672.
Kraemer, L. R. 1986. Biological basis of behavior in Corbicu/a
fluminea. I. Functional morphology of some trophic ac-
tivities. In: Britton, J. C. (ed). Proceedings of the Second
International Corbicula Symposium. American Malaco-
logical Bulletin, Special Edition No. 2(1986):193-201.
Morton, B 1983. Coral-associated bivalves of the Indo-Pacific.
In: Wilbur, K. (ed.). The molluscs. Vol. 6. .Academic Press,
New York, p. 139-224.
Oberling, J. J. 1964. Observations of some structural features
of the pelecypod shell. Mitteilungen der Naturforschem-
den Gesellschaft in Bern 20: 1-63.
Omori, M., I. Koba\ashi, M. Shibata, K. Mano, and H. Kamiya.
1976. On some problems concerning calification and fos-
silization of taxodont bivalves. In: Watabe, N. and K. Wil-
bur (eds. ). The Mechanisms of Mineralization in the In-
vertebrates and Plants. University of South Carolina Press,
Columbia, SC, p. 403-426,
Prezant, R. S. 1981. Comparative shell ultrastructure of l>on-
sid bivalves. Veliger 23(4):289-299.
Prezant, R. S. and K. Chalermwat. 1985. Flotation of the
bivalve Corbicula fluminea as means of dispersal. Science
225(4669): 1491 -1493.
Prezant, R. S. and A. Tan Tiu. 1986. Spiral crossed-lamellar
shell growth in the bivalvia Corbicula fluminea (Mollusca:
Bivalvia). Transactions of the .American Microscopical So-
ciety 105(4):338-347.
Robertson, j. L. and C. C. Coney. 1979. Punctal canal in the
shell of Musculium securis (Bivalvia: Pisidiidae). Mala-
cological Review 12:37-40.
Rosso, S. W. 1954. A study of the shell structure and mantle
epithelium of Musculium transversum (Say). Journal of
the Washington .Academy of Sciences 44:329-332.
Tan Tiu, A. 1987. Influence of environment on shell micro-
structure of Corbicula fluminea and Pohjmesoda carolin-
ana (Bivalvia: Heterodonta). Ph.D. dissertation. University
of Southern Mississippi, Hattiesburg, MS, 148 p.
Taylor, J. D., W. j. Kennedy, and A. Hall. 1969. The shell
structure and mineralogy of the Bivalvia. Introduction,
Nuculacea-Trigonacea. Bulletin of the British Museum of
Natural History, Zoology, Supplement 3:1-125.
Waller, T. A. 1980. Scanning electron microscopy of shell
and mantle in the order Arcoida (Mollusca: Bivalvia).
Smithsonian Contribution to Zoology Number 313. Smith-
sonian Institution Press, Washington, DC, 58 p.
Watabe, M. 1988. Shell structure. In: Wilbur, K. (ed.). The
Mollusca, Vol. 1 1 ; form and function. Academic Press, New
York, p. 69-104.
THE NAUTILUS 10.3(1):40-41, 1989
Page 40
Reoccurrence of Cyclonaias tuberculata in the
Huron River, Michigan
Kllen Scavia
Mark Mitchell
Huron River Watershed Council
-415 West Washington Street
Ann Arbor. \Ii 4S103
In 1935, mussels virtually paved the bottom of shoal-like
niches of the Huron River in southeastern Michigan.
Characteristic assemblages existed in the small, medium,
and large-river zones, as well as in creeks and river-lakes
(Van der Schalie, 1970). Since then, human activities
have brought about serious depletion among the 25 species
of mussels that occupied long stretches of the Huron.
One of those species, Cyclonaias tuberculata (Rafines-
que, 1820), was abundant between the cities of Dexter
and Ann Arbor in the 1930's but was designated as a
rare and endangered species by Michigan's 1975 Natural
Features In\entory.
Water quality in this river segment was improved in
recent years (Gannon & Meier, 1986) and currently sup-
ports a top quality, warm water fishery. In this note we
report findings of a surve\ conducted between .'^nn .Arbor
and De.xter during the summer of 1986 to determine if
Cyclonaias tuberculata had reestablished itself in this
portion of the Huron River.
The study area is a 16-km reach, between river miles
60 and 70. In this river segment, width ranges between
12 and 45 m and depth between 0.9 and 9.0 m (Sa\- &
Jansson, 1976), except for low flow conditions when depths
of 0.15 m were recorded. The 24-year annual mean dis-
charge ranges between 3.96 m'/sec and 16.7 m'/sec.
Six sampling locations were identified as having suit-
able substrate for Cyclonaias tuberculata. Preliminary
collecting established the presence of living or dead Cy-
clonaias at each of the sites. Where site conditions per-
mitted, 76 randomly sampled 1-m- plots were surveyed,
36 on each side of the river, six times between June and
August. Shells were hand-picked from the shallows, dig-
ging several inches below the surface. Dead valves de-
posited as middens by muskrats and raccoons were in-
cluded. The length of each shell was recorded, as well
as whether the specimen was live or dead. Living spec-
imens were returned to the river.
Of the 370 specimens of Cyclonaias tuberculata found,
23.5% were live animals. The three upriver stations \ ield-
ed 70% of the live specimens. Other genera found in this
reach of the river were Lampsilis, Villosa, Anodonta,
Elliptio, Ligumia, and Pisidium. The age of Cyclonaias,
approximated by tlie length of the shell, ranged from
about 3 to 20 years (2.6 to 11.0 cm); the average age for
both live animals and dead shells was approximately IS-
IS years (7.3 cm).
While the species has managed to recolonize this area
since its decimation some 50 years ago, its numbers re-
main well below prior population levels (Van der Schalie,
1970). A greater number of living specimens were lo-
cated in upriver sampling sites compared to downriver
sites; this ma\' correlate with impro\ed water qualit\ . In
recent surveys the water qualit) of those upper reaches
was rated as very good (Gannon & Meier, 1986). The
lower rated downriver area is impacted by Mill Creek,
the largest source of agricultural non-point pollution to
this river reach. During our study, plumes of silt and
suspended sediment were observed entering the river
from Mill Creek, possibly suffocating organisms. This
deleterious effect ma\ also be exacerbated by the effluent
of the Dexter Sewage Treatment Plant.
Theoretical life history data suggest that a healthy
population would contain greater numbers of young
valves relative to older valves than were found in this
survey. Three hypotheses ma\ explain the deviation be-
tween the size frequencN' distribution found in this sur\ey
(figure 1) and the expected theoretical distribution. (1)
Annual recruitment ma\ be low compared to 13 to 15
years ago. (2) (Greater selective mortalitv of voung, rel-
ative to older valves, may be occurring now w ith small
dead shells being washed to the nearest impoundment
where current is slowed. (3) Sample methodology used
may not be sensitive to the smallest mussels. Because
young mussels are approximateK 0.32 cm long at age 1
year (Neves, 1985), hand sampling is likely to miss this
group. In addition, the juvenile state may occup\- habitats
other than those of the adult that we sampled (D'Eliscu.
1972).
The latter hypotheses were tested in the summer of
1987 when the 1986 sample sites were revisited and
E. Sea via and M. Mitchell 1989
Page 41
40
35
(0
30-
<u
>
to
?5
>
o
20-
i-
(U
n
15-
h
D
Z
10
5-
0
2 4 6
Valve length (cm)
Figure 1. Size-frequenc> distribution of live and dead Cy-
clonaias tuberculata found in the Huron River study area.
sampled using a 2-mm mesh sieve. Three non-adult hab-
itat types were also sampled: macrophytes, loose substrate
downstream of boulders, and deep pool areas. Sampling
with sieves allowed a higher percentage of live adults
(53%) to be found compared to 23% in the 1986 survey.
However, only one early juvenile (less than 17 mm) and
four late juveniles (greater than 17 mm but less than 4
years old) were found in 450 0.25-m- samples. An abun-
dance of Sphaeriidae of the desired size category were
found, ruling out inefficient sampling as the problem but
raising the additional concern of species competition. It
has been postulated (Neves & Widlak, 1987) that the
juvenile stage of Unionidae is probably the most suscep-
tible stage to competitive interactions for food or space.
Because Sphaeriidae has a competitive advantage in terms
of adaptabilit\-, >oung Unionidae may suffer as Sphae-
riidae prosper.
This survey has determined that Cyclonaias tuber-
culata has been reestablished in the Huron River between
Ann Arbor and De.xter. However, the population size-
frequency distribution suggests there may be a reason
for additional concern. More information is needed to
assess if animal recruitment of Cyclonaias tuberculata
is sufficient to maintain a healthy mussel assemblage.
LITERATURE CITED
D'Elisou, P. N. 1972. Observation of the glochidium, meta-
morphosis, and juvenile of Andonta calijorniensis Lea,
1857. The Veliger 15(l):57-58,
Gannon, J. J. and P. G. Meier. 1986. Huron River water
quality study in vicinity of Ann Arbor, Michigan. Uni-
versitv of Michigan School of Public Health, Ann Arbor,
MI, 131 p.
Neves, R. J. 1985. Non-game update: endangered mussels.
Virginia Wildlife 46:30-31.
Neves, R. J and J. C. Widlak. 1987. Habitat ecology of ju-
venile freshwater mussels (Bi\alvia: Unionidae) in a head-
water stream in Virginia. .American Malacological Bulletin
5(l):l-7.
Say, E. W. and O. Jansson. 1976. The Huron River and its
watershed. Huron River Watershed Council, Ann Arbor,
MI, 34 p.
Van der Schalie, H. 1970 Mussels in the Huron River above
Ann Arbor in 1969. Sterkiana 39:17-22.
\'an der Schalie, H. 1975 .\n ecological approach to rare and
endangered species in the Great Lakes region. The Mich-
igan Academician 8:7-22.
THE NAUTILUS 103(1 ):42, 1989
Page 42
Occurrence of the Ribbed Mussel, Geukensia demissa, on the Book
Gills of a Horseshoe Crab, Limulus polyphemus^
Lewis E. Deaton-
Karen D. Kempler
Whitney Laborator\
University of Florida
9505 AlA South
St, Augustine, FL 32086
While collecting specimens of the flatworm, Bdelloura
Candida Girard, 1850, from the gills of the horseshoe crab,
Limulus polyphemus Linne, 1758, we observed a single
specimen of the .Atlantic ribbed mussel, Geukensia de-
missa granosissima Sowerby, 1914, attached by byssal
threads to a gill filament of one of the L. polyphemus.
This was the only occurrence of fouling by a mollusk in
well over 200 Limulus examined over a 2 day period.
The Limulus were collected in March of 1985 from the
Indian River in Florida near the \ olusia-Orange County
line.
It is of particular interest that the mussel was attached
to the gills of \.\\e Limidus since the latter species possesses
an appendage (the flabellum) specialized for cleaning
the gill filaments. Most, if not all, aquatic arthropods
groom the body surfaces, including the gills, extensively
(Bauer, 1981). The flabellae of the fouled Limulus were
fully functional. The mussel would not remain attached
to the Limulus beyond the next molt, as the byssal threads
did not penetrate the exoskeleton. The shell length of
the G. demissa, which probably settled on the horseshoe
crab as a larva, was 18 mm, suggesting that the animal
was about a year old (Lutz & Castagna, 1980). The Lim-
ulus had a carapace width of 15.5 cm.
Reports of fouling bivalves on arthropods are uncom-
mon. Wolff (1959) found 12 Mytilus edulis Linne, 1758
on a green crab, Carcinus maenas Linne, 1758, as well
' This is contribution number 282 from the Tallahassee, Sop-
choppy and Gulf Coast Marine Biological .Association.
- Department of Biologs , University of Southwestern Loui-
siana, Lafayette, L\ 70504.
as several 5-6 year old Crassostrea virginica on an in-
dividual of another crab species, Hyas araneus Linne,
1758. The age of the fouling oysters on this H. araneus
is probably explained by the fact that this species, like
most spider crabs, molts infrequently once mature (Hart-
noil, 1963). Marine turtles harbor a variety of epifaunal
mollusks, including both gastropod and bivalve species
(Frazier et al., 1985). The gastropod genus Crepidula
has been reported to occur on Limulus (Hoagland, 1984),
and while we found no Crepidula jornicata Linne, 1767
on any of the Limulus in the sample described here,
these gastropods are not uncommon on the ventral sur-
face of Limulus from this collection site.
LITERATURE CITED
Bauer, R. T. 1981. Grooming behavior and morphology in
the decapod Crustacea Journal of Crustacean Biologv 12:
153-173.
Frazier, J., D. Margantoulis. K Muldoon, C. U' Potter. J.
Rosewater, C. Ruckdeschel, and S. Salas. 1985. Epizoan
communities on marine turtles. I. Bivalve and gastropod
mollusks. Marine Ecology 6:127-140.
Hartnoll, R. G 1963. The biology of manx spider crabs Pro-
ceedings of the Zoological Society of London 1 4 1 423-496.
Hoagland, K. E. 1984. .Aerial exposure in the genus Crepidula
(Gastropoda: Prosobranchia) with comparisons to other taxa.
.American Malacological Bulletin 3:33-40.
Lutz, R. .A. and M. Castagna. 1980. .Age composition and
growth rate of a mussel (Geukensia demissa) population
in a Virginia salt marsh. Journal of Molluscan Studies 46:
106-115.
Wolff, T. 1959. Epifauna on certain decapod Crustacea, Pro-
ceedings of the .W International Congress of Zoology, pp.
1060-1061.
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THE NAUTILUS
Volume 103, Number 2
September 29, 1989
ISSN 0028-1344
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Delaware Museum of
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The American Museum of Natural
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Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
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Division of Mollusks
National Museum of
Natural History
Smithsonian Institution
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Department of Mollusks
Museum of Comparative Zoology
Harvard University
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Department of Geology
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Los Angeles County Museum of
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% Department of Mollusks
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THEt^NAUTILUS
CONTENTS
Volume 103, Number 2
September 29, 1989
ISSN 0028-1344
David G. Reid
Systematic revision ot the Recent species of Peasiella
Nevill, 1885 (Gastropoda: Littorinidae), with notes on the
fossil species 43
A new species of Coluzea (Gastropoda: Turbinellidae)
from off southeastern Africa 70
Genetics and shell morphology of hard clams (genus
Mercenaria) from Laguna Madre, Texas 73
Variation in size demography of lotic populations of
Corbicula fluminea (Miiller) 78
The nomenclatural status and phylogenetic affinities of
Syrinx aruanus Linne, 1758 (Prosobranchia: Turbinellidae) 83
M. G. Harasewych
Robert T. Dillon, Jr.
John J. Manzi
Barry S. Payne
Andrew C. Miller
Paul D. Hartfield
Robert F. McMahon
M. G. Harasewych
Richard E. Petit
Marine Biological Laboratory
LIBRARY
OCT 1 6 1989
Woods Hole, Mass.
THE NAUTILUS 103(2):43-69, 1989
Page 43
Systematic Revision of the Recent Species of Peasiella
Nevill, 1885 (Gastropoda: Littorinidae),
with Notes on the Fossil Species
David C. Reid
Department of Zoology
British Museum (Natural History)
London SW7 5BD, UK
ABSTRACT
Peasiella is a genus of the Littorinidae, whose members have
minute, trochoidal shells, and are found on rocky shores in the
Indo-Pacific region. Six Recent species are recognized herein,
one of which is described as new. Shell characters are highly
variable, but usually adequate for identification, and provide
characters for a key. The reproductive anatomy, paraspermatic
nurse cell, egg capsule, head-foot, and radula are described.
Distribution maps are provided, and the species can be classified
as oceanic, continental, or intermediate, according to their dis-
tribution and habitat. Within the subfamily Littorininae the
phylogenetic relationships of Peasiella are uncertain. Fossils
are known from France and New^ Zealand, the earliest from
the Middle Eocene. Four fossil species are briefly described,
but the account is probably not complete, because fossils may
have been misclassified in other families.
INTRODUCTION
Living species of Peasiella are restricted to the Indo-
Pacific region. Their shells are minute and trochoidal,
quite unlike those of most other httorinids. Consequently,
this is one of the most poorly known genera in the family
Littorinidae. Nevill (1885) listed seven species, and Tryon
(1887), in the first and hitherto only monograph, included
eight species. In the present revision si.x Recent species
are recognized, one of which is described as new.
Because of the trochoidal shell and multispiral oper-
culum, it is not surprising that six of the 19 available
names for Recent species were first described as members
of the Trochidae (Gould, 1849; Adams, 1853; Reeve,
1862; Pease, 1868; Issel, 1869; Turton, 1932). Fossil species
have been placed in the non-littorinid genera Trochus,
Pseudonina, Tornus, and Xenophora (Deshayes, 1824;
Morgan, 1915; Cossmann & Peyrot, 1917-19). The lit-
torinid affinities of these shells were first noted by Dunker
(1861), who described a species as a Risella Gray, 1842
(= Bembicium Philippi, 1846; see Reid, 1988). This clas-
sification was followed by most later authors, and indeed
the name Peasiella was first proposed as a subgenus of
Risella by Nevill (1885). Because the operculum of Pea-
siella is multispiral, while that of Bembicium is pauci-
spiral, Kesteven (1903) used it as a full genus, and re-
moved it to the Modulidae, together with the littorinid
genus Echinella Swainson, 1840 (= Tectarius Valen-
ciennes, 1832; see Reid, 1989), which also has a multi-
spiral operculum. The only other authors to indicate a
relationship between Peasiella and Tectarius were Sou-
verbie and Montrouzier (1879), who described a Pea-
siella species as an Echinella. Peasiella became widely
accepted as a full genus through the work of Japanese
authors (e.g., Habe, 1956, 1964, 1984; Oyama & Take-
mura, 1961; Yamamoto & Habe, 1962). Nevertheless, the
superficial resemblances between shells of Peasiella and
Bembicium continued to suggest a close relationship, and
Rosewater (1970) considered that they should be placed
together in a separate subfamily. The status and rela-
tionships of all the genera of the Littorinidae have re-
cently been revised, using cladistic analysis of anatomical
characters, and it has been shown that while Bembicium
is a member of the Lacuninae, Peasiella belongs in the
relatively derived Littorininae (Reid, 1989). However,
the relationships of Peasiella with other littorinine genera
remain uncertain.
An exhaustive survey of the fossil species is beyond
the scope of this work, for the material is widely scattered
in museum collections, and much of it is probably mis-
classified in other families.
Peasiella species are often common, but are incon-
spicuous, so that their ecology is poorly known. The avail-
able information is summarized below. Museum collec-
tions do provide some information about habitat
preferences, and give a relatively complete picture of
geographical distribution.
MATERIALS AND METHODS
This account is based on the collections of the following
museums: British Museum (Natural History), London
(BMNH); Museum National d'Histoire Naturelle, Paris
(MNHNP); National Museum of Natural History, Smith-
sonian Institution, Washington, DC (USNM); Academy
of Natural Sciences of Philadelphia (ANSP); Museum of
Page 44
THE NAUTILUS, Vol. 103, No. 2
Comparative Zoology, Harvard University (MCZ); Los
Angeles County Museum of Natural History (LACM);
Natal Museum, Pietermaritzburg (NM); National Sci-
ence Museum, Tok\o (NSMT); Australian Museum, Syd-
ney (AMS); Western Australian Museum, Perth (WAM);
New Zealand Geological Survey, Lower Hutt (NZGS).
Material has been collected personally (DOR, specimens
in BMNH) in Australia, Hong Kong, and Hawaii. In
addition, type specimens have been borrowed from the
Zoological Survey of India, Calcutta (ZSI); Oxford Uni-
versity Museum (OUM); Museum fiir Naturkunde, East
Berlin (ZMB) and Museo C^ivico di Storia Naturale "Gia-
como Doria', Genoa (MGD). All type specimens have
been examined unless otherwise stated. Lectotypes have
only been designated from syntypic series when only a
part of the original series was made available for ex-
amination.
Shell height was measured parallel to the axis of coil-
ing, and shell diameter perpendicular to this axis. The
height/diameter ratio was calculated as an index of shell
shape. The number of whorls of the teleoconch was
counted from the sinusigera ridge terminating the pro-
toconch, and the number of protoconch whorls counted
as described by Reid (1988:94). The numbers of ribs and
grooves given in parentheses are the rare extremes of the
range.
The amount of preserved material available was not
large, but 74 specimens were dissected, as listed in pa-
rentheses among the locality records, with the sex indi-
cated. No specimens were found to contain parasitic
trematodes. Serial histological sections were cut of three
male and two female P. roepstorffiana, and one female
each of P. tantilla and P. lutulenta. These were stained
with either Masson's trichrome (MT) (Culling, 1963) or
by the alcian blue-periodic acid-Schiff (ABPAS) tech-
nique for the histochemical differentiation of mucins
(Mowry, 1956). Spermatozoa from the seminal vesicle
were examined by light microscopy in four P. roepstorf-
fiana, three P. lutulenta (in each case from whole ani-
mals fixed and stored in 10% formalin in seawater), one
each of P. tantilla and P. isseli, and three P. roepstorf-
fiana (in these three cases samples from living animals
were fixed in 1% formalin in seawater, immediately be-
fore examination).
Radulae of 31 specimens were examined, from three
to eight for each species. They were treated with boiling
50% potassium hyroxide solution for 15 minutes, rinsed,
cleaned ultrasonically for ten seconds, and mounted flat
for examination by scanning electron microscopy. Pho-
tographs were taken of top and side views, and from the
anterior end at 45° to the horizontal; this last was adopted
as the standard view for showing cusp shape.
All collections listed in the locality records were ex-
amined, and have been used to construct distribution
maps.
GENERAL DESCRIPTION
The following descriptions of shell and anatomy of Pea-
siella are brief. A general, comparative account of all
the genera of the Littorinidae has been given elsewhere
(Reid, 1989).
Shell and protoconch: Shells of Peasiella are small (less
than 6.6 mm diameter) and almost always umbilicate.
The shape is usualK trochoidal (conical, with a keeled
periphery), but can be rounded at the margin and de-
pressed. The surface is usually sculptured by major spiral
grooves and fine spiral microstriae (figure 16). Conspic-
uous features such as carinate spiral ribs and radial folds
show considerable \ariabilit\ within species. The folds
may produce undulations of the peripheral keel, or in
extreme cases cause it to appear crenulated (scalloped).
The high degree of intraspecific variation can be ascribed
in part to large-scale geographical variation (see system-
atic section). In addition, the shells show allometric growth
(the apical angle decreasing with size), and a small degree
of sexual dimorphism (females being larger and slightly
lower-spired than males). Nevertheless, all the six Recent
species can be distinguished by their shells. The arrange-
ment of ribs on the shell base is an especialK useful
character. Peasiella isseli and P. infracostata may both
occasionally show small periostracal bristles on the ribs
around the umbilicus (figure 15).
The shell color pattern is basically of spiral rows of
dark spots, most prominent at the suture and periphery,
which sometimes form oblique or even spiral lines. Again
there is much intraspecific variation on a geographical
scale, but color patterns are nevertheless useful for iden-
tification (figures 1-14).
All the species show a small (0.21-0.28 mm diameter),
sculptured protoconch, terminated by a sinusigera ridge,
indicating planktotrophic development. The protoconch
consists of 2.3 to 2.8 whorls, of which the last 1 to 1.5
are sculptured by four or five zigzag or almost straight
spiral cords (figures 17, 18). It is only rarely preserved
intact in adult shells.
Operculum: The operculum is round, corneous and
multispiral, with five to seven revolutions (figure 19).
This type is found elsewhere in the Littorinidae only in
Tectarius (Echininus) Clench & Abbott, 1942.
Head-fool: Pigmentation of the head-foot is relatively
pale, and frequentK' there are one to three black bands
across the base of the snout. There ma\- be one or two
longitudinal black lines on the tentacles (figures 31-50).
From the limited amount of preserved material avail-
able, it appears that the pigmentation pattern of the head
may prove to be a useful taxonomic character in this
genus.
There is no evidence of longitudinal division of the
foot in preserved specimens, but this requires confir-
mation in living animals.
Male reproductive system: Both the prostate gland and
the anterior \as deferens along the side of the head are
open grooves, but the penial vas deferens is superficially
closed {i.e.. the duct is not surrounded by muscle, but
an epithelial connection to the surface remains). The
penis is long and usually with a single mamilliform penial
D. G. Reid, 1989
Page 45
Figures 1-14. Shells of Peasielta species, showing color patterns. 1. P. tantilla. Pupukea, Oahu, Hawaiian Is (BMNH). 2. P.
conoidalis. New Caledonia (BMNH). 3. P. roepstorffiana, Orpheus I., Queensland (BMNH). 4, 5. P. roepstorffiana, Ping Chau,
Hong Kong (BMNH). 6. P. roepstorffiana, Fiji (BMNH). 7. P. lutulenta, Barrow I., Western Australia (BMNH). 8. P. lutulenta.
Hoi Sing Wan, Hong Kong (BMNH). 9. P. lutulenta, Rowes Bay, Townsville, Queensland (holotype, AMS C149052). 10. P.
infracostata, Japan (BMNH). 11. P. infracoslata, Karachi (BMNH). 12. P. infracostata. Cape Ferguson, Queensland (BMNH).
13. P. isseli, Suez (BMNH 82.8.7.316). 14. P. isse/i, Grand Bay, Mauritius (possible syntype of Risella isseli var. mauritiana Viader,
1951; BMNH 1989004).
gland halfway along its length. Histological examination
reveals an additional type of penial gland, a small patch
of simple, subepithelial glands (not forming a glandular
disk, see Reid, 1989), at the base of the mamiUiform
gland. These two glandular types can be distinguished
by their staining reactions. The secretion of the mam-
illiform gland stains red in MT and magenta in ABPAS,
whereas that of the simple gland is colorless or pale blue
Page 46
THE NAUTILUS, Vol. 103, No. 2
in MT and stains darker magenta in ABPAS. The secre-
tion passes out through the hollow, raised papilla of the
mamilliform gland, but in the simple gland the secretion
passes in intercellular extensions through the o\erl\ ing
epithelium to the surface. Goblet cells (staining blue in
ABPAS) are abundant in the penial filament (i.e., distal
to the mamilliform gland).
In general, penial shape is similar in all Peasiella species,
and so is not a useful ta.xonomic character (figures 31-
50). Peasiella roepstorffiana is unusual, because in parts
of its range the mamilliform gland is absent. Penial shape
is normally rather constant within littorinid species, and
it is possible that two species may be involved here, as
discussed in the systematic section.
Spermatozoa: As in all members of the Littorininae,
the spermatozoa in the seminal vesicle are dimorphic.
The euspermatozoa are filiform and 36 to 50 iim in
length. The paraspermatic nurse cells are irregularly
rounded or oval and 11 to 23 ^im in diameter. They are
packed with conspicuous round granules (small and in-
distinct in P. roepstorffiana) and a nucleus is sometimes
visible, but they lack rod-shaped inclusions (figures 51-
56). In most other littorinines the euspermatozoa are
attached in bunches to the nurse cells, to form spermato-
zeugmata, which may function to prevent premature
dispersal of sperm from the largely open male pallial
gonoduct (Reid, 1989). However, in the few Peasiella
that have been examined, the euspermatozoa are not
attached to the nurse cells and the two cell types are
simply packed together in the seminal vesicle. This con-
dition is also found in Melarhaphe Menke, 1828, which,
like Peasiella, has a superficially closed penial duct.
Whether the apparent alssence of spermatozeugmata is
primitive, or a secondary loss as a consequence of closure
of the penial duct, is unknown. The former is more likely,
because elsewhere in the Littorininae spermatozeugmata
can be found in genera with superficially closed penial
ducts (e.g., Cenchritis von Martens, 1900, some species
of Littoraria Griffith & Pidgeon, 1834). As discussed in
the SNSteniatic section, some specimens of P. roepstorf-
fiana lack nurse cells in the seminal vesicle. Neither
intraspecific nor intrageneric variation in the presence
of nurse cells has been reported in any other littorinids.
Female reproductive system: The general arrangement
of the pallial o\iduct is the same as in other oviparous
members of the Littorininae. with the egg groove passing
successively through opaque albumen gland (colorless in
MT, magenta in ABPAS), translucent albumen gland
(colorless in MT, blue or magenta-purple in .\BP.\S),
opaque capsule gland (red in MT, pale purple in .•\BP.\S),
translucent capsule gland (blue in MT, dark purple and
magenta in ABPAS), and small jelly gland (colorless in
MT, magenta in ABPAS). Differentiation of these glands
can sometimes be seen in the whole animal and in gross
serial sections; the opaque capsule gland is especially
prominent, visible as a chalky white or cream crescent
on the lateral side of the pallial oviduct (figures 57-63,
65, 66). The spiral route of the egg groove (figures 58,
66), with a backward loop of albumen gland followed
by a larger loop of capsule gland, is unique to the genus,
and is the same in all six species.
The histological structure of the glands of the pallial
oviduct is unusual, in that there is not a clear separation
between a non-glandular epithelial lining of the egg
groove and the subepithelial glandular follicles beneath
[as found in all other Littorininae except Melarhaphe
(Reid, 1989)]. The posterior part of the jelly gland and
all of the capsule gland clearly show subepithelial glan-
dular tissue, but the albumen gland appears to be largeK
of epithelial structure. ProbabK' the glandular cells are
both epithelial and subepithelial throughout the pallial
oviduct.
Figures 15-22. Sculptural details, protoconchs. operculum, and radula of Peasiella species. Scale bars 15-19 = 0.2 mm; 20-22 =
0.02 mm. 15. P. isseli, detail of umbilicus showing periostracal bristles, Oman (BMNH). 16. P. conoidalis, detail of sculpture on
last whorl, Anaa, Tuamotu Is (BMNH). 17. P. roepstorffiana. protoconch, Orpheus I., Queensland (BMNH). 18. P. conoidalis.
protocoiich, Oneroa, Tuamotu Is (USNM 720753). 19. P. roepstorffiana, operculum, Samar I., Philippines (NSMT 56716). 20-22.
P. tantilla, three views (flat, side and anterior from an angle of 45°) of a single radula, Coconut I,, Oahu, Hawaiian Is (BMNH).
Figures 23-30. Radulae of Peasiella species, 23, 24, and 27 are anterior views from an angle of 45°; others are vertical \iews of
flat radulae. Scale bars = ().()2 mm. 23. P. conoidalis, Dunidu I,, Male, Maldive Is (ANSP 305263). 24. P. conoidalis. Anaa, Tuamotu
Is (BMNH). 25. P. roepstorffiana. Magnetic I., Queensland (BMNH). 26. P. roepstorffiana. Ping Chau. Hong Kong (BMNH) 27.
P. lutulenta, Barrow I., Western Australia (WAM 2347-67). 28. P. isseli, Mahe, Seychelles (BMNH). 29. P. infracostata, Okinawa,
Ryukyu Is (AMS C146902). 30. P. infracostata, Cape Ferguson, Queensland (BMNH).
Figures 31-50. Heads and penes of Peasiella species. All or part of penial vas deferens and duct of mamilliform gland indicated
by dotted lines when visible by transparency .\ll drawn from fixed material, except 34 and 50, drawn live and relaxed in 7.5%
magnesium chloride solution 31 -.36. /' tantilla. 31, 32. Coconut I , Oahu, Hawaiian Is (BMNH) 33-35. Pupukea, Oahu (BMNH),
36. Kahuku, Oahu (AMS C144407) 37. 3B. P. conoidalis. Otepipi, Anaa, Tuamotu Is (BMNH) 39-42. /'. roepstorffiana. 39.
Ping Chau, Hong Kong (BMNH) 'W, 41. Picnic Bay, Magnetic 1 , Queensland (BMNH). 42. Marine Biological Center, Phuket I.,
Thailand (BMNH). 43-46. P. lutulenta. 43. Shark Point, Barrow 1., Western Australia (WAM 2347 67). 44, 45. Bowes Bay,
Townsville, Queensland (BMNH). 46. Sai Kung, Hong Kong (BMNH). 47, 48. P. infracostata. 47. Kuchino, Shiznoka Pref., Japan
(BMNH). 48. Hoi Sing Wan, Tolo Channel, Hong Kong (BMNH). 49, 50. P. isseli. 49. Bale Ternay, Mahe, Seychelles (BMNH).
50. Eilat, Israel (BMNH),
D. G. Reid, 1989
Page 47
mmm
mm
Page 48
THE NAUTILUS, Vol. 103, No. 2
D. G. Reid, 1989
Page 49
Page 50
THE NAUTILUS, Vol. 103, No. 2
52
53
20 pm
Figures 5 1 -56. Paraspermatic nurse cells of Peasiella species.
All drawn from living material fixed in 1% solution of formalin
in seawater. 51. P. tantilla, Pupukea, Oahu, Hawaiian Is. 52,
53. P roepstorjfiana. Picnic Bay, Magnetic I., Queensland. 54,
55. P. lutulenta, Sai Kung, Hong Kong. 56. P. isseli, Eilat,
Israel.
Another unusual feature is that in there species (P.
tantilla, P. conoidalis, P. roepstorjfiana) the bursa cop-
ulatri.x is rudimentary or absent. In the remaining three
species (P. infracostata, P. isseli, P. lutulenta) the bursa
is large and anterior in position (figure 65). Situated at
the back of the spiral part of the pallial oviduct, the
seminal receptacle is constricted into two parts, an an-
terior coiled part and a posterior bulb, both of which
store sperm (figures 58, 66). This condition has not been
noted in other littorinids.
Egg capsule and development: The egg capsule has
been described only for P. infracostata. It was first il-
lustrated by Tokioka (1950) (figure 64) as 'Littorina-
capsula habei', and subsequently identified by Habe
(1956, as P. roepstorffiana). The pelagic capsule is 150-
200 ^m in diameter and contains a single egg 70 ^m in
diameter. It is of the "cupola' type, of basically hemi-
spherical shape with the domed surface sculptured by
five concentric rings. The capsule has also been illustrated
by Amio (1963); this less detailed drawing shows a pill-
box shaped capsule with a circumferential flange. All
littorinids with pelagic egg capsules undergo plankto-
trophic development, and this type of development is
confirmed by the shape and size of the protoconch of
Peasiella [see Reid (1989) for a review].
Radula: Characters of the radula are relatively constant
within the genus (figures 20-30), and, with one exception,
do not provide useful taxonomic characters. The central
tooth varies from rectangular to almost square in the
group (length/width at mid-point = 1.05-1.57), but does
not show consistent intraspecific differences. There are
two prominent basal projections and three tooth cusps
(sometimes with two extra denticles). .^11 the radular
cusps are moderately long, and are pointed or rounded.
The cusp patterns of the lateral and inner marginal teeth
are similar to each other, and there are four cusps on
each. The outer marginal tooth has three cusps in most
species (but five to six in P. infracostata) and there is no
outer projection on the base.
Alimentary system: The ducts of the salivary glands
pass through the nerve ring around the anterior esopha-
gus, and there is glandular material both anterior and
posterior to the ring.
ECOLOGY
Little is known about the ecology of Peasiella species.
The available information about their habitats is sum-
marized in the systematic section. All species are appar-
ently predominantK intertidal, occurring most com-
monly in the mid- to upper eulittoral zone on hard
substrates, often in the empty shells of barnacles. They
do, however, occur to lower levels on the shore than the
littorinids with which they are sympatric, and there are
reports of sublittoral occurrences. The> can be found on
various substrates, including limestone, granite and vol-
canic rocks, beach rock, coral rubble and mangrove trunks,
and occur in a range of conditions of exposure to wave
action. Ohgaki (1985) reported that an unidentified Pea-
siella species (probabK including both P. roepstorjfiana
and P. lutulenta) occurred only on sheltered shores in
Hong Kong. As discussed below, distribution records sug-
gest a marked preference by each species for either con-
tinental coasts, promontories and high islands, or atolls.
Peasiella species are usually common w here the)' occur.
In Japan, P. infracostata reached a density of 5 per 12.5
Figures 57-66. Female reproductive tract and egg capsule of Peasiella species. Abbreviations: b, bursa; cm, columellar muscle;
hg, hypobranchial gland; i, intestine; jg, jelly gland in straight section of pallial oviduct; k, kidney; oag, opaque albumen gland;
ocg, opaque capsule gland; r, rectum; ro, renal oviduct; sr, seminal receptacle; tag, translucent albumen gland; teg. translucent
capsule gland. 57-63. P. roepstorffiana. Picnic Bay, Magnetic I., Queensland (BMNH). 57. Pallial oviduct in situ, from right side
ot body whorl, showing relation to surrounding organs 58. Diagram of spiral route of egg groove in figure 57, show ing orientation
of sections in figures 59-6;3 (solid line, egg groove; thickened line, portion of egg groo\e through capsule gland; dashed line, renal
oviduct). 59-63. Sections through pallial oviduct in figure 57 (arrow in figure 63 indicates orientation from which figure 57 is
viewed). 64. Egg capsule of P. infracostata. Ago and Tanabe Bays, Japan (after Tokioka, 1950). 65, 66. P. lutulenta, Sai Kung,
Hong Kong (BMNH). 65. Entire pallial oviduct. 66. Diagram of spiral route of egg groove in figure 65 (conventions as in figure 58).
D. G. Reid, 1989
Page 51
teg tag ocg
63 62 61 60
sr oag
ocg
Page 52
THE NAUTILUS, Vol. 103, \o. 2
cm- on a moderately exposed shore at Amakusa (Mori
et al., 1985; Tanaka et ai, 1985), while Habe (1958)
recorded up to 4,200 per 25 cm- at Tanabe Bay. Taylor
(1976) noted that the nuiricoidean gastropod Drupella
cariosa included Peasiella in its diet.
The spawning season of P. infracostata in Japan is
from May to September (Kojima, 1960; Amio, 1963; Hi-
rai, 1963; Ohgaki, 1981 ). The heat resistance of this species
has been measured by Fraenkel (1966).
FOSSIL RECORD
Cenozoic fossils of Peasiella are not uncommon in France,
and at least one species, P. reyti, may have lived in a
sheltered, perhaps muddy, environment. The oldest are
of Middle Eocene age, the youngest from the Upper
Miocene. In addition, there is a single specimen known
from the Upper Oligocene of New Zealand. Among the
material that has been examined, four species have been
distinguished, and are described in the s\stematic sec-
tion. Two of these have rather long histories, P. reyti of
about 25 M\, P. rocpstorjfiana (if the solitary fossil is
correctly determined) of 30 My.
BIOGEOGRAPHY
The Recent species of Peasiella are restricted to the Indo-
Pacific region and this, together with the European fossil
occurrences, suggests an originally Teth\an distribution.
Most of the species are exclusively tropical, but P. in-
fracostata reaches a latitude of 33°S in South Africa and
41°N in Japan.
There is a close correspondence beween geographical
distribution and the environment in which the species
occur. Peasiella lutulenta inhabits sheltered and some-
what muddy sites, and is found only on continental coasts.
Peasiella roepstorffiana and P. isseli occur in cleaner
water, on promontories and high islands, and therefore
extend from the continental margins to high oceanic
islands in the Pacific and Indian Oceans respectively.
Peasiella tantilla also belongs in this ecological group,
and is endemic to the Hawaiian Islands. Peasiella infra-
costata appears to be intermediate between these two
groups; it does not tolerate such turbid conditions as P.
lutulenta, and occurs on both continental coasts and high
islands, but does not extend far into oceanic areas. Only
P. conoidalis is common on atolls, and has a wide, but
di.sjunct, distribution in the central Pacific and central
Indian Oceans. A similar situation occurs in the littorinid
genus Littoraria and in some other gastropods, in which
species within a genus can be classified as "oceanic or
'continental' (Reid, 1986), but the species of Peasiella
suggest a spectrum from one extreme to the other. There
is some overlap between Peasiella species in these cat-
egories of habitat and distribution, and the highest di-
versity is found in South East Asia and the western Pa-
cific, where P. roepstorffiana, P. lutulenta and P.
infracostata occur together on certain continental shores
{e.g., Queensland and Hong Kong).
PHYLOGENETIC RELATIONSHIPS
OF PEASIELLA
In a recent cladistic analysis of all subgenera of the Lit-
torinidae, it was shown that Peasiella is a member of the
relativeK deri\ed subfamiK Littorininae, based on the
synapomorphies of paraspermatic nurse cells, capsule
gland and egg capsules (Reid, 1989). The superficial re-
semblance of the shell to that of Bembicium (a member
of the primitive Lacuninael is therefore a case of con-
vergence. Within the Littorininae, the affinities of Pea-
siella remain uncertain. The trochoidal shell shape and
multispiral operculum are synapomorphies \Nith Tecta-
rius. Howe\er. these may be consergent, in \iew of the
lack of longitudinal division of the foot, the superficially
closed penial duct, absence of rods in the paraspermatic
nurse cells, and double spiral loop of the pallial o\iduct,
which are found in Peasiella but not in Tectarius. There
are no important synapomorphies with Cenchritis or
with Mainicaringia Ne\ill, 1885, two other littorinine
genera with uncertain relationships (Reid, 1989). One
possible interpretation is that the four characters listed
above are plesiomorphic, as the\- are believed to be in
Melarhaphe (the most primitive member of the Litto-
rininae). If so, Peasiella ma\ be the sister-taxon of all
the remaining littorinine genera (excepting Mela-
rhaphe) This suggestion is supported b>- the histology of
the glands of the pallial oviduct (which is intermediate
between the epithelial state of Melarhaphe and the sub-
epithelial state of other littorinines) and by the absence
of spermatozeugmata in Peasiella and Melarhaphe (if
this absence is indeed a plesiomorphic character). Clearly
further characters are required to test this hypothesis.
Within the genus Peasiella the three species P. tantilla,
P. conoidalis and P. roepstorffiana share the character
state of a vestigial or absent bursa copulatrix. Presence
of the bursa is a plesiomorphic character of the Litto-
rinidae, so its absence in these three species is apomor-
phic. Some similarities in shell shape and color pattern
are also shared by this group, which ma\ be a mono-
phyletic one.
SPECIES EXCLUDED FROM PEASIELLA
Confusion with the genus Bembicium (= Risella) has
been discussed in the introduction, and the species of
Bembicium have recentK been re\ised b\ Reid (1988).
The following species have in the past been assigned to
Peasiella, but are now excluded:
Fossarus caledonicus Crosse, 1874
Classified as Risella (Peasiella) b\ Tr\on (1887), this
species is a member of the Omphalotropinae (Assimi-
neidae, Truncatelloidea) (W. F. Ponder, personal com-
munication; Reid, 1988).
Tectarius luteus Gould, 1861
Grouped with Peasiella species b\' Smith (1884), this is
D. G. Reid, 1989
Page 53
a Plesiotrochiis (Cerithioidea) (Yen, 1944; see figure of
holotype in Johnson, 1964).
Peasiella roosevelti
Bartsch & Hehder, 1939
Described from the Galapagos Islands, this is a synonym
of Nodilittorina porcata (Philippi, 1846).
Trochus risella Benoist, 1874
(= Risella girondica Benoist, 1874),
Xenophora rhytida Cossmann, 1899
These were erroneously listed as members of Peasiella
by Reid (1988), and are discussed below.
SYSTEMATIC DESCRIPTIONS
Diagnosis of Genus Peasiella
Peasiella Nevill, 1885
Risella (Peasiella) Ne\'iil, 1885:159 [type by original designa-
tion Trochus tantillus Gould, 1849].
Teleoconch small (up to 6.6 mm diameter); usually tro-
choidal and umbilicate, up to 5 whorls. Protoconch 0.21-
0.28 mm diameter, 2.3-2.8 whorls, terminated by sinu-
sigera ridge. Operculum round, corneous, multispiral.
Mesopodial sole not longitudinally divided. Male repro-
ductive system: prostate and anterior vas deferens open;
penial vas deferens superficialK closed; penis long, not
bifurcate, some simple subepithelial glandular cells pres-
ent and usually a single mamilliform penial gland. Para-
spermatic nurse cells lacking rods and not forming sper-
matozeugmata. Female reproductive system: bursa
present or absent; egg groove coiled in 2 spiral loops,
each of half a whorl; glands of pallial oviduct at least
partly subepithelial; capsule gland present. Spawn of pe-
lagic, cupola-t\pe capsules containing single eggs; de-
velopment planktotrophic. Radula; rachidian tooth rect-
angular to almost square, 3 main cusps; 4 cusps on lateral
and inner marginal; outer marginal lacking basal pro-
jection, 3-6 cusps. Salivary glands constricted by nerve
ring around oesophagus.
Key to Recent Species of Peasiella
The following key employs only shell characters, and
because these are variable one species appears twice. The
only anatomical characters useful for identification are
the lack of the bursa copulatrix in P. tantilla, P. conoi-
dalis and P. roepstorffiana. and the presence of 5-6 cusps
on the outer marginal tooth of P. injracostata. The dis-
tribution maps may also assist with identification.
1. Shell color pattern of 45-55 oblique a.xial lines on
body whorl P. tantilla
- Shell color pattern otherwise 2
2. Dorsal surface smooth or with fine spiral micros-
triae only 3
- Dorsal surface with fine spiral microstriae, major
spiral grooves, and sometimes radial folds and ca-
rina at shoulder 4
3. Base with 4-5 spiral ribs, extending to edge of
umbilicus; dorsal color pattern of 7-13 brown spots
at periphery, often joined to suture by oblique
lines, or of 2-7 brown spiral lines P. injracostata
- Base with 1-6 spiral ribs, on outermost V2-% of
basal radius only; dorsal color yellow or orange,
sometimes with darker tessellation, or single spiral
brown band, or 9-12 peripheral spots only
P. roepstorffiana
4. Base of columella thickened and angled; basal
sculpture of 1-3 ribs near periphery, separated by
a gap from up to 3 smaller ribs around umbilicus
P. lutulenta
- Base of columella not thickened or angled 5
5. Outline equilaterally conical or taller (height/di-
ameter ratio > 0.87); often with radial folds and
crenulated margin; base with 5-8 ribs from pe-
riphery to umbilicus; color white or cream, some-
times with faint brown dots in spiral grooves
P. conoidalis
- Outline more depressed 6
6. Base with 1-6 spiral ribs, on outermost '/2-% of
basal radius only; no shoulder carina; periphery
sometimes crenulated P. roepstorffiana
- Base with 3-8 spiral ribs, becoming finer and more
closely spaced towards umbilicus; usually angled
or carinate at shoulder; periphery not crenulated
P. isseli
Recent Species of Peasiella
Peasiella tantilla (Gould, 1849)
(figures 1, 20-22, 31-36, 51, 67-73, 81)
Trochus tantillus Gould, 1849:118 [holotype USNM 5615: 34
paratypes MCZ 169392, not seen; Sandwich Is (= Hawai-
ian Is)],— Gould, 1852:184-185, 503, pi. 13, figs. 215,
215a, b.
Risella tantilla. — Martens & Langkavel, 1871:41; Smith, 1876:
552.
Risella [Peasiella] tantillus.— NeviW, 1885:159.
Ri.'iella (Peasiella) tantilla.— Tryon, 1887:263, pi. 50, figs. 32-
34.
Peasiella tantillus. — Kesteven, 1903:633, fig. 1 (radula, oper-
culum).
Peasiella tantilla.— Kny, 1979:74, fig. 24A.
Bembicium tantillum. —Cernohorsky, 1978:43-44, pi. 11,
fig. 5.
Margarita angulata A. .'Xdams, 1853: 190 [t\pe not found; Sand-
wich Is].
Risella parvula Dunker, 1861:42-43 [5 s\ntypes ZMB; Sand-
wich Is].
Trochus diminulivus Reeve, 1862: pi. 1 1, sp. 57 [t> pe not found;
Oahu Is (Hawaiian Is)].
Shell (figures 1,67-73): Dimensions: Adult size range
3.3-6.6 mm diameter, mean height/diameter ratio 0.877
(95% confidence limits ± 0.034, range 0,744-1.093, n =
30 from 13 localities).
Page 54
THE NAUTILUS, Vol. 103, No. 2
D. G. Reid, 1989
Page 55
Shape: Teleoconch 3-5 whorls. Outline equilaterally
conical or taller; sides convex; sometimes turreted by
carinate rib on shoulder; base flat. Peripheral keel with
strong rib; margin not undulating or crenulated; suture
indistinct or only slightK impressed. Umbilicus small or
closed. Columellar pillar gently concave and rounded at
base.
Sculpture: Protoconch 0.24 mm diameter, 2.8 whorls.
Teleoconch whorls with 5-10 spiral grooves above pe-
riphery, usually rather indistinct; single prominent rib
at shoulder in carinate forms. Microsculpture of fine spi-
ral striae over whole surface. Base with (1 )2-4 spiral ribs,
strong in carinate forms, outermost rib strongest, becom-
ing weaker and more closely spaced towards umbilicus.
Color: Shell whitish, golden \ellow or red brown. Pat-
tern of oblique axial lines of red brown or black, 45-55
at periphery of last whorl, but pattern sometimes faint;
spiral grooves sometimes darker, giving tessellated pat-
tern. Base white to red brown, alternating white and red
brown spots on ribs. Columella whitish, red brown at
base. Aperture cream to red brown, exterior pattern
showing through.
Animal: Head-foot (figures 31, 32, 35, 36): 2-3 black
bands across snout; tentacles unpigmented or with 2 lon-
gitudinal black lines; sides of foot black or mottled.
Penis (figures 31, 33, 34):
slender.
Single penial gland; filament
Paraspermatic nurse cells (figure 51): Irregularly
rounded, packed with round granules.
Pallial oviduct: Bursa vestigial or absent.
Radula (figures 20-22): Outer marginal with 3 cusps.
Distribution: Habitat: Crevices and pools on exposed
and sheltered rock> shores, usually in upper eulittoral
and supralittoral zones (Kay, 1979; personal observation),
also in shallow subtidal zone on corraline algae (S. Kool,
personal communication).
Range (figure 81): Hawaiian Is.
Records: Oahu: Waialee (MCZ); Kahuku (AMS, 16, 19);
Coconut I., Kaneohe Bay (DGR, 13, 29); Kuilima (DOR);
Pupukea (BMNH, 33, 19); Diamond Head (USNM); Maui:
Honokowai (USNM); Kauai: Kealia (USNM); Hawaii:
Honaunau, Kona (USNM); Hilo (USNM); Laupahoehoe
Point (USNM); Punaluu Black Sand Beach (USNM).
Remarks: This is the largest member of the genus, easily
distinguished by its color, pattern, commonly carinate
sculpture and few, strong basal ribs. It is a common
species, endemic to the Hawaiian Islands.
Peasiella conoidalis (Pease, 1868)
(figures 2, 16, 18, 23, 24,
37, 38, 74-80, 81)
Trochus conoidalis Pease, 1868:287, pi. 24, fig. 8 [4 syntypes
ANSP 18868; Paumotus (= Tuamotu Is); figure 76].
Risella (Peasiella) conoidalis.— NeviW. 1885:160; Tryon, 1887:
263, pi. 50, fig. 38; Couturier, 1907:161; Dautzenberg &
Bouge, 1933:359.
Risella conoidalis— Hed\ey, 1899:424.
Risella tantilla.—SmHh, 1876:552 [in part, not Gould, 1849].
Echinella gaidei Montrouzier in Souverbie & Montrouzier, 1879:
26-29, pi. 3, figs. 3, 3a, b [17 syntypes MNHNP; Lifou,
Loyalty Is; figure 78],
Risella (Peasiella) gaidei.— Trxon. 1887:263, pi. 50, figs. 35-
37.
Peasiella gaidei. — Oyama & Takemura, 1961: Peasiella and
Littorinopsis pl.^ figs. 4-6; Higo, 1973:46; Habe, 1984:11,
fig. 2.
Shell (figures 2, 16, 18, 74-80): Dimensions: Adult
size range 1.9-5.1 mm diameter; mean height/diameter
ratio 1.029 (95% confidence limits ± 0.045, range 0.773-
1.333, n = 42 from 22 localities).
Shape: Teleoconch 3-6 whorls. Outline equilaterally
conical or taller; sides lightly convex; whorls sometimes
shouldered; base flat to slightly convex. Periphery strong-
ly keeled, sometimes with straight, undulating, or cren-
ulated flange; suture indistinct except in strongly sculp-
tured and shouldered shells. Umbilicus usually small,
sometimes closed in small shells. Columellar pillar gently
concave and rounded at base.
Sculpture: Protoconch 0.21-0.22 mm diameter, 2.8
whorls. Teleoconch whorls with (4)6-8 strong, equidis-
tant grooves above periphery, occasionally increasing to
12 on last whorl; short radial plications sometimes present
at suture, giving 3 subsutural ribs a beaded appearance;
third rib below suture occasionally enlarged, making
whorls shouldered. Periphery sometimes with 9-15 short
axial folds, crenulating margin. Microsculpture of fine
spiral striae over whole surface. Base with (4)5-8 spiral
ribs, outermost largest; finer and more closely spaced
towards umbilicus.
Color: Shell white, cream or orange yellow; pattern
usually indistinct, but spiral grooves sometimes darker
or marked by brown dots, and peripher) occasionally
with orange or brown marks corresponding with axial
Figures 67-73. Peasiella tantilla. 67, 70. Pupukea, Oahu, Hawaiian Is (BMNH) 68. 69. Mokuoloe I , Oahu (USNM 346450).
71-73. Keaukaha, Hilo, Hawaii (USNM 339423). Figures 74-80. P. conoidalis. 74, 79. Otepipi, Anaa, Tuamotu Is (BMNH).
75, 80. Takume. Tuamotu Is (USNM 723721). 76. S\'ntype of Trochus conoidalis Pease, 1868, Tuamotu Is (ANSPT8868). 77.
Mataira 1., Raroia, Tuamotu Is (USNM 698654). 78. Syntype of Echinella gaidei Montrouzier in Souverbie & Montrouzier, 1879,
Lifou, Loyalty Is (MNHNP).
Page 56
THE NAUTILUS, Vol. 103, No. 2
Figure 81. Distributions of Peasiella tantilla (asterisks) and P. conoidalis (dots).
folds. Base unmarked. Aperture and columella orange
)ello\\ .
Animal: Head-foot: Unpigmented.
Penis (figures 37, 38): Single penial gland; filament slen-
der.
Pallial oviduct: Bursa apparently absent.
Radula (figures 23, 24): Outer marginal with 3 cusps.
Distribution: Habitat: ,\tolls and oceanic islands. On
and under coral rubble or under thin layer of sand, on
intertidal reef flats (X'iader, 1951; J. Trondle, personal
communication); on limestone in high intertidal zone (G.
J. Vermeij, personal communication); on beachrock in
barnacle zone (J. D. Taylor, personal communication).
Range (figure 81): Maldive Is, Seychelles, southern and
western Pacific, from southern Japan to Queensland,
Marshall and Tuamotu Is.
Records: Seychelles: Aldabra Atoll (BMNH); Maldive
Is: Dunidu L, North Male (ANSP); Fadiffolu (ANSP);
Chagos Arch.: Diego Garcia (Viader, 1951 ); Japan: Yaku-
jima (NSMT); Oku, Okinawa, Ryukyu Is (LACM); Tai-
wan: Yeh-Liu (.\NSP); Philippines: Binanga Port, Luzon
(USNM); Corregidor I., Luzon (ANSP); 10 km N of Ca-
lavag, Palawan (D. Kadolsky Colin); Mariana Is: Taga-
chan Point, Guam (G. J. Vermeij Colin); Caroline Is:
Ponape (AMS); Marshall Is: Engebi I., Eniwetok (USNM);
Bikini (USNM); Busch I., Rongelap (USNM); Enybor I.,
Jaluit (USNM); Gilbert Is: Tarawa (AMS); Ellice Is: Fu-
nafuti (AMS); Australia: Queensland: Lizard I. (ANSP);
Papua New Guinea: Rabaul, New Britain (AMS); Bou-
gainville Strait (AMS); Milne Bay (AMS); Solomon Is:
NW Sta Isabel I. (LACM); Laulosi I., Malaita I. (AMS);
New Hebrides: Malakula I. (AMS); Efate I. (LACM);
Loyalty Is: Lifou (AMS, MNHNP); New Caledonia: Pa-
nie (Hedley, 1899); Fiji: Deuba (AMS); Samoa; Swains I.
(USNM); Line Is: Palmvra (USNM); Society Is: Moorea
(LACM); Tahiti (BMNH); Tuamotu Is: Makatea (USNM);
Rangiroa (USNM); Manihi (LACM): Anaa (BMNH, 33,
39); Marutea du Sud (AMS); Takume (USNM); Oneroa
(USNM); Amanu (USNM).
Remarks: This wideK distributed species is distin-
guished from P. tantilla b\ its pale color, finer and more
numerous basal ribs, and frequent!) crenulated margin.
The tall outline, basal sculpture of fine ribs extending
right to the umbilicus, and the dorsal color pattern, sep-
arate it from P. roepstorffiana.
Peasiella roepstorffiana
(Nevill, 1885)
(figures 3-6, 17, 19, 25, 26,
39-42, 52, 53, 57-63, 82-89, 96)
'^Risella (Peasiella) tantillus var. parvula. — Nevill, 1885:159
[not Dunker, 1861].
Risella (Peasiella) roepstorffiana Nevill, 1885:161 [1 .syntype
seen, here designated lectotype, ZSI 1916; .\ndaman Is;
figure 89; 5 possible syntypes BMNH 1989005; S. Anda-
man].—Tryon, 1887:264, pi. 50, fig. 43.
Risella [Peasiella) templiana Nevill, 1885:161 [1 syntype seen,
here designated lectotvpe, ZSI 1913; Andaman Is; figure
83].
Risella (Peasiella) templiana var. nigrofasciata Nevill, 1885:
161-162 [1 svntvpe seen, here designated lectotype, 4.8
X 3,2 mm, ZSI M 17732 3; Andaman Is].
Risella (Peasiella) templiana \ar, sulnmhricata Nevill, 1885:
162 [1 s\ nt\ pe seen, here designated lectotype, ZSI 4.8 x
4.1 mm; .Andaman Is].
D. G. Reid, 1989
Page 57
Risclla haltcata Preston, 1908:197, pi. \4. fig. 13 [2 syntypes
HMNH 1908.7.2.68-69; Andaman Is].
?fi!6c//a (an(i7/a.— H«llfy, 1910:355 [not Gould, 1849].
Peasiella sp. Morton & Morton, 1983:62, figs. 5.3, 5.8.
Shell (figures 3-6, 17, 82-89): Dimensions: Adult
size range 1.6-5.0 mm diameter; mean height/diameter
ratio 0.757 (95% confidence limits ± 0.022, range 0.556-
1.000, n = 101 from 55 localities).
Shape: Teleoconch 3.5-5 whorls. Outline varying from
rather depressed to equilaterally conical; sides convex or
rarely almost straight; whorls usually gently rounded;
base flat to slightly conve.x. Peripheral keel strongK' an-
gled, usually a sharp flange, margin usualK- straight,
sometimes undulating or crenulated; suture slightly im-
pressed or indistinct. Umbilicus open. Columellar pillar
straight to slightly concave, rounded or slightly angled
at base.
Sculpture: Protoconch 0.23 mm diameter, 2.3 whorls.
Teleoconch whorls with (3)4-8 approximately equidis-
tant spiral grooves above periphery, sometimes increas-
ing to up to 13 on last whorl, sometimes obsolete. Short,
fine, radial plications occasionally developed at suture.
Sometimes 9-12 radial folds at periphery of last whorl,
producing undulating or crenulated margin. Microsculp-
ture of fine spiral striae over whole surface. Base with
(1)3-5(6) approximately equidistant spiral ribs, either
innermost or outermost strongest; innermost V3-V2 of bas-
al radius usually lacking spiral ribs, but rarely showing
1-2 small ribs.
Color: Shell cream, golden yellow or pinkish orange,
sometimes with faint tessellated pattern over whole dor-
sal surface; spiral grooves sometimes brown; peripheral
flange occasionally brown, either continuous or broken
into spots corresponding with marginal crenulations. Shells
occasionally with dark spiral band on lower half or mid-
dle of whorls. Base usually unpatterned; underside of
peripheral flange sometimes brown or spotted. Interior
of aperture golden yellow, exterior pattern showing
through; columella \ellow to purple brown. Distinctive
red brown, tessellated color pattern occurring in Fiji,
Samoa, southeastern Papua New Guinea and Philippines
(figure 6): red brown and white spots on ribs, 11-24 at
periphery of last whorl, making oblique tessellated pat-
tern over whole surface, sometimes darkest on lower half
of whorls; base and columella red brown; tessellations
showing through in pale band within aperture.
Animal: Head-foot (figures 39-42): Black band across
base of snout, single longitudinal black line on tentacles;
head sometimes entirely unpigmented; foot with faint
black speckling or unpigmented.
Penis (figures 39-42): Single penial gland in specimens
from Queensland, but this is absent in those from Malaya,
Thailand, and Hong Kong.
Paraspermatic nurse celts (figures 52, 53): Rounded to
irregularly oval, sometimes with produced tip; finely
granular contents with indistinct nucleus; nurse cells ab-
sent in specimens from Hong Kong.
Pallial oviduct (figures 57-63): Bursa absent.
Radula (figures 25, 26): Outer marginal with 3(4) cusps.
Distribution: Habitat: Islets and promontories on con-
tinental coasts; oceanic high islands. High eulittoral zone,
on granite and volcanic rock, amongst and above bar-
nacles and oysters. Rare or absent in turbid bays.
Range (figure 96): Central Indo-Pacific, from eastern
Bay of Bengal to Hong Kong, Philippines, Palau, Queens-
land, Fiji and Samoa.
Records: Burma: Arakan (BMNH, USNM); Andaman
Is (BMNH); Thailand: Marine Biological Center, Phuket
I. (BMNH, 2<5, 12); Surin Beach, Phuket I. (BMNH, 1<5);
Ko Tao (USNM); Pattaya (J. Le Renard Colin); Malaysia:
Langkawi Is (USNM); Port Dickson (ANSP, 1<3); Mersing
(BMNH); Vietnam: Baie d'Along (BMNH); Isle de la
Table (MNHNP); Hong Kong: Ping Chau, Mirs Bay
(BMNH, 33, 19); Hoi Ha, Mirs Bay (BMNH); Hoi Sing
Wan, Tolo Channel (BMNH, 13); Lamma I. (AMS, 13);
Sai Kung (DGR, 33, 19); Philippines: Binanga Port, Lu-
zon (USNM); Scott's I., Lingayen Gulf, Luzon (LACM);
Maricaban I., Luzon (USNM); Samar I. (NSMT); Zam-
boanga, Mindanao (NSMT); 10 km N of Calavag, Pa-
lawan (D. Kadolsky Colin); Caroline Is: Arakapesan, Pa-
lau (G. J. Vermeij Colin); Papua New Guinea: Nimoa,
Louisiade Arch. (AMS); Milne Bay (AMS); Port Moresby
(AMS); Queensland: Somerset (AMS); Lizard I. (AMS);
Cooktown (AMS); Low Isles (AMS); Green I. (AMS); Dunk
I. (ANSP); Orpheus I. (DGR); Picnic Bay, Magnetic I.
(DGR, 53, 29); Gladstone (AMS); Fiji (BMNH, MCZ,
MNHNP, ANSP, USNM); Nadi Bay, Viti Levu (AMS);
Samoa: Pago Pago Harbor, Tutuila I. (LACM, AMS).
Remarks: This is a highly variable species, with a dis-
tinctive red brown, tessellated color form in the eastern
and northeastern part of its range, and a more marked
tendency towards marginal crenulation in the northern
part. Nevertheless, considerable variation in shape and
color also occurs within local populations, and recogni-
tion of geographical subspecies appears unnecessary. The
unsculptured area around the umbilicus, and common
golden yellow color with lines or tessellations, are the
most useful diagnostic characters.
Of greater significance is the apparent geographical
variation in anatomical characters. The amount of ma-
terial available for dissection was limited, but the 12
males examined from Hong Kong, Malaya, and Thailand
all lacked the mamilliform penial gland, while this was
present in the 5 from Queensland. In addition, para-
spermatic nurse cells were absent in 4 specimens from
Hong Kong, but present in 3 from Queensland. Both
nurse cells and the mamilliform gland are uniformly
present in other Peasiella species. The lack of nurse cells
could be simply a developmental abnormality or seasonal
variation (although neither has been recorded elsewhere
in the family). Penial shape is usually rather constant in
Page 58
THE NAUTILUS, \ ol. 103, No. 2
D. G. Reid, 1989
Page 59
Figure 96. Distribution of Peasiella roepstorffiana.
littorinid species, and may differentiate between closely
related species, leading to the idea that it serves the
function of species recognition {e.g., Reid, 1986). The
geographical variation in penis shape in P. roepstorffiana
could therefore be evidence that two species are in-
volved. However, the number of mamilliform glands
does show intraspecific variation, especially in littorinids
with many such glands (Reid, 1989). In species of No-
ditittorina (which normally have a single gland), abnor-
mal animals can be found with two or none. A male P.
infracostata has been found without a mamilliform gland.
For this reason, and because of the absence of any con-
sistently correlated shell characters by which two species
could be distinguished in the P. roepstorffiana group, a
single nominal species is retained here until further in-
formation is available.
A single fossil from the Upper Oligocene of New Zea-
land may be referable to this species (see below).
Peasiella lutulenta new species
(figures 7-9, 27, 43-46, 54, 55,
65, 66, 90-95, 97)
Bembicium sp. Wells & Slack-Smith, 1981:269.
Peasiella lutulenta n. sp. Etymology: Latin lutulenta, muddy,
in reference to habitat of species. Types: holotype AMS
C149052, Kissing Point, Rowes Bay, Townsville, Queens-
land (figure 9); 10 paratypes AMS C149053, 16 paratypes
BMNH 1988138.
Shell (figures 7-9, 90-95): Dimensions: Adult size
range 1.7-3.8 mm diameter; mean height/diameter ratio
0.716 (95% confidence limits ± 0.031, range 0.500-0.909,
n = 36 from 26 localities).
Shape: Teleoconch 2.5-4 whorls. Outline varying from
rather flattened to almost equilateralK conical; sides con-
vex; base flat to slightly convex. Peripheral keel with
strong flange, margin not usually undulating or crenu-
lated; suture slightly impressed. Umbilicus open. Colu-
mellar pillar straight, with thickened angle at base, form-
ing margin of umbilicus.
Sculpture: Protoconch 0.24-0.27 mm diameter, 2.4
whorls. Teleoconch whorls with (4)5-7(9) strong, ap-
proximately equidistant spiral grooves above periphery.
Microsculpture of fine spiral striae over whole surface.
Peripheral keel usually entire, rarely with 9-11 slight
crenulations. Base with 2-6 spiral ribs, outermost 1-3
ribs most prominent, usually separated from smaller ribs
around umbilicus b\ marked gap.
Color: Shell cream to pale horn color; 7-16 brown to
black spots at periphery, usually also at suture, the two
series connected by fainter irregular axial stripes; occa-
sionally axial stripes complete and conspicuous. Periph-
Figures 82-89. Peasiella roepstorffiana. 82, 85. Orpheus I., Queensland (BMNH). 83. Lectotype of Risella templiana Nevill,
1885, Andaman Is (ZSI 1913) 84. Hong Kong (BMNH). 86. Viti Is (Fiji) (BMNH). 87, 88. Ping Chau, Hong Kong (BMNH). 89.
Lectotype of Risella roepstorffiana Nevill, 1885, Andaman Is (ZSI 1916). Figures 90-95. P. lutulenta. 90, 94. Hoi Sing Wan,
Hong Kong (BMNH). 91. Shark Point, Barrow I., Western Australia (BMNH). 92, 93, 95. Cockle Bav, Magnetic I., Queensland
(BMNH).
Page 60
THE NAUTILUS, Vol. 103, No. 2
Figure 97. Distributions of Peasiella isseli (asterisks) and P. lutulenta (dots).
eral spots usually showing through at periphery of base.
Columellar pillar whitish, red brown at base. Interior of
aperture cream, exterior pattern showing through.
Animal: Head-foot (figures 43-46): 0-2 black bands
across base of snout; black pigment above and below eye,
but tentacles unpigmented; foot unpigmented or black
mottled.
Penis (figures 43-46): Single penial gland.
Paraspermatic nurse cells (figures 54, 55); Irregularly
oval, packed with round granules.
Pallial oviduct (figures 65, 66): Large anterior bursa
present.
Radula (figure 27): Outer marginal with 3 cusps.
Distribution: Habitat: Sheltered, often muddy, con-
tinental shores. On rocks and outer fringes of mangrove
forests, frequently in empty barnacles.
Range (figure 97): Central Indo-Pacific, including Hong
Kong, Malaysia, Borneo, and northern Australia.
Records: Hong Kong: Hoi Sing Wan, Tolo Channel
(BMNH, 26, 22); Hoi Ha, Mirs Bay (BMNH); Sai Kung
(DGR, 3.5, 39); Vietnam: Isle de la Table (MNHNP, MCZ,
ANSP); Baie d'Along (BMNH); Malaysia: Mersing
(BMNH, 12; AMS); Tanjong Tuan (AMS)'; Port Dickson
(ANSP, IS); Borneo: Sandakan (USNM); Balikpapan
(AMS); Papua New Guinea: Eli Beach, Port Moresby
(AMS); Australia: Western Australia: Beagle I., Onslow
(AMS); Shark Point, Barrow I. (WAM, 26, 12); Dampier
(AMS); Port Hedland (AMS); La Grange Bay (ANSP);
Turtle I., King Sound (AMS); Northern Territory: East
Point, Darwin (AMS); Maningrida (AMS); Gove Penin.
(DGR); Queensland: Sweers I., Wellesley Is (AMS); Ma-
poon (AMS); Wednesday I. (AMS); Somerset (AMS); Port
Douglas (AMS); Bingil' Bay (AMS); Dunk I. (ANSP);
Cockle Bay, Magnetic I. (DGR); Rowes Bay, Townsville
(DGR, 2(5,' 12); Bowen (AMS); Island Head Creek, N of
Yeppoon (AMS); Gladstone (AMS); Facing I. (AMS); Point
Vernon, Hervey Bay (AMS); Moreton Bay (AMS).
Remarks: The thickened columellar base is diagnostic
of this species. Other useful characters include the basal
sculpture and usually coarse, dark color pattern on the
dorsal side (which separate it from P. roepstorffiana),
the coarse dorsal sculpture (separating it from P. infra-
costata) and the lack of a shoulder keel (separating it
from many individuals of P. isseli).
Late Pleistocene fossils have been found in Shark Bay,
Western Australia (G. \\. Kendrick, WAM), but the clos-
est Recent specimens are from Onslow, 500 km to the
north.
Peasiella isseli
(Semper in Issel, 1869)
(figures 13-15, 28, 49,
50, 56, 97-105)
Trochus sp. .■Kudouiii, 1826:42.
Risclla isseli Semper in Issel, 1869:194, 347 [refers to Savigny,
1817: pi. 5, figs. 35.1, 35.2: lectotype here designated
MNHNP 1.8 X 1.4 mm. figured by Bouchet & Danrigal,
1982: fig. 65; figures 101. 103 herein; Suez].— Pallary, 1926:
84, pi. 5, figs. 35.1, 35.2.
Risella (Peasiella) issc/i.— Nevill, 1885:160-161; Tryon, 1887:
263, pi. 50. figs. 39, 40.
Risella isseli yar carinata Pallary, 1926:84-85, pi. 5, fig. 36
[reproduced from Savigny, 1817; 1 syntypeseen, MNHNP
1.8 X 1.3 mm; Suez].
D. G. Reid, 1989
Page 61
Risella isseli var. undata Pallary, 1926:84, pi. 5, fig. 35,3 [re-
produced from Sasigii), 1817; type not found; Suez].
Risella isseli var. nuiuriliana \iader, 1951:1-49, pi. 3, figs. 10,
11 [2 syntypes in Mauritius Institute, not seen; Cannonier's
Point and Grand Bav, Mauritius; 3 possible syntypes BMNH
1989004, figure 14].
Trochus sismondae Issel, 1869:225-226, pi. 2, fig. 13 [holotype
MGD 4.0 X 4 1 mm; Suez].
Shell (figures 13-15. 98-105): Dimensions: Adult
size range 1.9-5.6 mm diameter; mean height/diameter
ratio 0.758 (95% confidence limits ± 0.044, range 0.545-
1.077, n = 45 from 26 localities).
Shape: Teleoconch 2.5-3.5 whorls. Outline varying
from depressed to almost equilaterally conical; sides con-
vex, sometimes with strong shoulder carina; base convex.
Periphery strongly keeled, sometimes with strong rib or
flange; margin usually entire, rarely slightly crenulated;
suture slightly impressed. Umbilicus small. Columellar
pillar concave, slightly angled at base.
Sculpture: Frotoconch 0.26-0.28 mm diameter, 2.3-2.5
whorls. Teleoconch whorls with (6)7-9(12) equidistant
or irregularly spaced spiral grooves above periphery, oc-
casionally obsolete; shoulder rib sometimes enlarged or
carinate. Microsculpture of fine spiral striae over whole
surface. Rarely, periphery and shoulder with 7-12 radial
folds, forming slight marginal crenulations. Base with
(3)4-6(8) spiral ribs, outermost largest, finer and more
closely spaced towards umbilicus; ribs around umbilicus
sometimes bearing periostracal bristles.
Color: Shell cream, tan, spire sometimes pink or lilac,
or entire shell dark brown or black. Pattern sometimes
absent. Patterned shells with (6)8-16 dark brown spots
at periphery, sometimes connected to sutural spots by
oblique brown lines; others with faint marbled pattern
over entire shell; base usually unpatterned. Columella
white to brown; aperture cream to brown.
Animal: Head-foot (figures 49, 50): Two black lines
across base of snout, two longitudinal black lines on ten-
tacles; sides of foot blackish.
Penis (figure 50): Single penial gland.
Paraspermatic nurse cells (figure 56): Round or oval,
packed with round granules.
Pallial oviduct: Large anterior bursa present.
Radula (figure 28): Outer marginal with 3-4 cusps.
Distribution: Habitat: Continental coasts and oceanic
high islands, only rarely on atolls. Upper eulittoral zone
of rocky shores.
Range (figure 97): Western Indian Ocean, including
Persian Gulf and Red Sea, south to Mozambique and
Madagascar.
Records: Egypt: Suez (BMNH); Israel: Eilat (BMNH,
15, 22; USNM, ANSP); Sudan: Dongonab Bav (BMNH);
Port Sudan (ANSP); Ethiopia: Massawa (BNINH, NM);
Dehui I., Dahlak Is (BMNH); Saudi Arabia: Jeddah
(USNM); Yemen: Aden (BMNH); Oman: Bandar Khay-
rhan (BMNH); Persian Gulf: Abu Dhabi (BMNH); Qatar
(BMNH); Kuwait (BMNH); Tanzania: Zanzibar (BMNH);
Mozambique: Mozambique I. (NM); Benguera I., Ba-
zaruto Arch. (NM); Madgascar: Nossi Be (USNM, ANSP);
Tulear (MNHNP); Mauritius: Grand Bay (BMNH); Sey-
chelles: Bale Ternay, Mahe I. (BMNH, USNM); North-
west Bay, Mahe I. (ANSP, 59); Aldabra Atoll (BMNH).
Remarks: This is another species with very variable
shell characters. The shoulder is often keeled, but the
species is separated from similarly shouldered shells of
P. tantilla and P. conoidalis by the color pattern, a gen-
erally more depressed outline and often by the basal
sculpture. Specimens without a shoulder keel are sepa-
rated from P. lutulenta by their narrower columellar
base, and from most specimens of P. roepstorffiana by
the basal sculpture. Distinction from P. infracostata is
discussed below.
Peasiella infracostata
(Issel, 1869)
(figures 10-12, 29, 30, 47, 48,
64, 106-112)
Trochus sp. .\udouin, 1826:42.
Risella infracostata Issel, 1869:195-196 [refers to Savigny, 1817:
pi. 5, fig. 40; lectotype, here designated, MNHNP 1.5 x
1.2 mm, figured bv Bouchet & Danrigal, 1982: fig. 64;
Suez; figure 109 herein].— Pallary, 1926:86, pi. 5, fig. 40;
Dautzenberg, 1929:496.
Risella (Peasiella) infracostata.— NeviW, 1885:160; Tryon, 1887:
264, pi. 50, figs. 41, 42.
Risella (Peasiella) tantillus var. subinfracostata Nevill, 1885:
160 [1 syntype seen, here designated lectotype, ZSI, 2.3 x
2.2 mm; Nicobar Is].
Ctjclostrema fuscopiperata Turton, 1932:198-199, pi. 51, fig.
1379 [2 syntypes, OUM; Port Alfred, South Africa].
Peasiella roepstorffiana. — Habe, 1956: fig. .\ (egg capsule);
Oyama & Takemura, 1961: Peasiella and Littorinopsis
pi., figs. 1-3; Yamamoto & Habe, 1962:16, pi. 3, figs. 3, 4,
34, 35; Amio, 1963:303, figs. 22a,b (egg capsule); Habe,
1964:28, pi. 9, fig. 23; Higo, 1973:46; Habe, 1984:11, fig.
1 [all not Nevill, 1885].
'Littorina-capsula habei' Tokioka. 1950: fig. 6.1 (egg capsule).
Shell (figures 10-12, 106-111): Dimensions: Adult
size range 1.1-3.5 mm diameter; mean height/diameter
ratio 0.846 (95% confidence limits ± 0.022, range 0.600-
1.120, n = 79 from 39 localities).
Shape: Teleoconch 2.5-4 whorls. Shell thin and trans-
lucent, or solid. Outline varying from almost equilat-
erally conical with rounded whorls to depressed globular
or almost flat with inflated whorls; in Japan, whorls flat;
base flat to convex. Periphery either uniformly rounded,
or with weak or strong rib, or (in Japan) with strong
flange; margin usually entire, rarely slightly crenulated;
suture usually impressed. Umbilicus open, sometimes very
small. Columellar pillar straight to uniformly rounded,
curved and not thickened at base. Aperture occasionally
somewhat patulous.
Page 62
THE NAUTILUS, Vol. 103, No. 2
D. G. Reid, 1989
Page 63
Figure 1 12. Distribution of Peasiella infracostata.
Sculpture: Protoconch not seen, apex usually eroded.
Teleoconch whorls commonly smooth above periphery;
spiral microstriae sometimes present and (in Japan) 7-
10(11) equidistant spiral grooves. Base with (2)4-5 ap-
proximately equidistant spiral ribs, becoming a little
stronger towards periphery. Basal and peripheral ribs
sometimes bearing small periostracal bristles.
Color: Shell white to horn color; spire sometimes dark
grey, black or purple brown. Shells from Indian Ocean
(figure 11): suture and periphery with alternating brown
and opaque white spots (9-13 brown spots at periphery
of last whorl); sutural and peripheral spots sometimes
connected by oblique brown lines; spiral brown line
sometimes placed just above suture on spire whorls; base
unmarked or with peripheral brown line. Shells from
Pacific Ocean (figure 12): 2-7 brown or black spiral lines
or dashes above periphery, with or without brown spots
at suture and periphery; 3-4 lines in grooves of base.
Shells from Japan (figure 10): black, eroded spire; pe-
riphery of last whorl paler, with 7-10 smudged brown
spots, sometimes forming oblique stripes or continuous
band; base with 7-10 brown spots or band in outermost
groove and brown band around umbilicus. Aperture
cream with exterior pattern showing through; columella
white to purple.
Animal: Head-foot (figures 47, 48): Head with 2 black
bands across base of snout, posterior one broad; tentacles
usually unpigmented, sometimes black pigment above
and below eye or single longitudinal black line; sides of
foot black, grey or unpigmented.
Penis (figures 47, 48): Single penial gland present, oc-
casionally absent.
Pallial oviduct: Large anterior bursa present.
Egg capsule (figure 64): Cupola type, with 5 concentric
rings.
Radula (figures 29, 30): Outer marginal with 5-6 slen-
der cusps.
Distribution: Habitat: Usually on continental coasts,
sometimes on high islands. On sheltered and moderately
exposed shores, from mean high water of spring tides to
below mid-tide level, in crevices and empty barnacles,
especialK on sheltered sides of rocks (Mori et at., 1985).
Range (figure 112): Margins of Indian Ocean, from
South Africa to Red Sea, margins of western Pacific Ocean,
from Japan to Queensland.
Records: Egypt: Suez (MNHNP); Yemen: Aden
(BMNH); Somalia: Mogadiscio (ANSP); Tanzania: Zan-
zibar (BMNH); Mofambique: Benguera 1., Bazaruto Arch.
(NM); South Africa: Mapelane, Zululand (NM); Um-
dhloti, Natal (NM); Mbotvi, Pondoland (NM); East Lon-
don (NM); Port Alfred (OUM); Pakistan: Karachi (BMNH,
Figures 98-105. Peasiella isseli. 98. Suez (BMNH 88.10.14). 99. Bale Ternay. Mahe, Seychelles (BMNH). 100. 102, 104. 105.
Bandar Khayrhan, Oman (BMNH). 101, 103. Lectotype of Riselta isseli Semper in Issel, 1869, Suez (MNHNP). Figures 106-
111. P. infracostata. 106-108. Karachi, Pakistan (BMNH). 109. Lectotype of Risella infracostata Issel, 1869, Suez (MNHNP).
110. Pattaya, Thailand (J. Le Renard Colin LR99012). 111. Zanpa Point, Okinawa, Ryukyu Is (AMS C146702).
Page 64
THE NAUTILUS, Vol. 103, No. 2
2<5); 7 km WNVV Bulegi Point, Sind Prov. (LACM); India:
Mada I., Bombay (ANSP); Nicobar Is (ZSI); Thailand:
Pattaya (J. Le Renard Colin); Borneo: Berhala Channel,
Sandakan (USNM); Bak-Bak. Kudat (USNM); Vietnam:
Isle de la Table, Tonkin (MNHNP); Hong Kong: Hoi
Sing Wan, Tolo Channel (BMNH, 13); Lok Wo Sha, Tolo
Channel (BMNH); Ping Chau, Mirs Bay (BMNH); Hoi
Ha, Mirs Bav (BMNH, 22); japan: Zanpa Point, Okinawa,
Rvukyu Is (AMS 19; USNM, NSMT); Kyushu: Tomioka,
Kumamotu Pref. (NSMT); Nagasaki (USNM); Shikoku:
Tatsukushi, Kochi Pref. (NSMT, USNM); Honshu: Ta-
nabe Bay, Wakayama Pref. (USNM); Kuchino, Shiznoka
Pref. (BMNH, 43); Arasaki, Kanagawa Pref. (NSMT,
USNM); Kominato, Aomori Pref. (LACM, ANSP); Asa-
mushi, Aomori Pref. (NSMT, USNM); Papua New Guinea:
Duke of York I., Rabaul, New Britain (.AMS); Australia:
Queensland: SW Lizard I. (LACM); Port Douglas (AMS);
Green I. (AMS); Halfmoon Bay (AMS); Ellis Beach (AMS);
Turtle Bay, Cape Ferguson (BMNH); Facing I., Port
Curtis (AMS).
Remarks: This species is the most variable in the genus
in its shell form and coloration, with recognizable forms
in several areas of its wide geographical range. Shells
from the Indian Ocean, South East Asia and Queensland
are globular, with inflated whorls lacking grooves or mi-
crostriae above the round or slightK angled periphery
(figures 106-110), and quite unlike any other members
of the genus. The difference between the oblique or
spotted color pattern of shells from the Indian Ocean
(figure 11) and the spiral lines of shells from the south-
western Pacific (figure 12) is rather consistent. However,
it is probably of minor importance, since shells from the
Indian Ocean may show a spiral line on the spire, and
on the body whorl the oblique lines may become almost
perpendicular to the apertural lip. In addition, shells
from the Pacific ma\ develop sutural and peripheral
spots like those in shells from the Indian Ocean.
Shells from Japan are very different, distinguished by
conical shape, peripheral flange, frequent presence of
microstriae and grooves above the periphers', and color
pattern of a blackish spire and peripheral spots (figures
10, 111). They are, however, connected to the typical
form of the species by intermediates from China and
Thailand (figure 110), which show the typical globular
shape and lined pattern, but with spiral grooves and
peripheral keel as seen in Japanese shells. Specimens from
Okinawa are especially variable, ranging from smooth
to strongly grooved, keeled to globular, and with black
or lined spire whorls. Keeled and faintK grooved shells
of F. infracostata do occasionalK occur in the Indian
Ocean (figure 106). The basal sculpture is similar in all
parts of the range. Radulae have been examined in single
specimens from Karachi and Cape Ferguson, Queens-
land, and in two specimens each from Kuchino and To-
mioka, Japan; all show the 5-6 sharp cusps on the outer
marginal teeth, which are diagnostic of this species (fig-
ures 29, 30).
The Japanese form of P. injracostata resembles those
shells of P. isseli without a shoulder keel. Both species
can show a similar color pattern of oblique brown lines.
Some collections of juvenile specimens from southern
Africa (here doubtfulK assigned to P. injracostata) con-
tain keeled, shouldered, grooved shells which resemble
P. isseli. Nevertheless, these two species are considered
to be distinct, for the following reasons. Typical forms
of both species are present in mi.xed samples from Zan-
zibar (BMNH), Aden (BMNH) and the Bazaruto Archi-
pelago (NM). Localit) records suggest that P. injracos-
tata is a continental species (at least over most of its
range in the Indian Ocean), while P. isseli is somewhat
more oceanic, extending to islands in the western Indian
Ocean. P. injracostata has not been reliabK recorded
from the Persian Gulf or Red Sea (the type locality, Suez,
is therefore doubtful), where P. isseli occurs commonly.
In areas where both species occur there are consistent
differences in shell shape. In adult P. injracostata there
is no shoulder keel, grooves are usualK absent above the
periphery, there are no more than 5 basal ribs, and the
periphery is usualK' rounded. In P. isseli the shoulder is
frequently keeled, groo\es are present above the pe-
ripherv, basal ribs ma\ number more than 5, and the
peripherv is sharpK keeled. Shell color patterns also dif-
fer: in P. injracostata the spire whorls often bear a spiral
brown line, or ma\ be entirely black; in P. isseli both
patterns are rare. The most important anatomical dif-
ference is the presence of 5-6 narrow cusps on the outer
marginal teeth of P. injracostata (3-4 in P. isseli). A
possible difference in coloration of the tentacles requires
confirmation in additional specimens.
Fossil Species of Pe.\siella
Peasiella minuta (Deshayes, 1824)
(figures 113-115)
Troclws minutus Deshayes, 1824:239-240. pi. 29, figs. 15-18
(.\tlas. 1S:37) [t> pe not seen; Valmondois]. — Deshayes, 1864:
956.
Risella minuta. — Cossmann, 1888:257-258; Cossmann. 1899:
325, pi. 23, fig. 24; Cossmann & Pis.sarro, 1907-1913: pi.
17, fig. 108.1; Cossmann, 1915:74, pi. :3, figs. 27-29.
Bemhicium minutum. — Gilbert, 1962:24.
?Xenophora bouryi Cossmann, 1888:189, pi. 7, figs. 36-38 [type
not seen; Neauphlette, Paris Basin, Eocene].
Shell (figures 113-115): Dimensions: Adult size range
3.1-6.6 mm diameter; mean height diameter ratio 0.810
(95% confidence limits ± 0.074, range 0.545-1.048, n =
17 from 6 localities).
Shape: Outline equilateralK conical or lower; periphery
sharpK angled or keeled, slight flange in juveniles; mar-
gins not crenulated; suture slightly impressed. Umbilicus
becoming closed in larger shells.
Sculpture: Teleoconch whorls with 6-9( 13) spiral grooves
above periphery; sometimes with up to 30 oblique radial
folds on last whorl. Base with 2-6(10) ribs, innermost
outlining umbilicus, iiuier 2 often most prominent.
Distribution: Range: Paris Basin (Upper Eocene) and
Loire (Middle Eocene) of France.
D. G. Reid, 1989
Page 65
Figures 113-120. Fossil species of Pfasie//a. 113-115. P. minuta. 113, 114. L'Ecaillette, Monneville, Paris Basin, Marinesian
(J. Le Renard Colin LR99004) 115. Ronquerolles, Paris Basin. Auversian (J. Le Renard Colin LR99002). 116. P. pontileviensis,
Ferriere-Larfon, Loire, Middle Miocene (MNHNP). 117. P aff roepstorffiana. Shell Gully. Chatton, Southland. New Zealand,
Duntroonian(NZGS9806). 118-120. P. reyti. 118, 120. LeBois-Gouet, Loire.\tlantique, Biarritzian(J. Le Renard Colin LR99010).
119.B7LSaubotte, Noaillan, Aquitanian (MNHNP).
Records: France: Paris Basin: Le Ruel, Bartonian
(MNHNP); Le Ruel, Marinesian; Le Quoniam, Marine-
sian; L'Ecaillette, Monneville, Marinesian; Ronquerolles,
Auversian; Baron, Auversian; Barisseuse, Auversian;
Mery-sur-Oise, Auversian; Loire: Saint-Lubin-de-la-Haye,
Lutetian (all J. Le Renard Colin).
Remarks: This relatively common species shows vari-
ation in the development of the peripheral flange, closure
of the umbilicus, and number of basal ribs. The com-
bination of conical shape, marked spiral grooves above
the periphery, and strong basal ribs extending to the
umbilicus, is diagnostic.
The identity of Xenopliora bouryi is uncertain; the
description of radial costules and 3 basal ribs, and ap-
parent lack of a thickened columellar base, resemble P.
minuta. However, the depressed shape, sinuous marginal
flange and apertural form are different, and, taken to-
gether, suggest that it may not be a Peasiella.
Peasiella reyti
(Cossmann & Peyrot,
(figures 118-120)
1917)
Trochus //iorinus. —Benoist, 1874b:316 [not Grateloup. 1832,
fide Cossmann & Pe\rot, 1917:63].
Pseudonina reyti Cossmann & Pe>rot, 1917:363-365, te.xt fig.
30, pi. 6, figs. 45-48 [type not seen; Merignac, Aquitaine;
Aquitainian].
Peasiella girondica. —Lozouel. 1986:229 [not Risella girondica
Benoist, 1874].
Page 66
THE NAUTILUS, Vol. 103, No. 2
Shell (figures 118-120: Dimensions: Adult size range
2.3-5.3 mm diameter; mean height/diameter ratio 0.627
(95% confidence limits ± 0.065, range 0.459-0.756, n =
9 from 3 localities).
Shape: Spire relativeK low, whorls conve.x at shoulder,
concave towards periphery; suture impressed; base con-
vex, becoming concave around umbilicus. Peripheral keel
bearing strong flange, rarely slightly undulating. Um-
bilicus large (up to 0.3 of basal diameter). Columellar
pillar uniformly rounded, thin.
Sculpture: Protoconch 0.30-0.36 mm diameter, 2 to
appro.ximately 3 whorls, terminated by sinusigera notch,
but sculpture not preserved on specimens examined. Te-
leoconch whorls usually rather smooth, with growth lines
and indistinct spiral microstriae only; sometimes coarser
striations forming 11-14 indistinct grooves above pe-
riphery; radial sculpture absent, or coarsely rugose at
shoulder, sometimes with 6-14 indistinct radial plications
producing undulations of marginal flange. Edge of um-
bilicus a thickened rib, usually outlined by broad, shallow
groove; rest of base almost smooth, but with radial growth
lines (strongly developed near umbilicus), sometimes up
to 10 faint spiral striae.
Color: Sometimes traces of a series of color spots at
suture and periphery.
Distribution: Halntat: Near Montpellier this species
occurred in an assemblage including barnacles, oysters,
and the potamidid gastropods Mesohalina, Potamides,
and Terebralia, together with driftwood, in a marl free
of sand grains, suggesting a sheltered, muddy, possibly
slightly brackish, intertidal, or lagoonal environment (D.
Kadolsky, personal communication).
Range: Loire (Middle Eocene), Adour (Middle and Up-
per Oligocene), Languedoc (Upper Oligocene), and
Aquitaine (Lower Miocene) of France.
Records: France: Le Bois-Gouet, Loire Atlantique, Biar-
ritzian (J. Le Renard Colin); Gaas, Adour Basin, Middle
Oligocene (MNHNP); Peyrere, Adour Basin, Chattian,
Upper Oligocene (Lozouet, 1986); La Paillade, near
Montpellier, latest Chattian (D. Kadolsky Colin); La Sau-
botte, Noaillan, Aquitanian (MNHNP).
Remarks: The nomenclature of this species is complex.
It was incorrectly listed as Trochus thorinus Grateloup
by Benoist (1874b), according to Cossmann and Peyrot
(1917), who had examined both the Grateloup and Beno-
ist Collections. The original description (Grateloup, 1832)
and figure (Grateloup, 1847) of Trochus thorinus were
both poor, but Cossmann and Peyrot (1917:280-282) ex-
amined the holotype and assigned the species (as a syn-
onym of Trochus biangulatm Eichwald) to the genus
Gihlmla (Trochidae); furtlii-rmort', their figure bears no
resemblance to Peasiella. The identity of Trochus heli-
cinus Grateloup, 1832 (not Gmelin; renamed subheli-
cinus d'Orbigny, 1852) is uncertain; in the absence of
type material in the Grateloup Collection, Cossmann and
Peyrot (1917) suggested that it was a synonym of Trochus
thorinus.
From the original description (Benoist, 1874a) it is
clear that Trochus risella (renamed Risella girondica by
Benoist, 1874b) is not a Peasiella, it was described as 15
mm in diameter, imperforate, with concave whorls and
tubercular sculpture. Xenophora rhytida. described and
figured b\ Cossmann (1899), bears a superficial resem-
blance to a Peasiella, but the scalloped or spiked margin,
projecting at the suture, the dorsal sculpture of 3 faint
cords, and the narrow umbilicus, are all unlike the pres-
ent species, and suggest that it may not be a littorinid.
The specimens examined during the present stud\ bear
a close resemblance to the original description and figures
of Pseudonina reyti by Cossmann and Peyrot (1917),
with the exception that the umbilicus is wider in the
former, but this character is somewhat variable in living
Peasiella species.
This species has a long stratigraphic range, from the
Middle Eocene to the Lower Miocene, but from the
material examined there seems no reason for subdivision.
It is compared with P. pontileviensis below. In habitat,
P. reyti may have resembled the Recent P. lutulenta.
Peasiella pontileviensis
(Morgan, 1915)
(figure 116)
Xenophora ? (Haliphaehua) pontileviensis Morgan, 1915:231-
232, figs. 14a-d [type not seen; N'allon de Charenton, Pont-
Levoy, Faiunian],
Circuhis pontileviensis. — Gilbert, 1949:71, pi. 5, fig. 1.
Torntis orthczensis Cossmann & Peyrot, 1918:27-28, pi. 7, figs.
39-41 [type not seen; Orthez, Aquitaine, Helvetian],
C.irculus orthczensis. — Gilbert, 1949:71,
Ri.'iella girondica. — Cossmann & Peyrot, 1919:437-438, pi. 17,
figs, 47, 48, 58 [not Benoist, 1874].
Shell (figure 116): Dimensions: 3.1-3.7 mm diame-
ter; height/diameter ratio 0.548-0.568 (n = 3, from 1
locality).
Shape: Depressed; whorls lightK rounded; base convex.
Strong flange at periphery, sometimes slightly undulat-
ing. Moderate umbilicus. Columellar pillar thickened
and angled at base.
Sculpture: Spire whorls of teleoconch smooth; 6-8 spiral
grooves and fine microstriae above periphery on last
whorl; no radial sculpture. Margin of umbilicus thick-
ened, but not outlined by a groove; 3-4 ribs on outer
half radius of base.
Color: Tw o specimens have 7 brown spots at suture and
12-20 smaller spots at periphery of penultimate whorl,
but little color on last whorl. A similar pattern covers the
last 2 whorls in fig. 14d of Morgan (1915).
Distribution: Range: Loire (Middle Miocene) and Aqui-
taine (Lower and Upper Miocene) of France.
Records: France: La Brede, Aquitaine, Aquitainian
(Cossmann & Peyrot, 1919); Orthez, Aquitaine, Helve-
D. G. Reid, 1989
Page 67
tian (Cossmanii & Peyrot, 1918); Ferriere-Lar9on, Loire,
Middle Miocene (MNHNP; Gilbert, 1949).
Remarks: This species is similar to P. reyti, but appears
to differ consistently in the presence of coarser spiral
grooves above the periphery on the last whorl (grooves
are absent or finer and more numerous in P. reyti), in
the markedly thickened columellar base and in the pres-
ence of ribs on the outer part of the base. Elsewhere in
the genus a thickened columellar base is found only in
the Recent species P. lutiilenta, from which P. pontilevi-
ensis differs in its smooth spire whorls and presence of
ribs on the outer part of the base only.
The three specimens examined showed 3 or 4 ribs on
the base; in the original description there were said to
be 2 (Morgan, 1915), and in Tornus orthezensis 5 (Coss-
mann & Peyrot, 1918). This range of variation is com-
parable to that of Recent species such as P. roepstorffiana.
Tornus orthezensis was said to lack spiral sculpture on
the dorsal surface (Cossmann & Peyrot, 1918), yet as
noted by Glibert (1949) the accompanying figure of the
holotype apparently shows faint spiral grooves.
This species appears to be generally rare; the authors
quoted in the synonymy (with the exception of Morgan,
1915) recorded a total of only 9 specimens. However,
Morgan (1915) recorded it as abundant at the type lo-
cality.
Peasiella aff. roepstorffiana
(NeviU, 1885)
(figure 117)
Shell (figure 117): Dimensions: 3.0 mm diameter;
height/diameter ratio 0.767 (n = 1).
Shape: Outline slightly convex; whorls almost flat-sided;
base slightly concave. Periphery sharply keeled, edge
straight; suture indistinct. Umbilicus almost closed. Col-
umellar pillar narrow, concave, rounded at base.
Sculpture: 6 strong, equidistant, spiral grooves above
periphery. No radial sculpture. Single strong rib on outer
part of base, with trace of another just within.
Distribution: Record: Shell Gully, Chatton, Okapua
Valley, near Gore, Southland, New Zealand, Duntrooni-
an. Upper Oligocene (NZGS).
Remarks: This single specimen from the Upper Oli-
gocene of New Zealand falls just within the range of
variation of the Recent species P. roepstorffiana. How-
ever, the single strong basal rib and very narrow um-
bilicus are atypical of that species, and until more spec-
imens are available, identification is tentative.
ACKNOWLEDGEMENTS
This work was partly carried out during the tenure of a
post-doctoral research fellowship at the Smithsonian In-
stitution, Washington, DC, and I should like to thank all
the staff there, particularly R. S. Houbrick. For loans of
specimens, 1 thank K. Way (BMNH), 1. Loch (AMS), P.
Bouchet (MNHNP), F. E. Wells and G. W. Kendrick
(WAM), N. V. Subba Rao (ZSI), R. N. Kilburn (NM), R.
Kilias (ZMB), G. Arbocco (MGD), and A. G. Beu (NZGS).
In addition, specimens were kindly provided by R. S.
Houbrick, S. Kool, T. Habe, J. Trondle, J. Dyson, A.
Mitchell, G. J. Vermeij and K. R. Smythe. For assistance
with fossil species I am most grateful to P. Lozouet
(MNHNP), J. Le Renard, and D. Kadolsky. The scanning
electron microscopes of the USNM were expertly oper-
ated by S. Braden and H. Wolf, and L. Cullen gave
assistance with histology. Light photography was by G.
N. G. Summons (BMNH).
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THE NAUTILUS 103(2):70-72, 1989
Page 70
A New Species of Coluzea (Gastropoda: Turbinellidae)
from off Southeastern Africa
M. G. Harasewych
Dfpartment of Invertebrate Zoology
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560, USA
ABSTRACT
Coluzea juliae, a new species of Columbariinae, is described
from bath) al depths oil northeastern South Africa and southern
Mozambique. It appears to be most close!) related to Coluzea
liriope Harasewych. 1986, from Makassar Strait in Indonesia.
Key words: Gastropoda; Turbinellidae; Columbariinae; Co-
luzea; Mozambique; South Africa; Indian Ocean.
INTRODUCTION
Although the genus Coluzea Allen, 1926 is represented
in Middle-Eocene shallow-water fossil beds of Europe,
post-Eocene records are restricted to deep water fades
of New Zealand. In the Recent fauna it is restricted to
bathyal depths along continental margins of the Indian
and southwestern Pacific Oceans. The southern coast of
Africa contains the richest and most diverse Recent col-
umbariine fauna known to date, to which is added the
new species described herein.
Repositories of examined specimens are indicated by
the following abbreviations:
DMNH — Delaware Museum of Natural History, Wil-
mington
NM — Natal Museum, Pietermaritzburg
SAM — South African Museum, Cape Town
USNM — National Museum of Natural History, Smith-
sonian Institution, Washington, DC
SYSTEMATICS
Genus Coluzea Allen, 1926
Coluzea juliae new species
(figures 1-4; table 1)
Description: Shell (figures 1-3) large for genus (to 89
nitn), thin, fusiform. Spire angle 37.0°-44.5°. Protoconch
deviated paucispiral, of 1% smooth, bulbous whorls.
Transition to teleoconch gradual, marked by develop-
ment of peripheral keel and broad axial ribs within Vz
whorl. Teleoconch with up to 9 whorls. Suture abutting
pronounced spiral cord anterior to w horl periphery. Spi-
ral sculpture of 3 equal threads or cords between suture
and periphery on early whorls. Cord nearest suture de-
creasing in prominence or disappearing b\ fifth post-
nuclear whorl; second and usually third spiral cords con-
comitantly thickening, each forming broad band. Second
band sufficientK elevated to form channeled suture. Two
strongly pronounced spiral cords between w horl periph-
ery and siphonal canal. Fourteen to 22 spiral cords along
proximal % of siphonal canal, decreasing in prominence
distally. Early whorls with 11-13 axial ribs/ whorl, each
forming tubercle along peripher\. Axial ribs becoming
less pronounced and tubercles forming short, open, an-
teriorly-directed spines on third post-nuclear whorl. Body
whorl with 15-16 spines. Aperture roughly semi-circular.
Outer lip thin, furrowed beneath periphery and major
cords. Inner lip smooth due to dissolution of portion of
outermost layer from parietal region. Siphonal canal long,
stout, axial, occasionally sinuate distalK (figure 3). Shell
color uniformly white. Periostracum thick, straw-colored
to tan, axially-bladed. Operculum (figure 4) corneous,
strongly ovate, with two straight sides converging on
terminal nucleus. Soft parts unknown.
Etymology: This species is named in honor of my wife
Julia, in recognition of her frequent assistance and sup-
port of my research.
Type locality: Off Inhaca Island, Mozambique, 26°07'S,
34°11'E, in 600-665 m, Anton Bruun Cruise 8, station
397C, September 29, 1964.
Holotype: USNM 718510, length 67.9 mm.
Paratypes: Paratypes 1-2, USNM 860174, from the type
localitv; paratvpe 3, SAM A36257, off northern Natal,
South Africa, '27°12.2'S, 32°56.0'E, in 660 in. May 20,
1976; paratypes 4-6, SAM A36256 off northern Natal,
South Africa, 27°14.8'S, 32°54.6'E, in 680-700 m. May
20, 1976; paratypes 7-8, DMNH 19068, trawled, deep
water off Ziiluland coast. South Africa, December, 1967;
paratypes 9-11, NM 5740, off Natal, South Africa, May,
1970; paratype 12, Gratz collection, off Natal, South Af-
rica, 200-300 fms [366-548 m].
M. G. Harasewych, 1989
Page 71
Figures 1-4. Coluzea juliae new species. 1. Holotype, USNM 718510, off Inhaca Island, Mozambique, 26°07'S, 34°irE, in 600-
665 m, Anton Bruun Cruise 8, station 397C, September 29, 1964 (1.0 x ). 2. Paratype 3, SAM A36257, off northern Natal, South
Africa, 27°12.2'S, 32°56.0'E, in 660 m. May 20, 1976 (1.0 x ). 3. Paratype 9, NM 5740, off Natal, South Africa, May, 1970 (1.0 x ).
4. Operculum of holotype (2.5 x ).
Distribution: This species is known only from the upper
continental slope off southernmost Mozambique and
northeastern Republic of South Africa. The confirmed
bathymetric range is 548-660 m.
Comparative remarks: Coluzea juliae new species most
closely resembles C. liriope Harasewych, 1986, a species
inhabiting similar depths along the Makassar Strait of
Indonesia, but differs in having: a periphery that is sharp-
ly keeled and lined with long, narrow, open, anteriorly-
directed spines rather than a rounded periphery with
low nodular spines; two extremely pronounced raised
cords between the periphery and siphonal canal, rather
that four weaker cords; and a crenulated outer lip rather
than one with a rectangular posterior margin. This new
species is also similar to Coluzea eastwoodae (Kilburn,
1971), with which it overlaps in geographic and bathy-
metric ranges, but can be readily distinguished on the
basis of its characteristic spiral sculpture between the
suture and periphery.
Discussion: Of the 13 specimens of Coluzea juliae new
species examined, 10 had repaired breaks along the prox-
imal portion of the siphonal canal, indicating crab pre-
Table 1. Coluzea juliae new species. Measurements of shell characters. Linear measurements in mm (n = 10, e.xcept for mea-
surements involving the siphonal canal, where n = 7).
Character
Mean
Range
SD
Shell length (apex to proximal end of
siphonal canal)
Aperture length
Aperture length shell length
Siphonal canal length
Siphonal canal length/aperture length
no. whorls teleoconch
no. axial ribs/spines on body whorl
Spire angle
41.3
13.8
0.33
34.5
2.82
8.6
16.6
41.5°
33.1-51.3
10.6-17.3
0.31-0.36
28.2-45.8
2.44-3.34
8.0-9.0
14-23
37.0°-44..5°
6.0
2.3
0.02
6.7
0.31
0.4
2.7
2.2°
Page 72
THE NAUTILUS, Vol. 103, No. 2
dation. The thick spiral cords characteristic of this species
strengthen the shell and increase its resistance to crushing
and peeling predators. These expanded cords also result
in an altered apertural shape that accommodates a taller,
more voluminous mantle cavity, especialK in the regions
that contain the h> pobranchial gland, pallial gonoducts
and rectum Similar modifications of apertural shape oc-
cur in Coluzea liriope (Harasewvch, 1986; pi. 2, figs. 3,
4), and to a lesser extent in Coluzea cingulata (Martens,
1901) (see Darragh, 1969: pi. 6, figs. 106, 1 16) and in the
early whorls of C. diatephanotis (Melvill, 1891) (see Har-
asewych, 1986: pi. 1, figs. 1-3). The similarities between
Coluzea juliae new species and C. liriope are sufficiently
compelling to suggest a close relationship between these
two species. The relationships of these species to C. cin-
gulata and C. distephanotis are less clear. The dorsal
expansion of the aperture in the early whorls of C. diste-
phanotis occurs anterior to the periphery, and may be
a convergent adaptation to increase the volume of the
mantle cavity.
LITERATURE CITED
Darragh, T \. 1969. A revision of the family Columbariidae
(Mollusca: Gastropoda). Proceedings of the Roval Society
of Victoria 83(11:63-1 19.
Harasew\th, M G. 1986. The Columbariinae iGastropoda:
Turbinellidae) of the eastern Indian Ocean Journal of the
Malacological Societ\ of .Australia 7(3-4): 15.5- 170.
Kilburn, R. N. 1971. On some species of the families Ton-
nidae, Hipponicidae. Buccinidae, Columbariidae, Fascio-
lariidae, Psammobiidae and Vlactridae (.Mollusca) in South
African waters. Annals of the Natal Museum 20(3):483-
497.
von Martens, E. 1901. Neue Meer-Conchylien von der
deutschen Tiefsee-Expedition Sitzungsberichte der Ge-
sellschaft naturforschender Freunde zu Berlin 1901 14-
26,
Melvill, J, C. 1891. Descriptions of eleven new species be-
longing to the genera Columbarium, Pisania, Minolia.
Liotia, and Solarium. Journal of Conchology 6(12):405-
411, pi. 2.
THE NAUTILUS 103(2):73-77, 1989
Page 73
Genetics and Shell Morphology of Hard Clams
(Genus Mercenaria) from Laguna Madre, Texas*
Robert T. Dillon. Jr.
Department of Biology
College of Charleston
Charleston, SC 29424, USA
John J. IVlanzi
South Carolina Marine Resources
Research Institute
PO Box 12559
Charleston, SC 29412, USA
ABSTRACT
Texas Mercenaria were originally described as a subspecies of
M. campechiensis. but are now generally regarded as a sub-
species of M. nxercenaria. primariK' based on aspects of shell
ridging. We used isoz> me frequencies at seven loci, six mor-
phometric variables, shell ridging, and nacre color to compare
Texas populations to reference populations of M. campechien-
sis and M. mercenaria. Texas populations were indeed distinct,
but much more similar to the former. Hard clams from Texas
should be considered Mercenaria campechiensis texana (Dall,
1902).
Key words: Hard clams; Texas; electrophoresis; morphomet-
INTRODUCTION
Venerid clams of the genus Mercenaria (variously known
as quahogs, cherr\stones, hard clams, etc.) are of such
commercial importance that it is surprising their system-
atic relationships are not better understood. Most authors
follow Abbott (1974) in recognizing two North American
species, the northern Mercenaria mercenaria (Linne,
1758) and southern Mercenaria campechiensis (Gmelin,
1791). Three criteria have been used to distinguish the
species. Mercenaria campechiensis is supposed to have
thick concentric ribs, white nacre, and a lunule at least
as wide as it is high, while M. mercenaria has thin, easily
eroded ribs, purple nacre, and a narrower lunule (figure
1).
.\bbott (1974:523) noted that M. campechiensis hy-
bridizes with M. mercenaria in the wild, and "could well
be considered a subspecies. It has recently been shown,
however, that some reproductive isolation exists between
the two species where they occur sympatrically in the
Indian River, Florida (Dillon & Manzi, 1989). Thus we
consider these species distinct.
* Contribution No. 266 from the SC Marine Resources Cen-
ter.
Dillon and Manzi (1989) selected one population each
from central portions of the ranges of M. mercenaria
and M. campechiensis to serve as "references". These
populations appeared to be typical both genetically and
morphologically, with no evidence that either contained
any hybrid genomes. We found that 100?c of the M.
mercenaria shells had thin, easily eroded concentric ribs,
while over 995t of M. campechiensis had thick, resistant
concentric ribs. Nacre color was also a useful discrimi-
nator— 80% of the M. mercenaria had distinct purple
color, while 92% of M. campechiensis were completely
white. The ratio of lunule width to lunule height proved
to be of limited utility. Over 86% of M. campechiensis
in our sample had ratios less than 1.0, and thus would
have been misclassified as M. mercenaria using this tra-
ditional criterion. We did find, however, that if measures
of lunule height and width were combined with overall
shell length, width, height, and weight, very accurate
morphometric discrimination between the two species
was possible (Dillon & Manzi, 1989).
Hard clam populations inhabiting the Texas coast of
the Gulf of Mexico were originally described by Dall
(1902) as Venus (now Mercenaria) campechiensis tex-
ana. Dall viewed the presence of thin, easily eroded
concentric ribs in the Texas populations (figure 1) as
justification for recognizing the subspecies. The subspe-
cies was transferred to M. mercenaria by .\bbott (1954),
at least partly because the inshore fauna of the Gulf of
Mexico is generally Carolinian, rather than Caribbean
in affinity (personal communication to Joy Goodsell). Here
we show that based on isoz\me frequencies and all other
shell characteristics besides ridging, texana is a subspe-
cies of M. campechiensis as originally described, not M.
mercenaria.
METHODS
We were able to obtain 29 Mercenaria of the subspecies
texana from Laguna Madre, in the vicinity of Corpus
Christi, Texas. Samples were taken of both siphon and
foot tissue, and electrophoretic analysis performed as
described elsewhere (Dillon, 1982, 1985; Dillon & Manzi,
Page 74
THE NAUTILUS, Vol. 103, No. 2
Figure 1. Left and anterior aspects of tfie three Mercenaria
taxa. Left — M. campechiensis (Gmelin, 1791), Center — the
subspecies M. campechiensis texana (Dall, 1902), Right — M.
mercenaria (Linne, 1758) The scale bar is 50 mm.
1987). We estimated allele frequencies at the same seven
enzyme loci that have been examined previously: glucose
phosphate isomerase (GPI), leucine aminopeptidase
(LAP), superoxide dismutase (SOD), 6-phosphogluconate
dehydrogenase (6PGD), mannose phosphate isomerase
(MPI), and phosphoglucomutase (two loci — PGMS and
PGMF). Simple Mendelian inheritance of codominant
alleles has been demonstrated at GPL LAP, 6PGD, PGMS,
and PGMF by Adamkewicz et al. (1984).
Gene frequencies at individual loci were compared
using chi-square tests for two independent samples, cor-
rected for continuity in 2 x 2 cases. Alleles with expected
frequencies less than 5 were combined with other rare
classes if possible, otherwise they were eliminated. The
genetic distance over all 7 loci between each pair of
populations was calculated using the method of Nei
(1972).
Six measurements were made on the shells of most
individuals. Maximum shell length, shell height (maxi-
mum dimension in the plane of symmetry perpendicular
to shell length), shell width (maximum dimension per-
pendicular to the plane of symmetry), lunule height (along
the line separating the valves), and lunule width (the
maximum dimension perpendicular to lunule height)
were measured with vernier calipers. The weight of both
valves combined was recorded to the nearest gram. Nacre
Table 1 . .■\llele frequencies at seven enzyme loci for clams of
the subspecies texana compared to reference populations of M.
mercenaria and M. campechiensis (data of Dillon & Manzi,
1989). Sample sizes were approximately 29 texana, 194 M,
campechiensis, and 224 M. mercenaria.
M M.
.Allele mercenaria texana campechiensis
GPIUO
0.023
0.0
0 0
105
0.014
0.0
0.0
100
0.901
0.0
0.0
90
0.021
0.0
0.008
85
0.0
0.034
0.023
80
0.002
0.310
0.221
70
0.038
0.414
0.432
65
0.0
0.0
0.013
60
0.0
0.190
0.271
50
0.0
0.052
0.031
LAP 104
0.095
0.063
0.074
100
0.412
0.042
0.127
96
0.46;3
0.521
0.333
94
0.019
0.354
0.407
90
0.012
0.021
0.059
SOD 100
0.768
0.0
0.018
90
0.232
0.609
0.702
80
0.0
0.391
0.281
6PGD 110
0.030
0.077
0.084
100
0.622
0.481
0.517
95
0.0
0.0
0.011
90
0.348
0.442
0.388
MPIllO
0.0
0.0
0.032
108
0.059
0.370
0.484
105
0.389
0.304
0.267
100
0,300
0.304
0.174
95
0.253
0.022
0.043
PGMS 103
0.012
0.0
0.0
100
0.844
0.060
0.161
97
0.043
0.040
0.078
95
0.0
0.0
0.075
92
0.077
0.860
0.578
87
0.024
0.040
0.056
82
0.0
0.0
0.035
77
0.0
0.0
0.016
PGMF 103
0.148
0.021
0.061
100
0.852
0.417
0.282
97
0.0
0.562
0.636
95
0.0
0.0
0.021
color and strength of concentric ridges were also noted.
Five whole individuals and two single-valves were lost
subsequent to tissue sampling. Thus sample sizes were N
= 29 for isozyme frequencies, but only N = 24 for the
morphological analyses.
We compared the Texas clams to the reference pop-
ulations of 224 M. vwrcenaria and 194 M. campechiensis
analyzed by Dillon and Manzi (1989). The M. mercenar-
ia were sampled from a tributary of the Stono River, 15
km south of Charleston, South Carolina. The M. cam-
pechiensis were collected at Cedar Key, on the north-
R. T. Dillon, Jr. and J. J. Manzi, 1989
Page 75
Table 2. Results of the principal component analysis of shell morphometric data from Mercenaria mercenaria, M. campechiensis,
and tlie subspecies texana.
Morphological
E:
igenvectors
character
PCI
PC2
PC3
PC4
PC5
PC6
Shell length
0.42
0.16
-0.43
-0.59
0.41
0.32
Shell height
0.44
-0.08
-0.25
-0.15
-0.27
-0.80
Shell width
0.42
-0.33
-0.10
0.10
-0.66
0.50
Lunule width
0.40
-0.05
0.85
-0.30
0.09
-0.02
Lunule height
0.34
0.84
0.05
0.40
-0.10
0.05
Shell weight
0.41
-0.39
-0.06
0.61
0.55
-0.01
Eigenvalue
4.96
0.56
0.25
0.11
0.08
0.03
Cumulative variance
0.83
0.92
0.96
0.98
0.99
1 00
central Gulf coast of Florida. All shells examined in this
study have been deposited at the Academy of Natural
Sciences of Philadelphia. Catalog numbers are as follows:
Texas population 373466, M. campechiensis 373467, M.
mercenaria 373468.
Following Dillon and Manzi (1989), we performed a
discriminant analysis on principal component scores ex-
tracted from the six measurement variables. First a prin-
cipal component analysis was performed on the corre-
lation matrix calculated over all 442 individuals (the
Princomp procedure, SAS, 1985). We disregarded vari-
ance on the first principal component (PC) as a method
of factoring out size variance, and used factor scores on
the remaining 5 PC s as new variables for nearest-neigh-
bor discriminant analysis (the Neighbor procedure, SAS,
1985). This is a nonparametric discriminant analysis, not
involving the calculation of discriminant functions. In
our application there were 418 known clams, and only
the 24 Texas clams were entered as unknowns. Each clam
was classified as M. mercenaria if at least 19 of its 20
nearest Euclidean neighbors of known affinity were M.
mercenaria, M. campechiensis if 19 of 20 were M. cam-
pechiensis, and intermediate if otherwise.
RESULTS
Table 1 compares allele frequencies at seven enzyme loci
in the Texas clams to reference frequencies established
for M. mercenaria and M. campechiensis by Dillon and
Manzi (1989). The two reference populations are strik-
ingly distinct at GPI, SOD, MPI, and PGMF, and in
these four cases, the texana sample is not significantly
different from M. campechiensis by chi-square. The two
reference populations are also distinct at the LAP and
PGMS loci, but although the texana sample is much more
similar to M. campechiensis, significant differences exist.
The frequency of LAP 100 is significantly lower in tex-
ana (chi-square = 8.01, 2 d.f), and there seems to have
been a significant loss of allelic diversity at the PGMS
locus (chi-square = 13.5, 1 d.f). The Texas population
was not significantly different from either reference pop-
ulation at the 6PGD locus.
Calculated over all 7 polymorphic loci, Nei's genetic
distance (D) between the two reference populations was
0.82. The Texas population showed D = 0.041 to M.
campechiensis but D = 0.83 to M. mercenaria.
Results of the principal component analysis on shell
morphometries are given in table 2. Factor loadings were
somewhat different from those obtained by Dillon and
Manzi (1989), since 24 clams of the subspecies texana
have replaced 170 individuals from the Florida hybrid
zone. We discarded PCI, representing 83% of the vari-
ance, and used the remaining 17% for discriminant anal-
ysis.
Figure 2 shows that the two reference populations are
quite distinct on PC2, even though this is not a discrim-
inant function, with M. campechiensis scoring lower.
Judging from the factor loadings on PC2 (table 2), M.
campechiensis would seem to have a wider, heavier shell
than M. mercenaria. In contrast to our previous findings,
lunule height loads very strongly on PC2, while the con-
tribution of lunule width is negligible. It would appear
that M . campechiensis does not have an especially wide
lunule for its size, but rather a distinctively short (or
'"lower") one. Nearest-neighbor discriminant analysis
confirmed that the two reference populations are very
distinct (table 3). One M. campechiensis was misclassi-
fied, to 95% confidence, as M. mercenaria, but no M.
mercenaria were misclassified. The reference popula-
tions were both about 80% distinct.
Although more similar to M. campechiensis than M.
mercenaria, Texas clams were quite diverse morphomet-
rically (figure 2). The three lowest PC2 scores all be-
longed to texana specimens, suggesting that Texas pop-
Table 3. Classification (to 95% confidence) of clams by nearest-
neighbor discriminant analysis on principal component scores,
given the two reference populations as knowns and specimens
of the subspecies texana as unknowns.
M. mer- Inter- M. cam-
cenaria mediate pechiensis
M. mercenaria reference
texana
M. campechiensis reference
180
44
0
2
14
8
1
37
156
Page 76
THE NAUTILUS, Vol. 103, No. 2
1.8
1.5
1.2
n 09
LU 0.6
z
o
I 0.3
o
^ 0.0
i-0 3
z
I -0 6 -
-0.9
-1.2 I-
-1.8 -
oat, o
8o
o o
••^ A A
O A A OA A O AA
O AA ^^
A A
A
A AA
AAA
AAA
A A
o o
o o
A
O A O O
• o o
O O A O AAA A A A
O O O A •gOA* AA O A ^ A A
OO O OO O OO AO A AAA A AA A AA
oo OO o aao o o a aaaaaa*
O O OOtO O O OOOA iVJO IWSA AAA AAA i
O O OO O • A O^ AA AAAA AAA
O OOOO 0« O A OO AAA A A AA AAAA
OOOO OOOOOOO AOAA AAA A A
OO OO O O O A AA AAA
OOOO O OOOOAA AAA i
O 09 A O O A A ^lA A
CC
oi o o
CO
o oo OO
O O O A OO
O
O
o
o
o o
o
Oe£,
oo
AA AA AAA A
-3.0
-1.8
06
18
PRINCIPAL COMPONENT 2
Figure 2. Factor scores on principal components 2 and 3. Triangles are M. mercenaria reference, open circles are M. campechiensis
reference, and closed circles are M. campechiensis texana. A total of 51 reference individuals are obscured by o\erlap.
ulations may be distinguished by greater relative width
and weight than reference M. campechiensis. and by
even shorter lunules. Specimens of the subspecies texana
also tended to be distinct on PCS, showing wider lunules
and shorter shells overall. Table 3 shows that most shells
from the Texas population could not be identified, to
95% confidence, as coming from either reference pop-
ulation. Among classifiable shells, however, those indis-
tinguishable from M. campechiensis outnumbered those
from M. mercenaria by a ratio of 4 to 1.
All individuals from the Texas population showed the
typical M. campechiensis trait of purely white nacre.
But the striking feature of the texana shells was the
presence of thin, easily eroded concentric ridges or ribs.
Ribs were eroded to leave bald patches on all 24 indi-
viduals examined, even though over 99% of the reference
M. campechiensis population had strong, resistant ribs.
DISCUSSION
Isozyme frequencies clearly show that the Texas popu-
lations are much more similar to M. campechiensis than
M. mercenaria. Considering overall genetic distance, it
was in fact the reference M. campechiensis population
that was intermediate, not the Texas population. It is
difficult to compare our values of D to those collected
from other taxa, since monomorphic loci were excluded
from this study. But it appears that isozyme divergence
between both species and subspecies of Mercenaria is
unusually low (Avise, 1976).
It might be argued that the geographic distance be-
tween Texas and South Carolina populations makes a
comparison of isozyme frequencies unfair. But extensive
dispersal is apparently possible during Mercenaria' s ve-
liger stage. Dillon and Manzi (1987) reported only a
single significant difference at these se\en loci in a com-
parison of Massachusetts and X'irginia M. mercenaria.
Onh two significant differences were apparent between
X'irginia and South Carolina, and the approximately 20
clams from the Atlantic coast of Florida identified as
pure M. mercenaria were not strikingK different from
South Carolina populations (Dillon & Manzi, 1989). It
seems unlikely that a difference of the magnitude re-
ported here between the reference M. mercenaria and
the Texas clams could be due to distance alone. The
minor differences shown at two loci between the Texas
population and the reference M. campechiensis popu-
lation from northern Gulf Florida are of the magnitude
we have observed from isolation b\ distance.
The reference populations were quite distinct in shell
morphometries, and again individuals of the texana sub-
species tended to sort out with M. campechiensis. The
nacre color of the Texas clams also clearly places them
with M. campechiensis. But the presence of some pe-
culiarities of shell shape, together with thin, easily eroded
concentric ridges, makes Texas populations so distinct
that the\ do warrant recognition as a subspecies, Mer-
cenaria campechiensis texana (Dall, 1902).
Preliminary results from hybridization studies be-
tween standard M. mercenaria and M. campechiensis
conducted in our facilities suggest that shell ridge thick-
ness is primariK , perhaps entirely, under genetic control.
The ridge thickness of F, h\brids (measured by me-
chanical filing) is intermediate between that of pure off-
spring from the two species spawned at the same time
and reared in the same environment. The thinner, finer
R. T. Dillon, Jr. and J. J. Manzi, 1989
Page 77
ribs shared by M- mercenaria and M. campechiensis
texana may be an adaptation for burrowing in the fine,
terrigenous silt and mud found in the estuaries of the
American Atlantic and northern Gulf coasts. The thicker,
heavier ribs of typical M. cavipechiensis may be an
adaptation for the coarser, carbonate sands offshore, in
peninsular Florida, and the Caribbean Sea. Thin ridges
are probabh' ancestral, with thicker ridges evolving after
the divergence of M. mercenaria and A/, campechiensis.
Otherwise, one would need to postulate that thin ridges
evolved separately in M. mercenaria and M. campe-
chiensis texana.
Some attention has focused on M. campechiensis tex-
ana as a candidate for mariculture, especially in the
Texas environment to which it is adapted (Craig et al.,
1988). Another possible source of commercial interest is
in the hybridization of Texas populations to M. merce-
naria as a method of increasing genetic variability (re-
view by Dillon & Manzi, 1988). Recently Goodsell (1989)
has made all reciprocal crosses between M. mercenaria,
M. campechiensis, and M. campechiensis texana, dem-
onstrating the feasibility of this approach. In any such
future studies, the genetic relationships among these three
taxa should be kept in mind.
ACKNOWLEDGEMENTS
We thank David McLean, John Wise, and Nancy Hadley
for their technical assistance. Joy Goodsell helped with
sampling and the literature search. Mercenaria campe-
chiensis texana were provided by Tom Bright, Texas Sea
Grant. This work was sponsored by NOAA, National Sea
Grant College Program Office, US Department of Com-
merce under Grant No. NA85-AA-D-SG121, and the
South Carolina Sea Grant Consortium.
LITERATURE CITED
Abbott, R. T. 1954.
York, 541 p.
American seashells. Van Nostrand, New
Abbott, R. T. 1974. American seashells, 2nd ed. Van Nostrand
Reinhold, New York, 663 p.
Adamkewicz, L., S. R. Taub, and J. R. Wall. 1984. Genetics
of the clam Mercenaria mercenaria. I. Mendelian inher-
itance of allozyme variation. Biochemical Genetics 22:215-
219.
Avise, J. C. 1976. Genetic differentiation during speciation.
In: Avala, F. J. (ed.). Molecular evolution. Sinauer, Sun-
derland, MA, p. 106-122.
Craig, M. A., T. J. Bright, and S. R. Gittings. 1988. Growth
of Mercenaria mercenaria and Mercenaria mercenaria
texana seed clams planted in two Texas bays. Aquaculture
71:193-207.
Dall, W. H. 1902. Synopsis of the family Veneridae and the
North American recent species. Proceedings of the United
States National Museum 26:335-411.
Dillon, R. T, Jr. 1982. The correlates of divergence in isolated
populations of the freshwater snail, Goniobasis proxima.
Unpublished Ph.D. thesis, University of Pennsylvania,
Philadelphia, 183 p.
Dillon, R. T., Jr. 1985. Correspondence between the buffer
systems suitable for electrophoretic resolution of bivalve
and gastropod isozymes. Comparative Biochemistry and
Physiology 826:643-645.
Dillon, R. T, Jr. and J. J. Manzi. 1987. Hard clam, Mercenaria
mercenaria, broodstocks: genetic drift and loss of rare al-
leles without reduction in heterozygosity. Aquaculture,
Amsterdam 60:99-105.
Dillon, R. T, Jr. and J. J. Manzi. 1988. Enzyme heterozygosity
and growth rate in nursery populations of Mercenaria
mercenaria (L.). Journal of Experimental Marine Biology
and Ecology 116:79-86,
Dillon, R. T., Jr. and J. J. Manzi. 1989. Genetics and shell
morphology in a hybrid zone between the hard clams,
Mercenaria mercenaria and M. campechiensis. Marine
Biology 100:217-222.
Goodsell, J. G. 1989, Shell morphology, growth and survival
of larval and early post-larval Mercenaria and their hy-
brids. Unpublished M.S. thesis, Clemson University, Clem-
son, SC.
Nei, M. 1972. Genetic distance between populations, Amer-
ican Naturalist 106:283-292.
SAS Institute, Inc. 1985. SAS users guide, version 5 ed. SAS
Institute, Gary, NC, 956 p.
THE NAUTILUS 103(2):78-82, 1989
Page 78
Variation in Size Demography of Lotic Populations of
Corbicula fluminea (Miiller)
Barry S. Payne
Andrew C. Miller
Environmental Laboratory
US Army Engineer Waterways
Experiment Station
PO Box 631
Vicksburg, MS 39180
Paul D. Hartfield
Mississippi Museum of Natural Science
111 North Jefferson Street
Jackson, MS 39202
Robert F. McMahon
Department of Biology
University of Texas at Arlington
Arlington, TX 76019
ABSTRACT
Patterns of size demography among 14 lotic populations of the
Asiatic clam, Corlncida fluminea. in the state of Mississippi
were compared based on samples collected between May 24
and June 22, 1984. Individuals greater than 20 mm in shell
length (SL) and greater than 1 year old comprised a substantial
fraction (13-64%) of only six populations. These populations
had distinct multiple cohorts. Four of the six populations oc-
curred among longer-lived native unionids, suggesting that
streambed stabilit> is crucial to the establishment of lotic pop-
ulations of C. fluminea « ith complex size (and age) structure.
Individuals greater than 20 mm SL comprised a small fraction
(0-5%) of the other eight populations; these populations were
characterized by a heavily dominant, single cohort of small
clams. Unusually high stream discharge throughout the state
from December, 1982 through May, 1983 and again in De-
cember, 1983 exaggerated ordinarily high winter and spring
flows The simple size demography of the eight populations
with few large individuals probably reflected recolonization of
flood-decimated assemblages by small individuals transported
downstream from more stable upstream sites.
Key words: Corbicula fluminea; unionids; size demography;
lotic habitats; flood scour; streambed stability.
INTRODUCTION
The Asiatic clam, Corbicula fluminea. is clearly adapted
to well-oxygenated sand and gravel shoals of lotic habitats
and is intolerant of reducing sand, mud, or silt (Mc-
Mahon, 1983 and references within). Sand and gravel
shoals are often shifted b\ flood flows in lotic habitats.
As reviewed by McMahon (1983), C. fluminea shows
several adaptations to unstable substrata, including: a
massive and concentrically ridged shell to resist abrasion
and anchor individuals in substrata exposed to swift flow;
extensive mantle fusion, narrow pedal gape, a foramen
around the posterior adductor muscle, and highly de-
veloped statocysts providing for rapid locomotion and
burrowing; and hermaphroditism plus production of
crawling pediveliger larvae (also capable of byssal thread
attachment) allowing rapid establishment of dense pop-
ulations by just a few colonists. Despite larval adaptations
to enhance local recruitment, C. fluminea is passively
transported downstream in large numbers. Pediveligers
(e.g., Sickel, 1979) and post-larval juveniles less than 2
mm SL (Williams & McMahon, 1986) drift on water
currents. Furthermore, small individuals (up to 15 mm
but mostly less than 5 mm SL) are passively transported
downstream in large numbers, evidently by tumbling
along the bottom (Williams & McMahon, 1986). Passive
transport of individuals from viable upstream popula-
tions allows rapid recovery of downstream populations
that have been severely reduced in size or destroyed (e.g..
Cherry et ai, 1980; McMahon & Williams, 1986).
Despite extensive information on biological adapta-
tions of C. fluminea to lotic habitats, only one study has
been published detailing the life history of C. fluminea
in a lotic habitat in the southern United States (Sickel,
1979). Most detailed information about population dy-
namics of this species comes from studies of lentic pop-
ulations (McMahon, 1983 and references within). Cor-
bicula populations in lotic habitats are likely to be affected
by seasonal patterns of stream discharge affecting sta-
bility of sand and gravel shoals. The purpose of this paper
is to describe interpopulation variation in size demog-
raphy of C. fluminea at 14 lotic sites in Mississippi rep-
resenting a range of substrata stabilit\' conditions.
MATERIALS AND METHODS
A large number of individuals, ranging from 98 to 413,
were collected from 14 lotic populations at locations
throughout Mississippi (table 1). All samples were taken
by scooping or pushing substrate into a dip net with a
mesh size of 4.0 mm. The maximum shell length (SL,
the greatest anterior-posterior dimension across the
valves) of each individual was measured to the nearest
0.1 mm using a Vernier caliper, and SL frequency his-
tograms were plotted for each population. All samples
except one were collected within an 18 day period in
B. S. Payne et ai, 1989
Page 79
Table 1. Description of samples and habitats of 14 lotic populations of C. fluminea sampled in Mississippi during late spring and
early summer of 1984.
Site
Stream
Date
Sample Den-
size sity' Flow-
Substrate
BYP
Bavou Pierre
5 Jun
136
L
L
TAN
Tangipahoa R.
5 Jun
129
M
M
PAS
Pascagoula R.
21 Jun
202
H
N
LEA
Leaf R.
21 Jun
196
M
L
CHU
Chunkv R.
18 Jun
119
M
L
YOC
Yockanookanv R.
7 Jun
136
M
M
NOX
Noxubee R.
18 Jun
280
L
M
PRL
Pearl R
7 Jun
213
L
L
BBL
Big Black R.
19 Jun
98
L
L
STR
Strong R
24 Mav
145
L
H
BUT
Buttahatchie R.
18 Jun
102
L
M
CHI
Chickasawhav R.
21 Jun
413
H
N
TAL
Tallahala Cr.
21 Jun
192
M
M
FMC
Fourteen Mile Cr,
22 Jun
141
L
N
Along stumps and snags; with unionids
Sandy gravel along snags and macropytes; with unionids
Mud, sand, and silt; with unionids
Muddy sand; with unionids
Mud over bedrock; with pisidiids
Gravel where stabilized by stumps and snags
Unconsolidated riffles of mud, sand, gravel, and chalk flakes
Detritus and sand among roots of cypress trees
Sand bar in association with snags, limbs, and sticks
Pockets of detritus and sand in scour holes in bedrock
Gravel shoals
Mud, sand, and broken clay at end of long run
Sand and gravel stabilized by snags
Thick silt over sand and some gravel
' Density was estimated as high (H; >300 individuals/m^), moderate (M; 50-300 ind/m^), or low (L; <50 ind/m-).
-Flow velocity was estimated, during relatively low discharge at all sites, as high (H; >0.5 m/sec), moderate (M; 0.2-0.5
m/sec), low (L; detectable but <0.2 m/sec), or neglible (N).
late spring (June 5-22, 1984) to minimize the influence
of different dates of sampling on intersite variation in
size demography. One sample was collected slightly ear-
lier, on May 24. All samples were collected during rel-
atively low flow conditions.
Stream discharge in Mississippi responds to seasonal
patterns of rainfall, and is typically high in winter and
early spring and low during summer and early fall (figure
1, US Geological Survey, 1981; Tate et ai, 1982, 1983;
Tharpe et ai, 1984). Winter and spring discharge was
unusually high throughout the state from December,
1982 through mid- May, 1983 and in December, 1983.
High flows in 1983 broke historic records at two of the
14 sites (BBL and BYP) and approached record levels at
several other locations (Tate et ai, 1983). Ratios of ma,\-
imum-to-minimum daily discharge averaged 1,535 from
October, 1982 through September, 1983 for gage sites
near our sampling locations (Tate et ai, 1983), reflecting
the extreme shifts in hydrologic conditions characteristic
of these lotic habitats.
RESULTS
Individuals greater than 20 mm SL comprised a sub-
stantial fraction of six populations (figure 2). The percent
abundance of individuals greater than 20 mm SL equaled
12.5, 13.8, 13.9, 19.5, 46.5, and 63.9 at TAL, BYP, PAS,
LEA, TAN, and CHU, respectively. Two distinct cohorts
were evident at TAL, and the other five populations
included three or more cohorts (figure 2). Both the abun-
dance of individuals greater than 20 mm SL and the
complex size demography indicated longevity of 2 to 3
years for a substantial fraction of each population. These
age estimates are based on Sickel's (1979) detailed study
of a single southern lotic population in the Althahama
River in Georgia. Individuals in the Altahama River grew
to modal SL's of 14 and 22 mm in the first and second
year of life, respectively. The largest individual collected
(38 mm SL) was from the LEA population, and the CHU,
TAN, and PAS populations also included individuals
greater than 30 mm SL.
Four of the six populations that included a substantial
number of 2 and 3 year old individuals (TAN, PAS, LEA,
and BYP) occurred with adult unionacean bivalves (table
1). The most common unionids were Quadrula pustulosa
(Lea), Fusconaia flava (Rafinesque), and Pluerobema
beadleanum (Lea) at TAN and BYP. Unionids at PAS
were dominated by Lampsilis teres (Rafinesque) and Q.
pustulosa. Dominant species at LEA were Toxolasma
texasensis (Lea) and L. teres.
Eight populations were heavily dominated by a single
cohort of individuals (1983 recruits) less than 20 mm SL
(figure 3). The percent abundance of C. fluminea greater
than 20 mm SL among these eight populations was sig-
nificantly less (t = 3.53; p < 0.01) than among the other
six populations. No individuals greater than 20 mm SL
were collected at YOC or NOX. Percent abundance of
clams greater than 20 mm SL equaled 0.2, 0.5, 2.0, 2.1,
2.8, and 4.9 at CHI, PRL, BBL, STR, FMC, and BUT,
respectively. The paucity of large individuals in these
eight populations did not allow cohorts other than the
1983 generation to be clearly distinguished (figure 3).
The average SL of the dominant 1983 generation ranged
among the eight populations from 8.8 mm at YOC to
15.9 mm at FMC. This wide range reflected intersite
differences in timing of recruitment and/or SL growth
rates. The dominant cohort of the FMC population was
considerably larger than the main cohort of the other
eight populations. This population was from a small
stream in a drainage of mostly agricultural land, while
all other populations were in larger streams or rivers that
drain primarily forested land. Thus, the FMC site was
Page 80
THE NAUTILUS, Vol. 103, No. 2
TOMBtOBEE n AT COtUMBUS
(NORTHEAST MiSStSSIPPlI
Figure 1 . Median monthly discharge recorded from October,
1980 through September, 1984 at three representative gage
stations in Mississippi (based on data in Tharpe et al. (1984),
Tate et al. (1982, 1983), and US Geological Survey (1981). The
dotted line shows the median monthly discharge averaged for
the period 1951-80.
a comparatively eutrophic habitat with sustained periods
of warm water temperature during winter (due to in-
creased solar insolation and smaller stream size). These
different habitat conditions at FMC probably accounted
for the higher than average SL growth of clams at this
site.
DISCUSSION
The general lack of large clams greater than 1 year in
age in Mississippi stream populations of C. fluminea is
unusual. Most populations of C. fluminea (studied main-
ly in lentic habitats) have multiple cohorts, including a
substantial number of larger individuals ranging in SL
from 20 to 40 mm (Britton et al, 1979; McMahon, 1983).
In our samples, four of the six C. fluminea populations
with complex age structure and relatively abundant large
individuals occurred with native unionacean clams. These
native unionids require long-term substratum stability to
establish populations due to their long lifespans and low
recolonization capability (Kraemer, 1979; McMahon,
1983; Miller et a/., 1986; Payne & Miller, 1989).
Thus, the greater abundance of large C. fluminea at
TAN, PAS, LEA, and BYP relative to most of the other
Mississippi populations is probably due to greater sta-
bility of the stream bottom at these four sites. Diamond
(1982) reported that the complexity of age structure in
lotic populations of the pluerocerid snail, ]uga plicefera,
in the Pacific Northwest was inversely related to the
likelihood of scour by flood flows. In addition, Sickel's
study of a single population of C. fluminea in the Al-
tahama River (1979) revealed differences in size demog-
raphy of samples taken on opposite sides of the river at
sites only 2,250 m apart, suggesting that variation in SL
PERCENT FREQUENCY
Figure 2. Shell length (SL) frequency histograms for six
Mississippi populations of Corbicula with a substantial number
of individuals greater than 20 mm SL and distinct multiple
cohorts.
distributions is important even within closely adjacent
assemblages in a lotic population. Sand and gravel shoals
of Mississippi streams and rivers are typically unconso-
lidated, and the distribution of unionids in the state is
limited to portions of streams with stable substratum
(Hartfield & Rummel, 1981; Hartfield & Ebert, 1986).
The effects of stream bottom stability on bivalve distri-
bution were especially evident during sampling at TAN
and BYP, where C. fluminea was most dense and union-
ids were restricted to gravelly sand (TAN) or sand (BYP)
shoals that were stabilized by trunks of fallen trees or
stands of the submersed macrophyte Vallisneria sp.
Spring and fall peaks in production of pediveliger
YOG NOX PRL BBL STR BUT CHI FMC
PencENr fwfoufnc.
Figure 3. Shell length (SL) frequency histograms for eight
Mississippi Corhiciila populations dominated b\ indi\ iduals less
than 20 mm SL that comprise an essentially single cohort of
1983 recruits. The slanted line through each histogram sepa-
rates 1983 recruits from older individuals.
B. S. Payne et a/., 1989
Page 81
larvae by C. fluminea have been observed in several
populations that have been repetitively sampled through-
out the year (reviewed by Britton et al., 1979; McMahon,
1983). In most lotic habitats in North America, spring
release of pediveligers during high flows would lead to
extensive downstream drift and little local recruitment.
Late summer and fall release of pediveligers during low
flow would be more likely to lead to local settlement near
the site of the adult population. Sickel (1979) observed
a major peak in density of drifting larvae in early May
in the Altahama River. However, major recruitment of
juvenile clams was not evident in his samples until mid-
July and these recruits attained a modal SL of only 4
mm by late October. These results suggest that the main
recruitment period occurred substantially later and dur-
ing lower flows than the late spring period of maximum
larval drift. Patterns of pediveliger release, transport, and
recruitment need to be studied in lotic habitats in relation
to both stream hydraulics and season. In addition, down-
stream transport and recruitment of juvenile and small
adult clams should be considered in studies of C. flu-
minea population dynamics in lotic systems.
The ubiquity of C. fluminea populations almost ex-
clusively comprised of small individuals (figure 3) during
the late spring of 1984 may reflect recent decimation of
those populations by flood flows during December, 1982
through mid-May, 1983 and December, 1983. Certainly,
the eight populations portrayed in figure 3 were com-
prised mainly of clams small enough to have been pas-
sively transported downstream from more stable up-
stream populations. Several investigations have shown
that larvae and young specimens (the latter with SL up
to 5.0 mm but generally less than 2.0 mm) of C. fluminea
are transported downstream in large numbers on water
currents (Sinclair, 1964; Goss & Cain, 1977; Sickel, 1979;
Williams & McMahon, 1986). McMahon and Williams
(1986) showed that populations decimated by thermal
stress in a lotic channel receiving heated effluent from
an electricity-generating plant in Texas were quickly
reestablished by downstream transport of both juveniles
(average SL = 3.5) and small adults (average SL = 9.0
mm SL). Large juveniles and small adults were evidently
rolled along the bottom by water currents, because no
individuals greater than 2.0 mm SL were recovered in
water column samples of drift (Williams & McMahon,
1986). Similarly, populations of C. fluminea killed during
winter by low water temperature in the New River,
Virginia rapidly reestablished the next spring by passive
transport of juveniles from a viable upstream population
(Cherry et al., 1980). Removal by downstream transport
of entire C. fluminea populations was noted by one of
us (RFM) after severe floods of the San Gabriel River in
Texas.
Ecological studies of intersite variation in life history
of North American C fluminea populations are espe-
cially valuable because interpopulation genetic variation
is low for this introduced species (Smith et al., 1979;
McLeod, 1986). Thus, comparative studies of C. flumi-
nea life history offer an opportunity to evaluate the in-
fluence of environmental factors on manifestation of life
history features in a relatively invariant genetic stock.
Lotic populations should be a major component of these
comparative studies, because C. fluminea is adapted to
such habitats (Britton & Morton, 1979; McMahon, 1983).
Our single but nearly simultaneous samples of 14 lotic
populations provide strong circumstantial evidence that
stream discharge and substratum stability are among the
most important environmental factors affecting C. flu-
minea population dynamics in lotic habitats. Compre-
hensive studies of the life history of this species in lotic
systems must focus on hydraulic ecology (see Statzner et
al., 1988) and include multiple stream locations as well
as frequent and long-term sampling programs.
ACKNOWLEDGEMENTS
Thanks are extended to Ms. Sharon Waites and Dr. Carl
Way for shell length measurements and review of a draft
of the manuscript, respectively. The data presented herein
were obtained from research conducted under the En-
vironmental Impact Research Program of the United
States Army Corps of Engineers by the Waterways Ex-
periment Station. Permission was granted by the Chief
of Engineers to publish this information.
LITERATURE CITED
Britton, J. C, D. R. Coldiron, L. P. Evans, Jr., C. Golightly, K.
D. O'Kane, and J. R. TenEyck. 1979. Reevaluation of
the growth pattern in Corbicula fluminea (Muller), In:
Britton, J. C. (ed). Proceedings, First International Cor-
bicula Symposium. Texas Christian University Research
Foundation. Fort Worth, TX, p. 177-192,
Britton, J. C. and B. Morton. 1979. Corbicula in North .\mer-
ica: the evidence reviewed and evaluated In: Britton, J.
C. (ed.). Proceedings, First International Corbicula Sym-
posium. Texas Christian University Research Foundation,
Fort Worth, TX, p. 249-287.
Cherry, D. S., J. H. Rodgers, Jr., R. L. Carney, and J. Cairns,
Jr. 1980. Dynamics and control of the Asiatic clam in
the New River, Virginia, N'irginia Polytechnic Institute
and State University, Bulletin No 123, Blacksburg, VA,
72 p.
Diamond, J. M. 1982. Stream geomorphology and benthic
habitat predictability as determinants of the population
dynamics and life history of the snail Juga plicifera. Jour-
nal of Freshwater Ecology 1:577-588.
Goss, L. B. and C. Cain, Jr 1977, Power plant condenser and
service water s\stem fouling by Corbicula, the Asiatic clam.
/(!: Jensen, L. D. (ed). Biofouling control procedures. Mar-
cel Dekker, Inc., New York, NY, p. 11-17.
Hartfield. P, D. and D. Ebert. 1986, The mussels of southwest
Mississippi streams. American Malacological Bulletin 4:
21-23,
Hartfield, P D. and R. G. Rummel. 1981. Mussels of the Big
Black River. Mississippi Academy of Science .\bstracts 26:
128.
Kraemer, L. R. 1979. Corbicula (Bivalvia: Sphaeriacea) vs.
indigenous mussels (Bivalvia: Unionacea) in U.S. rivers: a
hard case for interspecific competition? American Zoolo-
gist 19:1085-1096.
Page 82
THE NAUTILUS, Vol. 103, No. 2
McLeod. M. J. 1986 Electrophoretic variation in North
.American Corbicula. In. Proceedings, Second Internation-
al Corbicula Ssmposium. .•\merican .Vlalacological Bulle-
tin, Special Edition No. 2, p. 125-1.32.
McMahon, R. F. 1983. Ecology of an invasive pest bivalve,
Corbicula. In: Russell-Hunter, \V. D. (ed.). The Mollusca
(Volume 6): ecologv. Academic Press, Orlando, FL, p.
505-561.
McMahon, R. F. and (.' J Williams. 1986. Growth, life cycle,
upper thermal limit and downstream colonization rates in
a natural population of the freshwater bivalve mollusc.
Corbicula fluminea (Muller) receiving thermal effluents
In: Proceedings, Second Intenational Corbicula Sympo-
sium. American Malacological Bulletin, Special Edition
No. 2, p. 231-239.
Miller, A. C, B. S. Payne, and I) W Aldridge. 1986. Char-
acterization of a bivalve community in the Tangipahoa
River, Mississippi. The Nautilus 100 18-23.
Payne, B. S. and A. C. Miller. 1989 Growth and survival of
recent recruits to a population of Fusconaia ebena (Bi-
valvia: Unionidae) in the lower Ohio River. American
Midland Naturalist 121:99-104.
Sickel, J. B. 1979. Population dynamics of Corbicula in the
Altahama River, Georgia. In: Britton, J. C. (ed.). Proceed-
ings, First International Corbicula Symposium. Texas
Christian L'niversitv Research Foundation, Fort Worth.
TX, p. 69-80
Sinclair, R M 1964. Clam pests in Tennessee water supplies.
Journal of the American Water Works Association 56:592-
599.
Smith, M H . J Britton. P Burke, R K Chesser, M W Smith,
and J Hagen 1979 (ienetic \ariability in Corbicula. an
invading species. h\: Britton, J. C. (ed.). Proceedings, First
International Corbicula S\ mposium. Te.\as Christian Uni-
versity Research Foundation, Fort Worth, TX, p. 24.3-248.
Statzner, B., J. A. Gore, and V. H. Resh 1988. Hydraulic
stream ecology: observed patterns and potential applica-
tions. Journal of the North .American Benthological Society
7:.307-360.
Tate. C. H., E. J. Tharpe, G. A. Bednar, and W. T Oakley.
1982. Water reources data: Mississippi water year 1982.
US Geological Survey. \\ater-Data Report MS-82-1, Jack-
son, MS, 344 p.
Tate, C. H., E. J. Tharpe, G. A. Bednar, and W. T. Oakley.
1983. Water resources data: Mississippi water \ear 1983.
US Geological Surve\. U'ater-Data Report MS-83-1. Jack-
son, MS, 360 p
Tharpe, E. J., F. Morris III, and W T. Oakley. 1984. Water
resources data: Mississippi water year 1984. US Geological
Surve\, Water-Data Report MS-84-1, Jackson, MS, 340 p.
US Geological Survey. 1981 . Water resources data: Mississippi
water vear 1981. Water-Data Report MS-81-1, Jackson,
MS, 410 p.
Williams, C. J. and R F McMahon. 1986. Power station
entrainment of Corbicula fluminea (Muller) in relation to
population dynamics, reproductive cycle and biotic and
abiotic variable In: Proceedings, Second International
Corbicula S\mposium .American Malacological Bulletin,
Special Edition No. 2, p. 99-111.
THE NAUTILUS 103{2):83-84, 1989
Page 83
The Nomeiiclatural Status and Phylogenetic Affinities of
Syrinx arnanus Linne 1758 (Prosbranchia: Turbinellidae)
M. G. Harasewych
Richard E. Petit
Di'partinfiit of Imertebrate Zoology
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560 USA
The largest gastropod ever to have lived inhabits shallow
waters along the northern coast of Australia and adjacent
islands. Although this species, which reaches nearly a
meter in length (Anonymous, 1982), was known to pre-
Linnean authors (Buonanni, 1681: fig. 101; Rumphius,
1705: pi. 28, fig. A), its nomenclatural status and phy-
logenetic affinities have remained unclear during the
intervening centuries.
In erecting the ta.xon Murex aruanus, Linne (1758:
753) included references to two figures. The first, that
of Rumphius (1705: pi. 28, fig. A), represents a specimen
from the Aru Islands [between New Guinea and Austra-
lia] that Rumphius had called "Buccinum aruanum".
The second figure (Gualtieri, 1742: pl. 47, fig. B) depicts
a smaller, unrelated species endemic to the eastern coast
of the United States. Linne apparently never realized
that his taxon encompassed two species, as he repeated
the same figure references in his catalogue of the Mu-
seum Ulricae (Linne, 1764:641), which contained a spec-
imen of the western Atlantic species (Hollister, 1958:75-
78). In a later work (Linne, 1767:1222), he added a third
figure reference (Buonanni, 1681: fig. 101) that illustrates
the species from New Guinea and Australia. In all his
references to Murex aruanus, Linne listed the habitat as
New Guinea. There was no specimen of either species
in Linne's collection at the time the species was described
that could be regarded as the holotype (Hanley, 1855:
302).
The failure of Linne to differentiate between the two
disparate species included in his description has resulted
in the application of the taxon aruanus predominantly
to the Australian species [usually as Syrinx aruanus], and
less frequently to the western Atlantic species [as Fulgur
or Busycon aruanus]. The nomenclature was further
complicated when Hollister (1958:78) invalidly desig-
nated as lectotype of Murex aruanus Linne, 1758, the
specimen of the western Atlantic species from the Mu-
seum Ulricae collection that Linne examined and re-
ferred to in his 1764 work. This specimen was not avail-
able for lectotype designation as it was not referred to
by Linne in the original description (Linne, 1758:753)
and thus could not be considered part of the type series
as defined by ICZN article 72(b)(i). Nor was a lectotype
designation necessary, as the species had been effectively
restricted by previous revisers.
Born (1778:314-315), the first to refer to Murex aru-
anus Linne, cited the Buonanni and Rumphius, but not
the Gualtieri figures, and added a reference to figures
from Chemnitz [as "Martini"] (1780: table 138, figs. 1295
and 1296) [both are Neptunea antiquata (Linne, 1758)].
In 1780, Born repeated these references, but omitted
figure 1296. Schroter (1783:520) cited the Buonanni and
Rumphius figures, added a figure from Chemnitz (1780:
143, vignette 36, fig. D) that depicts the same species,
gave the correct locality, and specifically excluded the
Gualtieri figure from synonymy. It could be argued, and
was indeed accepted by some authors, that Born had
effectively restricted the specific name aruanus to the
species from new Guinea and Australia. However, by
specifically excluding the Gualtieri figure, Schroter un-
questionably restricted the species concept to include
only the Australian species. His action clearly meets the
first reviser provision of the International Code of Zoo-
logical nomenclature (ICZN article 24b).
As Roding's (1798:121) erection of the genus Syrinx
(type species: Murex aruanus Linne, 1758, by subsequent
designation of Winckworth, 1945:144) remained little
known until the twentieth century (Dall, 1915), this species
appears in earlier literature predominantly as "Fusus"
proboscidiferus Lamarck, 1816, or as Megalatractus pro-
boscidiferus (Lamarck, 1816). The family assignment of
Syrinx aruanus has generally followed that of the west-
ern Pacific species of Hemifusus, based on their super-
ficial conchological similarity. Thus, in most twentieth
century literature (e.g., Thiele, 1929:321; Wenz, 1943:
1217), this species is referred to the family Melongenidae.
Published observations on the anatomy of this species
have been few (Kesteven, 1904), yet the data presented
clearly indicate that Syrinx aruanus is referable to the
family Turbinellidae. The radula of S. aruanus (Keste-
ven, 1904: pl. 42, fig. 3) is nearly identical to that of
Turbinella pyrum (Dall, 1885: pl. 19, fig. 1; Harasewych,
Page 84
THE NAUTILUS. Vol. 103, No. 2
1987: fig. 21), but differs from the radulae of all melon-
genids in having monocuspid lateral teeth. Syrinx aru-
anus also has a long thin proboscis that folds into a non-
evaginable proboscis sheath (Kesteven, 1904:424, pi. 42,
fig. 2), a turbinellid character, and lacks the long snout
formed of fused tentacle bases that is diagnostic of melon-
genines. The protoconch, operculum and egg cases of
Syrinx are also most similar to those ot Turbinella an-
gulata. .\s noted by Kesteven (1904 448) the figures of
the alimentary system of "Fuaus proboscideus" pub-
lished by Haller (1888) do not accurately represent Syr-
inx aruanus. and ma\ represent an unrelated species.
In summar\, the ta.\on Murex aruanus Linne, 1758
has been clearly restricted to the northern Australian
species figured in Buonanni, Rumphius, and Chemnitz
by Schroter (1783) who may be regarded as the first
reviser. This species serves as the type of the genus Syrinx
Roding, 1798 [Si/rinx Bohadsch, 1761 having been de-
clared nonbinomial, ICZN Opinion 185]. Based on the
structure of its radula, proboscis, proboscis sheath, pro-
toconch, operculum and egg case. Syrinx is regarded as
a moiiot\ pic genus in the subfamily Turbinellinae of the
Turbinellidae.
LITERATURE CITED
.■\nonvmous. 1982. This may be the world's biggest. Hawaiian
Shell News :30(7): 12.
Born, I. 1778. Index reruni naturalium Musei Caesarei \in-
dobonensis. Pars I. Testacea. Kraus, N'ienna, [38], 458[82],
Ipi
Born, 1. von. 1780. Testacea Musei Caesarei \ indobonensis.
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T H Ef7N AUT I L U S
CONTENTS
Marine Biological Laboratory
LIBRARY
JAN 8 1990
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Volume 103, Number 3
December 28, 1989
ISSN 0028-1344
Ronald H. Karlson
John A. Sullivan
Effects of pagurid density and size on the behavior of
Crepidula convexa Say
85
Geerat J. Vermeij
Habitat and form of Crepidula grandis in Japan, with
comments on habitat specialization in calyptraeid
gastropods
89
Edward J. Petuch
New species of Malea (Gastropoda: Tonnidae) from the
Pleistocene of southern Florida
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Carl M. Way
Andrew C. Miller
Barrv S. Pavne
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the lower Tennessee River
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W. B. Saunders
Paul N. Bond
Lee C. Hastie
David Itano
On the distribution of Nautilus pompilius in the Samoas,
Fiji and Tonga
99
Faigel K. Vale
Michael A. Rex
Repaired shell damage in a complex of rissoid gastropods
from the upper continental slope south of New England
105
Bradford H. Burnham
Paul E. Fell
Distribution of Melampus bidentatus (Say and Succinea
wilsoni (Lea) within a tidal marsh in eastern Connecticut
109
Alan R. Kabat
The "Gray Catalogues" [Mollusca] of the British Museum
113
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THE NAUTILUS 103(3):85-88, 1989
Page 85
Effects of Pagurid Density and Size on the Behavior of
Crepidula convexa Say
Ronald H. Karlson
John A. Sullivan
Eculog) Program tSLHS)
Universit\ of Delaware
Newark, DE 19716, USA
ABSTRACT
The calyptraeid gastropod Crepidula convexa Sa\ is commonh
found on sfieiis occupied by hermit crabs. Although this fa-
cultative association is more common on small shells occupied
b\ Pagurus longicarpus Sa\ than on large shells occupied by
P. pollicaris Sa\ . field experiments clearU show that C. convexa
does colonize shells occupied b\ the latter hermit crab and that
densities of C. convexa can be maintained on such shells at low
pagurid densit\ (Shenk, 1986). In this contribution, we report
the results of a field experiment that illustrates the negati\'e
effect of a high densit> of P. pollicaris on C. convexa. In the
laboratory, we attempt to determine whether the decline in
numbers of C. convexa at a high density of P. pollicaris is a
behavioral response to the hermit crabs or the result of mor-
tality. We also examine the movement of C. convexa onto
alternative shell substrates and the effect of the size of P. pol-
licaris on these movements. Our results indicate that move-
ments of C. convexa are significantK affected b\ pagurid den-
sit) and size. We infer that this relativel)- motile calyptraeid
probably escapes mortality in the field by moving off of shells
occupied by large P. pollicaris and that shells occupied by P.
longicarpus provide a favorable substrate which may represent
a refuge from predation.
Key words: Escapes; refuges; predation; hosts; phoresis; epi-
zoism; Crepidula: Pagurus
INTRODUCTION
Along the east coast of the United States, the calyptraeid
gastropod Crepidula convexa occurs on a wide variety
of hard substrates. In addition to living on inanimate
objects, this organism can often be found living on shells
in facultative associations with other moilusks and two
species of hermit crabs (Hendler & Franz. 1971; Hoag-
land, 1977, 1978, 1979, 1984; Karlson & Cariolou, 1982;
Shenk, 1986; Shenk & Karlson, 1986; McGee, 1988). Cre-
pidula convexa is a relatively motile, nongregarious
species that lacks planktonic larvae (Hoagland, 1978).
Juveniles and adult males are not strongly attracted to
females (Hoagland, 1978), dispersion patterns are not
highly aggregated, and densities are relatively low com-
pared with more gregarious members of this genus
(Hoagland, 1978; Shenk, 1986; McGee, 1988).
In spite of not being highly substrate-specific nor
strongly attracted by conspecifics, juvenile and adult Cre-
pidula convexa can rapidly colonize small shells occupied
bv the hermit crabs Pagurus longicarpus (Karlson &
Cariolou, 1982) and P. pollicaris (Shenk, 1986; Shenk &
Karlson, 1986). Crepidula convexa can also colonize large
shells occupied by P. pollicaris. but densities on these
larger substrates have been found to be an order of mag-
nitude less than those on smaller shells (Shenk, 1986;
Shenk & Karlson, 1986). In a series of field experiments
designed to evaluate several alternative hypotheses ex-
plaining Crepidula distributions on shells occupied by
hermit crabs, Shenk (1986) noted that 1) a relatively high
density of C. convexa, resulting from colonization of
large shells occupied by P. pollicaris, could be main-
tained at low densities of P. pollicaris, 2) at high densities
of P. pollicaris, densities of C. convexa on these large
shells declined, 3) the density of C. convexa on small
shells occupied b\ P. longicarpus did not decline even
when placed together with high densities of P. pollicaris,
and 4) the differential response of co-occurring C. con-
vexa and its congeners C. plana Say and C. fornicata
(Linne) to high densities of P. pollicaris coincided with
observed distributional difl^erences; C. convexa was most
common on small shells, C. plana on the inside of large
shells, and C. fornicata on the outside of large shells (see
also Karlson & Shenk, 1983). All three species readily
colonized the outside of large shells, but only C. fornicata
could persist there with high densities of P. pollicaris.
After rejecting hypotheses invoking habitat selection
and interspecific competition as explanations for ob-
served distributional differences, Shenk (1986) inter-
preted his experimental results as support for a refuge
diversification hypothesis. Under this hypothesis, pre-
dation by P. pollicaris results in differential exploitation
of prey refuges by the congeners; they only coexist at
low predator densities. Since predation usually acts to
promote species coexistence by reducing prey densities
and the intensity of interspecific competition (Paine,
1966), the refuge diversification hypothesis suggests an
Page 86
THE NAUTILUS. \ol. 103. \o. 3
Table 1. Mean mimher n( Crepidula convexa per shell {± SD)
in cages with eitlier one or five Pagurus poUicaris. There was
one hermil crab per cage in 15 cages and five per cage in 5
cases
Nuniljer of hermit crabs per cage
Da\ of census
1
5
0
1.53 ± 0.9.3
1.20 ± 0.77
7-8
1.40 ± 1.07
1.00 ± 0.87
10-11
1.27 ± 0 89
072 ± 0.80
IV 14
1,07 ± 0 5.5
0.28 ± 0,53***
*••( = 3.716, p < 0 001
unusual role for predators in this assemblage (Shenk,
1985, 1986).
In this contribution, we focus on the interaction be-
tween Crepidula concexa and the two hermit crab species.
First, we illustrate the densit\ -dependent decline of C.
convexa from shells occupied b\ Pagurus poUicaris. We
then attempt to determine whether this decline is a be-
havioral response or the result of mortalitv of C convexa
at high densities of P. poUicaris. We also examine how
C. convexa responds to the presence of small shells oc-
cupied by P. longicarpus and how this response varies
with the size of P. poUicaris.
METHODS AND RESULTS
Our first experiment was conducted in the field to illus-
trate the decline in numbers ot Crepidula convexa in
response to a high densitv of Pagurus poUicaris (desig-
nated below as the densitv experiment). We conducted
a laboratory experiment in which alternative shell sub-
strates were provided in aquaria with C. cotivexa and a
high density of P. poUicaris (designated belov\ as the
alternative substrate experiment). In another laboratory
experiment, we evaluated the effect of the size of P.
poUicaris on the beliavior of C. convexa (designated be-
low as the size experiment).
Density experiment: During JuK 1987, we placed 20
wire cages (80 x 80 x .30 cm) witli 0.6 cm galvanized
mesh approximately 50 m from shore at a depth of 1.0
m below mean low water at Cape Henlopen, Delaware.
We used the small mesh to inhibit colonization of the
shells by nonexperimental animals. During \o\v tides over
a .D-day period, we used a dipnet to collect hundreds of
shells occupied by Pagurus poUicaris. Most commonly
encountered were the shells of the gastropods Busycon
carica (Gmelin), Polinices duplicatus (Sa\), and \atica
clausa Rroderip and Sowerbv. From these collections,
we used 40 hermit crabs in shells with 1-5 Crepidula
convexa each. We placed a single hermit crab into each
of 15, randomly designated cages and five hermit crabs
each into the remaining 5 cages; these densities are equiv-
alent to 1.6 and 7.8 hermit crabs/m-. We cleaned the
cages and inspected all shells for C. convexa after the
experiment had been in place for 7-8, 10-11, and 13-
14 da vs.
.■\fter 13-14 days, there was a significant, densitv-de-
pendent decline in the number of Crepidula convexa
per shell (table 1). In response to higher densities of
Pagarus poUicaris, 77"^ of the original 30 C. convexa
disappeared while only 301 of the original 23 disap-
peared from the shells at low pagurid density. The den-
sit\ -dependent decline in numbers of C. convexa is com-
parable to those reported b\ Shenk (1986) for predation
experiments conducted in Long Island Sound during 1984
and 1985.
.41ternative substrate experiment: In this experiment,
we placed three Pagurus poUicaris into each of 10 aquar-
ium compartments (30 x 20 cm) to achieve a high den-
sity equivalent to 50 hermit crabs/m-. Such densities
occur naturalK especialK at sites where hermit crabs
acquire new shells from dying gastropods (RHK, personal
observation). Each hermit crab-occupied shell had 1-4
Crepidula convexa. We also placed six Ilyanassa obsoleta
(Say) shells with no C. convexa into each compartment;
five raiidomK' selected compartments received Ilyanassa
shells occupied b\ P. longicarpus and five received shells
which had been plugged with non-toxic silicon sealant
to prevent occupancv- b\ hermit crabs. These alternative
shell substrates were examined for C. convexa everv 1-
2 days for 8 days.
.At the end of this experiment, 36?c of the original 39
Crepidula convexa left their large shell substrates after
8 davs (table 2). Comparisons betv\ een aquaria receiving
plugged vs. hermit crab-occupied Ilyanassa shells indi-
cate that there was no significant difference in the mean
number of C. convexa per shell remaining on large shells
occupied by Pagurus poUicaris (t = 0.841, p > 0.40,
table 2), but the small Ilyanassa shells occupied b\' P.
longicarpus accumulated significantK more C convexa
than did plugged shells (t = 2.594, p < 0.02, table 2).
Of the fourteen individuals that left the large shells over
the 8-da\ period, 10 accumulated on small, hermit crab-
occupied shells, one on a small plugged shell, and three
were unaccounted for.
Size experiment: .-Kfter collecting 30 Pagurus poUicaris
from the field, we divided them into large (18.5 ± 4.3
[SD] mm) and small (9.3 ±2.1 [SD] mm) size classes
based on anterior shield lengths. Maximum anterior shield
lengths are reported to be 21.3 mm for P. poUicaris and
7.25 mm for P. longicarpus (Williams, 1984). We then
placed three P. poUicaris from a single size class into
each of five, randomly selected aquarium compartments
for each size class. Each of these hermit crab-occupied
shells had one C. convexa at the start of the experiment.
We also placed six Ilyanassa shells occupied by P. lon-
gicarpus (again with no C. convexa) into each com-
partment to provide C. convexa with alternative sub-
strates. In order to monitor net movement of C. convexa
among substrates, we examined all shells at 1-2 day
intervals lor 10 davs.
Over this 10 day period as many as 12 Crepidula
convexa had moved onto shells occupied by Pagurus
longicarpus and, bv the end of the experiment, 709c of
R. H. Karlson and J. A. Sullivan, 1989
Page 87
Table 2. Mean number of Crepidula convexa per shell { ± SD) in 10 laborator>- aquarium compartments containing three Pagurus
pollicam each and either si.\ plugged (PL) Ilyanassa obsoleta shells (I) or six shells occupied 1)\ P iongicarpus (HC:) P, pollicaris
were in Busycon carica, Polinices dupHcatus, or Natica clausa shells (B, P, or N).
Treatment
and shell
t>pe
HC
PL
Day of census
B, P, or N
I
B. P. or N
1
0
1.40 ± 0.84
0
1.20 ± 0.41
0
1
1.07 ± 0.80
0.20 ± 0.48
1.13 ± 0.35
0
2
1.13 ± 0.91
0.23 ± 0.51
1.00 ± 0.53
0
3
1.10 ± 0.85
0.20 ± 0.48
0.87 ± 0.64
0
4
0.93 ± 0.71
0.20 ± 0.48
0.93 ± 0.60
0.03
± 0.19
5
0.93 ± 0.71
0.23 ± 0.51
0.73 ± 0.60
0.03
± 0.19
7
0.87 ± 0.64
0.30 ± 0.54
0.67 ± 0.62
0.03
± 0.19
,S
0 93 ±0 71
0.33 ± 0,62
0,73 ± 0,60
0.03
± 0.19
Table 3. Mean number of Crepidula convexa per shell (± SD) in the size experiment. Each aquarium compartment contained
three Pagurus pollicaris in either Busycon carica. Polinices duplicatus. or yatica clausa shells (B, P, or N) and six Pagurus
longicarpus in Ilyanassa ohsolcia shells (I),
nsus
P, pollicaris
size
class and shell type
Small
Large
Day of ce
B. P. or N
I
B. P. or N
I
0
1
2
3
5
6
7
8
9
10
1
0.67 ± 0.62
0.60 ± 0.64
0.27 ± 0.60
0.40 ± 0.64
0.47 ± 0.75
0.47 ± 0.64
0.58 ± 0.79
0.58 ± 0.79
0,58 ± 0,79
0
0.13 ±
0.13 ±
0.33 ±
0.23 ±
0.20 ±
0.20 ±
0.21 ±
0.21 ±
0.21 ±
0.35
0.35
0.48
0.43
0.40
0.40
0.40
0.40
0.40
1
0.60 ± 0.51
0.60 ± 0.51
0.47 ± 0.51
0.47 ± 0.51
0.47 ± 0.64
0.47 ± 0.51
0.25 ± 0.45
0.17 ± 0.38
0,17 ± 0.38
0
0.07 ± 0.24
0.07 ± 0.24
0.07 ± 0.24
0.07 ± 0.24
0.03 ± 0.19
0
0,12 ± 0.33
0.08 ± 0.28
0,08 ± 0,28
the original 30 C.
convexa had left shells
i occupied b> P.
DISCUSSION
pollicaris (table 3). However, comparisons of the mean
number of C. convexa per shell indicate that there were
no significant differences between treatments for shells
occupied by either P. pollicaris (t = 1.620, p > O.IO) or
P. longicarpus (t = 1.304, p > 0.10) after 10 days. Never-
theless, there are some interesting trends in the data.
Over the course of the experiment, there was a steady
decline in numbers of C. convexa on shells occupied by
large P. pollicaris (from 15 to 2), while se\eral C. convexa
moved back and forth between shells occupied by small
P. pollicaris (from 15 to 4 to 7) and those occupied by
P. longicarpus (from 0 to 10 to 5). Such active movement
by C. corivexa among t\ pes of substrates has been noted
previously by Hendler & Franz (1971) and Karlson &
Cariolou (1982). By the end of the experiment, 16 C.
convexa were still on shells, 8 had moved to the walls of
the aquaria (7 of these were part of the large P. pollicaris
group), and 6 had died. These deaths occurred in two
aquarium compartments (one from each of the two
groups) in which hermit crabs had died and fouled the
water.
The above field and laboratory experiments corroborate
earlier reports by Shenk (1986) that the number of C.
convexa on shells occupied by P. pollicaris declines in
response to high densities of this hermit crab. In our
laboratory experiments, we observed this decline as a
total of 38 individuals (out of a possible maximum of 69)
moved off their original shells and as many as 23 moved
onto shells occupied by P. longicarpus (tables 2 and 3).
The absence of man\- deaths in these experiments indi-
cates that the density-dependent decline in numbers of
C. convexa observed in the field (Shenk, 1986; table 1)
was probably not the result of mortality due to predation.
Instead, our results are consistent with the h>pothesis
that the decline in numbers of C. convexa is a density-
dependent behavior such as an escape response from P.
pollicaris.
An alternative explanation for why Crepidula convexa
tend to move off shells occupied by Pagurus pollicaris
at high, but not low densities of this hermit crab might
involve sex ratio biases introduced into the design of these
Page 88
THE NAUTILUS, Vol. 103, No. 3
experiments. Since females tend to be less motile during
the summer when the\ brood (Hoagland, 1978), it is
possible that disproportionately more females were used
in treatments \\ ith low densities of P. pollicaris b\ Shenk
in 1984 and 1985 (Shenk, 1986) and b\ ourseKes in the
density experiment (table 1). Sex determinations were
not made in any of these experiments, so we cannot reject
this possibility. However, we consider a sex ratio bias to
be an unlikely explanation, because densit\- manipula-
tions have yielded consistent field results on three sep-
arate occasions.
The movement of Crepidula convexa onto small shells
occupied by Pagurus longicarpua has now been docu-
mented in field experiments (Karlson 6i Cariolou, 1982;
Shenk, 1986) and in the laboratory (Karlson & Cariolou,
1982; tables 2 and 3). However, it continues to be unclear
why they exhibit this noiiraiidom beha\ior. In the field
experiment conducted b\ Karlson and Cariolou (1982)
in a shallow subtidal area at Cape Henlopen, C. convexa
rapidly colonized shells which had been plugged as well
as shells occupied by P. tongicarpus. In the presence of
high densities of P. pollicaris in the alternative substrate
experiment reported above, C convexa appeared to pre-
fer shells occupied by P. longicarpus over plugged shells.
Alternative explanations for this phenomenon include
the notions that 1 ) these small shells represent refuges
from predation (Shenk, 1986), 2) the\ represent a limited
spatial resource in an environment characterized b\ high
sedimentation rates (Karlson & Cariolou, 1982), and 3)
these shells are preferred substrates whose selection by
C convexa in\ol\ es cues and sensory adaptations specific
to this association. Given the wide variety of substrates
utilized by this species, the facultative nature of the
associations with hermit crabs, and the field experiment
demonstrating no preference for shells occupied b\' P.
longicarpus (Karlson & Cariolou, 1982), we consider this
last option unlikely. However, additional studies are
clearK needed to discriminate among these alternatives.
Our data indicate that the sliell-to-shell movement of
Crepidula convexa is affected by the size of Pagurus
pollicaris; larger hermit crabs appeared to inhibit the
movement of C. convexa back onto hermit crab-occupied
shells and to favor their movement up the sides of the
aquaria. This size-dependent eftect on mobilit\ ma\ ex-
plain the large effect of shell size on colonization by C.
convexa (Shenk, 1986; Shenk & Karlson, 1986); in field
experiments, they colonized small shells (< 100 cm-)
occupied by P. pollicaris at much higher rates than larger
shells occupied by larger P. pollicaris.
Crepidula convexa is a member of the family Calyp-
traeidae, a group containing species that are commonly
found on the shells of other mollusks. Although the exact
nature of most of these associations remains subject to
speculation, an escape response and refuge exploitation
have been reported for interactions involving Crepidula
adunca (Sowerb\), the host snail Calliostoma ligatum
(Gould), and the predator Leptasterias hcxactis (Brandt)
(X'ermeij et al., 1987). These associations should continue
to provide useful models for investigation of the ecolog-
ical and evolutionary effects of predators on the behavior
of their pre\ and the exploitation of host species as ref-
uges.
ACKNOWLEDGEMENTS
We thank S. Karlson, D. Levitan, A. Shenk, and two
anonv mous reviewers for comments on this manuscript
and the College of Marine Studies, Uni\ersit\ of Dela-
ware, for laboratorv space. The Peter White Fellowship
program at the University of Delaware provided finan-
cial support. This is Ecology Program contribution no.
127.
LITERATURE CITED
Hendler, G. and D. R. Franz. 1971. Population dvnamics and
life history of Crepidula convexa Sa\ (Gastropoda; Pro-
sobranchia) in Delaware Ba\ The Biological Bulletin 141:
514-526.
Hoagland, K. E. 1977. Systematic review of fossil and recent
Crepidula and discussion of the Calvptraeidae. Malaco-
logia 16;.353-420.
Hoagland, K, E. 197S, Protandrv and the evolution of en-
vironmentalU -mediated sex chance: a stud\ ot the Mol-
lusca. Malacologia 17:365-391
Hoagland, K E. 1979. The behavior of three sympatric species
of Crepidula (Gastropoda: Prosobranchia) from the At-
lantic, with implications for evolutionary ecology. The
Nautilus 94:143-149.
Hoagland. K. E. 1984. Use of molecular genetics to distinguish
species of the gastropod genus Crepidula iProsobranchia:
Calvptraeidae). Malacologia 25:607-628.
Karlson, R. H. and M. A Cariolou. 1982. Hermit crab shell
colonization In Crepidula convexa Sav Journal of Exper-
imental Marine Biologv & Ecology 65:1-10.
Karlson, R. H. and M. A. Shenk. 1983. Epifaunal abundance,
association, and overgrowth patterns on large hermit crab
shells. Journal of Experimental Marine Biologv & Ecologv
70:55-64.
-McGee, B. L. 1988. Gregariousness in Crepidula: fact or
fiction.'' Masters thesis, University of Delaware, Newark,
Delaware. 83 p.
Paine, R. T. 1966. Food web complexitv and species diversity.
The American Naturalist 100:65-75.
Shenk, M. A. 1985. Effects of competition and predation on
the distribution of Crepidula species .American Zoologist
25:87A (Abstract 457).
Shenk, M. A. 1986. Ecological determinants of Crepidula
distributions. Doctoral dissertation, Universitv of Dela-
ware, Newark. Delaware, 91 p.
Shenk, M. A. and R, H. Karlson. 1986. Colonization of a shell
resource by calvptraeid gastropods: tests of habitat selec-
tion and preemption models. Journal of Experimental Ma-
rine Biology &i Ecologv 99:79-89.
Vermeij, C; J., R. B. Lowell, L. J. Walters, and J. A. Marks.
1987 Good hosts and their guests: Relations between tro-
chid gastropods and the epizoic limpet Crepidula adunca.
The Nautilus 101:69-74.
\\ illiams. .\. B. 1984. Shrimps, lobsters, and crabs of the
.Atlantic coast of the eastern L nited States. Maine to Flor-
ida, Smithsonian Institution Press. Washington, D.C.,
550 p.
THE NAUTILUS 103(3):89-91, 1989
Page 89
Habitat and Form of Crepidula grandis in Japan, with
Comments on Habitat Specialization in Calyptraeid Gastropods
Geerat J. Vermeij
Department of Geology
University of California at Davis
Davis, CA 95616
ABSTRACT
Crepidula grandis (Middendorff, 1949) is the only member of
the Cal\ ptraeidae in the vicinity of Akkeshi, Hokkaido, Japan.
It occupies the full range of substratum types that on many
other coastlines are divided among three or more species of
Calyptraeidae. Crepidula grandis li\ing inside hermit-crab shells
are relativeK flatter than are indi\ iduals on shell exteriors, but
they are less flattened than are species such as C. perforaiu
(Valenciennes, 1846) or C. ivalshi (Reeve, 1859), which are
more or less restricted to the interiors of hermit-crab shells.
Great unexplained differences among regions exist in the man-
ner in which calyptraeids have become morphologically spe-
cialized to the occupation of gastropod shell substrata.
Key words: Crepidula, Calyptraeidae, substratum specializa-
tion, hermit crabs, biogeography .
Crepidula grandis (Middendorff, 1849) is an abundant
shallow-water calyptraeid gastropod in the cool-temper-
ate waters of the northwestern Pacific. It is the single
living member of a distinctive North Pacific group of
large-bodied species that during the Miocene, Pliocene,
and early Pleistocene occurred in both the western and
eastern Pacific Ocean (Hoagland, 1977a; Vermeij, 1989).
Despite the fact that C. grandis is both biogeographicalK'
and ecologically interesting, almost nothing is known of
its biolog). Habe (1958) pointed out that the species in
the vicinity of Akkeshi, Hokkaido (Japan) occurs on the
lower valves of the commercial scallop Mizuhopecten
yessoensis (Jay, 1856), which like C. grandis belongs to
a lineage that underwent a range restriction to the north-
western Pacific (Masuda, 1971; Kafanov, 1986; Vermeij,
1989). Soviet workers (Golikov & Gulbin, 1978; Golikov
& Kussakin, 1978) have summarized the depth distri-
bution of C. grandis, which extends from the low inter-
tidal zone to a depth of more than one hundred meters.
During the summer of 1988, I had the opportunitv' to
make some observations on C. grandis in the vicinity of
the Akkeshi Marine Biological Station on the southeastern
coast of Hokkaido. Together with some comments on
patterns of substratum specialization among other calyp-
traeids, these observations are presented in this brief
account.
Among cool-temperate northern marine regions with
native species of Crepidula, the northwestern Pacific is
unique in lacking small flat-shelled species that are more
or less specialized for life on the inner surfaces of gas-
tropod shells inhabited by hermit crabs. Three species
of Crepidula have been recorded from the northwestern
Pacific: C. grandis (which also occurs along the northern
and western coasts of Alaska); C. lingidata (Gould, 1846),
a species also found in the northeastern Pacific; and the
endemic C. derjugini Golikov and Kussakin, 1962. Noth-
ing is known about the habitats of C. lingulata in the
northwestern Pacific, but in Puget Sound (Washington)
in the northeastern Pacific I have found it on stones and
on the inner surfaces of empty pelecypod valves. Golikov
and Kussakin (1978) report that C. derjugini lives on
stones and especially on the outer surfaces of shells of
the gastropod Neptunea.
Crepidula grandis is the only species of Crepidula
known from the vicinity of Akkeshi (see also Habe, 1958).
In Akkeshi Bay, the species occurs sporadically in the
low intertidal zone and is abundant in the shallow sub-
littoral. It occupies a wide variety of hard substrata,
including open rock surfaces, the abandoned cavities of
pholad pelecypods, the lower valves of Mizuhopecten
yessoensis, the outer surfaces of opercula of Fusitriton
oregonensis (Redfield, 1846), the outer surfaces of shells
of living gastropods and hermit crabs, and the inner
surfaces of hermit-crab shells. The pattern of occurrence
of C. grandis on outer and inner surfaces of shells at
Akkeshi is shown in Table 1.
Species of Crepidula that are usually found inside large
shells of hermit crabs are usually concave when seen
from above, closely conforming with the curvature of
the host shell. Examples include C. perforans (Valen-
ciennes, 1846) in the cool-temperate and warm-temper-
ate northeastern Pacific, C. nivea (C. B. Adams, 1852)
in the tropical eastern Pacific, C. plana (Say, 1822) in
the western Atlantic, and C. (Siphopatella) walshi (Reeve,
1859) in southeast Asia and northern Australia. When
settling on shells, C. plana apparently alwavs begins life
on the shell exterior and then migrates or becomes re-
stricted to the inside (Shenk & Karlson, 1986). The species
also occurs under stones and on the horseshoe crab Lim-
ulus polyphemus (E. C. Dudley, personal communica-
tion).
Although individuals of C. grandis taken from the
Page 90
THE NAUTILUS. Vol. 103, No. 3
Table 1. Occurrence of Crepidula grandis on exterior and
interior surfaces of gastropod shells at .Akkeshi, Hokkaido.
Host shell
Per-
N(, Ng centage
Euspira pila (Pilsbury, 1911) hermit
crabs, e.xterior 9 2 22%
E. pila hermit crabs, interior 9 0 0
Buccinum miranduni Smith, 1895, living 67 6 9%
B mirandum hermit crabs, e.xterior 22 5 23%
B. mirandum hermit crabs, interior 22 1 5%
Septunea arlhritica Bernardi, 1S5S, living 9 0 0
N. arlhritica hermit crabs, exterior 20 2 10%
N. arlhritica hermit crabs, interior 20 .5 2.5%
Ni,: .Number of host shells.
Ng! Number of C. grandis.
Percentage: number of C. grandis divided by number of host
shells multiplied by one hundred.
inner surfaces of hermit-crab shells at Akkeshi are rel-
atively flatter (mean height : length ratio 0.30 ± 0.025,
n = 18) than are individuals living on gastropod shell
exteriors (mean height : length ratio 0.38 ± 0.047, n =
20, p < 0.001 b\ T-test), the former are still convex
when seen from above and are therefore much less flat-
tened than are shells of the specialized hermit-crab as-
sociated species. In fact, shells of C. grandis from the
inner surfaces of hermit-crab shells are indistinguishable
in profile from shells collected from the lower valves of
Mizuhopecten yessoensis (mean height : length ratio 0.30
± 0.022, n = 8).
These observations show that C. grandis at Akkeshi is
a classic habitat generalist, occupying the full range of
substrata that on many coastlines are divided among at
least three species of Crepidula, some of which are spe-
cialized with respect to the type of substratum occupied.
Interesting regional differences exist in the extent of this
specialization. In the vicinity of Puget Sound, C. adunca
(Sowerby, 1825) is found almost exclusively on exterior
surfaces of the shells of living gastropods (Vermeij et ai,
1987), whereas C. perforans occurs only on shell inte-
riors; C. lingulata and C. nunimaria (Gould, 1846) and
the related Calyptraea fastigiata (Gould, 1846) are found
on stones as well as on empty valves of pelecypods. In
the northwestern Atlantic, C. plana is usually found on
shell interiors; whereas C. convcxa (Say, 1822) is found
on shell exteriors, stones, and eelgrass blades, as well as
on the larger stones that are the typical habitat of C.
furnicata (Linnaeus, 1758) (Franz & Hendler, 1970;
Hendler & F"raiiz, 1971; Hoagland, 1977a, b; Shenk &
Karlson, 1986). On the Pacific coast of Panama, my ob-
servations indicate that C. incurva (Broderip, 1834) is a
specialist on exterior surfaces of gastropod and hermit-
crab shells, C. nivea is an interior-surface specialist, and
Calyptraea mamrnillaris (Broderip, 1834) is found on
interior (rarely on exterior) surfaces of empty pelecypod
valves. The larger calyptraeids, such as Crepidula strio-
lata (Menke, 1851), C. lessonii (Broderip, 1834), C. onyx
(Sowerby, 1824), C. aculeata (Gmelin, 1791), Crucibu-
lum umbrella (Deshaves. 1830), C. spinosum (Sowerby,
1824), C. personatuni Keen, 1958, and C. scetellatum
(Wood, 1828), occur mainly on stones and solid rocks
and only occasionally on shells. A division of substrata
similar to that in Panama has been qualitatively de-
scribed by Bandel and W'edler (1987) for the caKptraeids
of the Caribbean coast of Colombia. On the west side of
the York Peninsula of northern Queensland (Australia),
I have found Crepidula walshi on the interior surfaces
of hermit-crab shells; it was the only calyptraeid species
I was able to find localK .
.\lthough there is remarkabh little published infor-
mation about the ecologv- of cal\ ptraeids except for some
of the .American species, available information indicates
that southeastern Hokkaido is not the onl\- region where
there is no substratum specialization by calyptraeids to
shells. Several calyptraeids coexist on the temperate west
coast of South .America, but none of the species appears
to be found only on either the exterior or interior surfaces
of shells (see e.g.. Marinco\ich, 1973). None of the species
is flattened in a way suggesting specialization for life on
shell interiors. Only one calypytraeid, Cheilea equestris
(Linnaeus 1758), occurs on the islands of the tropical
western Pacific. Like species of Cheilea in tropical .Amer-
ica, C. equestris is found under rocks and never occurs
on shells. The hipponicid Sabia conica (Schumacher, 1817)
is found only on gastropod and hermit-crab shell exteriors
in the tropical Indo-Pacific and in temperate .Australia,
but no oceanic western Pacific or Indian Ocean gastropod
is specialized for life on shell interiors. This situation
applies to the shallow waters of the Caribbean islands as
well.
These patterns of specialization are puzzling. .Although
calyptraeids that are specialized for life on shell interiors
appear to be restricted to productive waters on the shores
of continents and large islands, not all producti\e shores
harbor such species. Hermit-crab specialists occur either
sympatrically with many other calyptraeids (as in the
eastern Pacific and western .Atlantic) or alone (C. walshi
in Queensland). The absence of hermit-crab associates
cannot be attributed to the lack of suitable quantities of
large gastropod shells, for the latter are abundant in the
tropical western Pacific and West Indies, as well as in
northern Japan where C. grandis facultatively occupies
shell interiors. In the case of C. graridis. facultative oc-
cupation of shell interiors is accompanied b\ relati\ e shell
flattening in a species with considerable morphological
plasticity, but this specialization is much less extreme
than in interior specialists, with the result that C. grandis
is likely to be more intrusive to its hermit-crab hosts.
Northwestern Europe lacked species of Crepidula un-
til C fornicata was introduced with oysters from eastern
North America (see Hoagland, 1985). It will be inter-
esting to determine \\ liether C. fornicata in Europe oc-
cupies the interior surfaces of hermit-crab shells, a hab-
itat rarely if ever occupied by C. fornicata in its native
.American range.
G. J. Vermeij, 1989
Page 91
ACKNOWLEDGMENTS
The University of California at Davis and the University
of Maryland at College Park partially funded my re-
search in Japan. I thank Janice Cooper for technical
assistance, Elizabeth Dudle> and Edith Zipser for field
assistance, the director and staff of the Akkeshi Marine
Biological Station for greatly facilitating our visit to Ja-
pan, and K. Elaine Hoagland for critically reviewing the
manuscript.
LITERATURE CITED
Bandel, K. and E. VVedler. 1987 Hydroid, amphineuran and
gastropod zonation in the littoral of the Caribbean Sea,
Columbia. Senckenbergiana Maritima 19:1-129.
Franz, D. R. and G. Handler. 1970. Substrate diversity and
the taxonomy of Crepidula convexa (Say) (Gastropoda:
Prosobranchia). University of Connecticut Occasional Pa-
pers (Biological Sciences Series) 1:281-289.
Golikov, .\. \. and V. V. Gulbin. 1978. Prosobranchia! gas-
tropods of the Kurile Islands. I. Orders Docoglossa-Ento-
mostoma. In: Kussakin, O. G. (ed.l. Fauna and vegetation
of the shelf of the Kurile Islands. Academy of Sciences,
USSR, Far East Science Center, Institute of Marine Biol-
ogy: 159-223.
Golikov, A. N. and O. G. Kussakin. 1978. Rakovinnye briu-
khonogie molliuski litorali morei SSSR. "Nauka", Lenin-
grad, 256 p.
Habe, T. 1958. Fauna of Akkeshi Bay XXV. Gastropoda.
Publications of the .Akkeshi Marine Biological Station 8:
1-39.
Hendler, G. and D. R. Franz. 1971. Population dynamics and
life history of Crepidula convexa Say (Gastropoda: Pros-
obranchia) in Delaware Bay. Biological Bulletin 141:514-
526.
Hoagland, K. E. 1977a. Systematic review of fossil and Recent
Crepidula and discussion of evolution of the Calyptraei-
dae. Malacologia 16:353-420.
Hoagland, K. E. 1977b. A gastropod color polymorphism: one
adaptive strategy of phenotypic variation. Biological Bul-
letin 152:360-372.
Hoagland, K. E. 1985. Genetic relationship between one Brit-
ish and several North American populations of Crepidula
jornicata based on allozyme studies (Gastropoda: Calyp-
traeidae). journal of Molluscan Studies 51:177-182.
Kafanov, A. I. 1986. Comparison of the geographical and
stratigraphical ranges of Fortipectininae and Patinopec-
tininae (Bivalvia: Pectinidae). Monographs of the Mizu-
nami Fossil Museum 6:23-40.
Marincovich, L., Jr. 1973. Intertidal mollusks of Iquique,
Chile. Natural History Museum of Los Angeles County
Science Bulletin 16:1-49.
Masuda, K. 1971. On some Patinopecten from North Amer-
ica. Transactions and Proceedings of the Palaeontological
Society of Japan (n.s.) 83:166-178.
Shenk, M. A. and R. H. Karlson. 1986. Colonization of a shell
resource b\ calyptraeid gastropods: tests of habitat selec-
tion and preemption models. Journal of Experimental Ma-
rine Biology and Ecology 99:79-89.
\'ermeij, G. J. 1989. Geographical restriction as a guide to
the causes of extinction: the case of the cold northern
oceans during the Neogene. Paleobiology 15. In press.
\ermeij, G. J., R. B. Lowell, L. J. Walters, and J. A. Marks.
1987. Good hosts and their guests: relations between tro-
chid gastropods and the epizoic limpet Crepidula adunca.
Nautilus 101:69-74.
THE NAUTILUS 103(3):92-95, 1989
Page 92
New Species of Malea (Gastropoda Tonnidae) from the
Pleistocene of Southern Florida
Edward J. Peluch
Department of Geology
Florida Atlantic University
Boca Raton, FL 33431, USA
ABSTRACT
Two new large tonnids of the genus Malea \'alenciennes, 1832
are described from the early and middle Pleistocene of the
Everglades Basin of southern Florida. These are Malea springi
new species from the Caloosahatchee Formation (Calabrian
Pleistocene) along the Miami Canal, Palm Beach Count) , and
Malea petiti new species from the Bermont Formation (Af-
tonian Pleistocene) in extreme southwestern Palm Beach Coun-
t\. With a length of 189 mm (holot\pe), Malea springi is the
largest of the known fossil western Atlantic Malea species. Due
to its stratigraphic position in the middle Pleistocene Bermont
Formation, Malea petiti may have been the last living North
Atlantic Malea species.
INTRODUCTION
In the Neogene formations of Florida, the tonnid genus
Malea Valenciennes, 1832 is rarely seen, with only a few
complete specimens ever having been collected. The ge-
nus was unrecorded from the Floridian Peninsula until
1964, when Olsson and Petit documented the first records
of Malea species in the Pliocene and early Pleistocene
of the Everglades region. The best preserved specimens
were collected in the early Pliocene (Zanclian Stage)
"Pinecrest Beds" (= Buckingham Formation of Mans-
field, 1939; see Petuch, 1986, 1988, for stratigraphic no-
menclature and dating) from near Lake Okeechobee,
and were assigned to the Pliocene \'enezuelan species
Malea densecostata (Rutsch, 1934) (figure 4). The pres-
ence of this characteristic South and Central American
index fossil in Florida was used by Olsson and Petit (1964;
554) to correlate the "Pinecrest Beds" with the Punta
Gavilan Formation of Venezuela.
Within the same paper (1964:553), Olsson and Petit
also mention in passing the occurrence of a second, larger
Malea species that had been found in dredgings from
the early Pleistocene ((Calabrian Stage) (see Petuch, 1988
for correlation and dating) Caloosahatchee Formation
along the Miami Canal in the central Everglades Basin.
This Pleistocene Floridian Malea, however, was only
collected as tantalizing fragments, and could not be as-
signed to any known taxon. Olsson and Petit (1964;553)
also mention the presence of poorly -preserved internal
molds of a possible third Floridian Malea species. These
were said to be exposed in the limestone of the late
Pliocene (Piacenzian Stage) Tamiami Formation, pre-
sumably at the stratotype area along the Tamiami Trail
in Collier County. In the subsequent literature on Flo-
ridian molluscan paleontology , however, little attention
has been given to these large and stratigraphically im-
portant gastropods.
In 1981, the Miami Canal, particularly in the area just
north of the levee at the Broward-Palm Beach County
line, was deepened to allow better water Dow during
times of drought. At that time, several complete speci-
mens of a large new Malea species were dredged during
the canal deepening, from approximately 20 meters depth
below the Everglades surface. This large, un-named ton-
nid was collected in an undescribed coral reef facies of
the Caloosahatchee Formation, along with classic Ca-
loosahatchee index fossils such as Siphocypraea prohlem-
atica Heilprin. 1886, Hystrivasum horridum (Heilprin,
1886), and Tiirhinella scolymoides Dall, 1890. Judging
from its occurrence in the Caloosahatchee Formation
along the Miami Canal, the un-named Malea appears to
represent the same species that Olsson and Petit had
collected as fragmentary specimens in the 1960's. This
new reef-associated Caloosahatchee species is the largest
Malea known from North America and is one of the
largest gastropods found in the Caloosahatchee Forma-
tion.
Road fill quarries in the central Everglades region,
along the Palm Beach- Broward County line, have re-
cently yielded yet another large, un-named Malea, in
this case from the early middle Pleistocene (Aftonian
Stage) Bermont Formation. Unlike the Miami Canal Ca-
loosahatchee specimens, however, the Bermont speci-
mens were all encased in a semifriable limestone and
were only partially complete. U'ithin this imdescribed
indurated member of the Bermont Formation, Malea
specimens were collected along with classic Bermont in-
dex fossils such as Fasciolaria oheechobeensis Tucker and
Wilson, 1932, Melongena (Rexmela) bispinosa (Philip-
pi, 1844), Vasum floridanuru McGinty. 1940, Lindoliva
E. J. Petuch, 1989
Page 93
K.^"^ 1
Figures 1-4. Fossil Malea species from the Pliocene and Pleistocene of soutfiern Florida. 1, 2. Malea springi new species, dorsal
and ventral views of holotype, length 189 mm, Caloosahatchee Formation, Calabrian Pleistocene, UP 21455 3. Malea petiti new-
species, dorsal view of holotype, length (incomplete) 130 mm, Bermont Formation, Aftonian Pleistocene, UP 21456. 4. Malea
densecostata (Rutsch, 1934), ventral view of 109 mm specimen, Buckingham Formation (= Pinecrest Beds) at Indian Prairie Levee,
Zanclian Pliocene. Taken from Olsson and Petit (1964, pi. 79, fig. 5).
spengleri Petuch, 1988, and Strombus (Eustrombus)
mayacensis Tucker and \\'ilson, 1933. Fragmentary spec-
imens of this large Bermont Malea were also collected
at a quarry west of Miami, in Dade County, along with
the same molluscan assemblage as that found in the cen-
tral Everglades rock pits.
In this paper, two new Floridian fossil Malea species
are described; Malea petiti n.sp. from the Bermont For-
mation and Malea springi n.sp. from the Caloosahatchee
Formation. These species, along with the Buckingham
Malea densecostata and the un-named, moldic Tamiami
species, show that four different species of Malea oc-
Page 94
THE NAUTILUS, Vol. 103, No. 3
curred in southern Florida during Plio-Pleistocene time.
The relationship of the new ta.xa to other Plio-Pleistocene
Malea species from elsewhere in the western Atlantic
are discussed under the respective descriptions. Institu-
tional abbreviations, for the deposition of type material
include: UF (Florida Museum of Natural History, Uni-
versity of Florida, Gainesville, Florida), ANSP (Paleon-
tology collection. Department of Malacology, .Academy
of Natural Sciences of Philadelphia), and F.AU (Paleon-
tolog) collection. Department of Geology, Florida At-
lantic University, Boca Raton, Florida).
SYSTEMATICS
Gastropoda
Prosobranchia
Tonnacea
Tonnidae
Tonninae
Malea Valenciennes, 1832
Malea petiti new species
(figure 3)
Material examined: HOLOTYPE — Length (incom-
plete) 130 mm, width 98 mm, dredged from appro.xi-
mately 17 m depth in Griffin Brothers road fill pit, 11
km due west of US Highwa)- 27, on Broward County-
Palm Beach County line, Florida, lower member of the
Bermont Formation, Aftonian Pleistocene, UF 21456;
PARATYPES — fragment, length 89 mm, dredged from
20 m depth in Capeletti Brothers pit mine #11, 7 km
west of Florida Turnpike, northeastern Dade County,
Florida, Bermont Formation, UF 23800; length 56 mm
(mold of juvenile), from same locality and depth as ho-
lotype, FAU 414.
Description: Shell inflated, subcylindrical, very thin and
fragile; sides of bodv whorl onK slightly rounded, giving
shell barrel-shaped appearance; shoulder slightly angled,
producing flattened subsutural area; spire (of juvenile
mold) low, flattened; body whorl (of holotype) orna-
mented with 22 wide, very flattened ribs; thin, flattened
secondary rib present betw een each pair of wide primary
ribs; secondary ribs widest and best developed on pos-
terior half of body w horl, becoming thinner and almost
obsolete on anterior half; siphonal canal proportionally
small, recurved; because of fragmentar\ nature of type
material, shape and form of outer lip, aperture, parietal
shield, and columellar region unknown.
Straiigraphic range: Known only from the lower beds
(un-named member?) of the Bermont Formation in the
Everglades Basin, Aftonian Stage of the Pleistocene.
Etymology: Named for Mr. l\ichard E. Petit, of North
Myrtle Beach, South Carolina, w ho, along with the late
Dr. A. A. Olsson, documented the first records of Malea
from southern Florida.
Discussion: Although similar in size to the Caloosa-
hatchee Malea springi, M. petiti differs in being a more
cylindrical shell with straighter sides, and in having a
distinctK lower spire and flatter subsutural area above
the shoulder. The form of the ribs and rib count also
differ between the two species; those of A/, petiti are
wider and less numerous (22 on the holot\pe), \\ hile those
of M. springi are narrower and more numerous (26 on
the holotype). Based on its overall similarit> to the older
M. springi, M. petiti is most probabK the direct de-
scendant of the finer-ribbed Caloosahatchee species. Tak-
ing into account the geologically young age and high
stratigraphic position of M. petiti, this new species may
have been the last-living Malea s.s. in North .\merica.
Malea springi new species
(figures 1, 2)
Material examined: HOLOTYPE — Length 189 mm,
width 130 mm, dredged from 20 m depth along the
Miami Canal, due west of the Talisman Sugar Refinery,
10 km north of the Broward-Palm Beach Levee, south-
western Palm Beach County, Florida, uppermost beds
(Ayers Landing Member?) of the Caloosahatchee For-
mation, Calabrian Pleistocene, UF 21455; PARA-
TYPES— length 178 mm, from same depth and locality
as holotype, ANSP 1133; length 170 mm, from same
depth and locality as holot\pe. Spring collection, Stuart,
Florida; length 156 mm. from same depth and locality
as holotype, collection of author.
Description: Shell inflated, globose, thick and heavy;
sides of body whorl distinctly rounded; shoulder and
subsutural area rounded; spire whorls elevated, slightK
protracted; suture impressed, minutely canaliculate; body
whorl ornamented with 26 (on holot\ pe) thin, flattened
ribs; small, very thin secondary ribs sometimes present
between large primary ribs, especially in midbody re-
gion; siphonal canal proportionally well-developed, re-
curved; parietal region overlaid b> wide, thick, smooth
shield; columellar notch proportionalK large, well-de-
veloped, "U"-shaped, deep; edges of notch bordered by
large knobby bosses, one on either side, that project into
aperture; outer lip very thickened, wide, with 23 (on
holot\pe) thin, elongated denticles along facing edge;
medial portion of inner edge of lip wider than rest of
lip, projecting into aperture; projecting bosses on either
side of columellar notch and wide medial area of lip
produce slightK sigmoidal shape within aperture.
Etymology: Named for Mr. Keith Spring, marine bi-
ologist at Continental Shelf, Inc., Tequesta, Florida, who
collected the holotype along the Miami Canal.
Discussion: Malea springi is most similar to the ances-
tral Pliocene M. densecostata, but differs in being a much
larger, more inflated shell with a higher, more protracted
spire, and in having finer and more numerous ribs. The
columellar notch of M. springi is proportionalK larger
than that of M. densecostata. being more indented and
wider. The knobby bosses on either side of the columellar
notch of M. springi are also larger and more developed
than those of M. densecostata. and e.xtend farther into
E. J. Petuch, 1989
Page 95
the aperture. The shape of the inner edge of the Hp also
differs between the two species; with the edge being
rounded and arcuate in M. densecostata, but subarcuate
in M. springi. being shghtly deformed by the wider
media! area that projects inwardly, into the aperture.
The new Caloosahatchee Malea also resembles the well
known M. camura Guppy, 1866, from the late Miocene-
Pliocene of the Caribbean Basin (the Bowden Formation
of Jamaica, the Gatun Formation of Panama and Costa
Rica, and the Cercado and Gurabo Formations of the
Dominican Republic), but differs in being a much larger
and more inflated shell with more numerous and thinner
ribs, and in having a much wider and better developed
columellar notch. The 35.8 mm specimen of M. camura
illustrated by Woodring (1928: pi. 20, figs. 7, 8) is typical
of the species, and can be used for comparison with M.
springi.
Malea springi also resembles M. mareana Weisbord,
1962 from the late Pliocene (early Pleistocene?) Mare
Formation of Venezuela (Weisbord, 1962). Although
similar to M. springi in shape and in having an elevated
spire, the possibly-contemporaneous M. mareana differs
in being a much smaller shell (holotype 47 mm), and in
having a proportionally smaller and narrower columellar
notch. The t\ pe of M. mareana (illustrated by Weisbord,
1962, pi. 24, figs. 1, 2) also has a sharply-angled, sub-
carinated shoulder, a feature that is missing in the dis-
tinctly round-shouldered M. springi.
At 189 mm length, the holotype of Malea springi is
the largest specimen of any of the known western At-
lantic Malea species. Previously, this title was held by a
specimen of M. goliath Pilsbry and Johnson, 1917 from
the Gurabo (Cercado?) Formation of the Dominican Re-
public, which reached a length of 129 mm (holotype).
This was followed by the Floridian specimen of M. den-
secostata illustrated bv Olsson and Petit (1964), with a
length of 109 mm. The Bermont M. petiti, with an in-
complete holotype of 130 mm, and the Caloosahatchee
M. springi, then, together probably represent the greatest
development in shell size in the fossil American Malea
species. The "Malea sp." that I illustrated previously
(Petuch, 1988, pi. 22, figs. 5, 6) is M. springi.
ACKNOWLEDGEMENTS
I thank the following for their help in collecting Malea
species in the Everglades Basin: Mr. Keith Spring, Con-
tinental Shelf Associates, Inc., Tequesta, Florida, Mr.
Jack Spengler, Lake Worth, Florida, and Mr. Donald Ash-
er, Mechanicsville, Maryland.
LITERATURE CITED
Mansfield, W. C. 1939. Notes on the Upper Tertiary and
Pleistocene moilusks of peninsular Florida. Geological Bul-
letin No. 18, State of Florida Department of Conservation,
75 p.
Olsson, A. A. and R. E. Petit. 1964. Some Neogene Mollusca
from Florida and the Carolinas. Bulletins of American
Paleontology 47(217):509-575.
Petuch, E. J. 1986. The Pliocene reefs of Miami; their geo-
morphological significance in the evolution of the Atlantic
Coastal Ridge, southeastern Florida, U.S.A. Journal of
Coastal Research 2(4):.391-408.
Petuch, E. J. 1988. Neogene history of tropical American
moilusks. Coastal Education and Research Foundation,
Charlottesville, VA, 217 p.
Weisbord, N. E. 1962. Late Cenozoic gastropods from north-
ern Venezuela. Bulletins of American Paleontology 42(193):
7-672.
Woodring, W. P. 1928. Miocene moilusks from Bowden, Ja-
maica. Carnegie Institute of Washington, Publication 385:
1-564.
THE NAUTILUS ia3(3):96-98, 1989
Page 96
The Influence of Physical Factors on the Distribution and
Abundance of Freshwater Mussels (Bivalvia: Unionidae)
in the Lower Tennessee River
Carl M. Way
Andrew C. Miller
Barry S. Payne
U.S. Army Engineer Waterways
E.xperiment Station
Environmental Laboratory
Vicksburg, MS 39180-6199, USA
ABSTRACT
Sixteen quantitative samples of mussels were taken by SCUBA
divers at each of two inshore and two offshore sites from a
mussel bed in the lower Tennessee River in July, 1987. Sedi-
mentation (measured with in situ sediment traps), sediment
type, and current velocity were measured at all sites. Fusconaia
ebena (Lea, 1831) was the dominant mussel at both the inshore
and offshore sites representing 72 and 53% of the community,
respectively. Total mussel densits . species diversity, and even-
ness were all greater at the inshore sites. Sedimentation rates
were significantly greater and current velocities were approx-
imatel) half as great (11 and 19 cm/sec, respectively) at the
inshore versus the offshore sites. We h\ potliesize that physical
factors are the dominant influence structuring this mussel com-
munity in the lower Tennessee River.
Key words: Mussels; Unionidae; sedimentation; lower Ten-
nessee River.
nessee River to water velocity, sedimentation, and sub-
strate type.
STUDY AREA
Sampling sites were located on the left bank of the Ten-
nessee River (RM 18.6), approximateK 6 km below the
Kentucky Lock and Dam. Sites 1 and 3 were 61 m apart
and 31 m from shore (inshore sites); sites 2 and 4 were
61 m apart and 61 m from the shore (offshore sites). Sites
1 and 2 were located 200 m upstream of sites 3 and 4.
Mussels have been collected from this bed, w hich appears
to extend throughout most of the river between RM 18.6
and 11.0 (Sickle, 1985), since 1931 (van der Schalie, 1939).
Current velocity at the sediment-water interface was
11.4 cm/sec (s = 1.1; N = 8) and 19.2 cm/sec (s = 1.4;
N = 8), and water depth was 3-4 m and 5-6 m at the
inshore and offshore sites, respective!) .
INTRODUCTION
Freshwater mussels (Family Unionidae), being filter-
feeders and essentially non-motile, have long been con-
sidered intolerant of poor water quality and sedimen-
tation (Hynes, 1960, 1970; Pennak, 1978). Laboratory
experiments conducted by Ellis (1933), in which he bur-
ied mussels in various types of substrates, provided early
evidence on the negative effects of sediment deposition.
In addition, Stansbery (1970) considered that sediment
from agricultural practices, reservoir construction, main-
tenance dredging, and pollution eliminated many species
of mussels. However, the distribution and abundance of
many species of mussels is at least partly dependent upon
low water velocities and low to moderate levels of sedi-
mentation for the successful settlement of glochidia.
The objective of this study was to relate community
composition and density of the dominant species (Fus-
conaia ebena) at a large mussel bed in the lower Ten-
MATERIALS AND METHODS
Sampling was conducted on July 22-23, 1987. Using
SCUBA, a diver randomly placed and secured a 16-cell
PVC grid (each cell 1 x 1 ni) to the substrate at each
sampling site. A 0.25 m- quadrat was placed in the lower
left corner of a cell and all substrate was removed by
the diver to a depth of 15 cm. Substrate was returned to
the surface in a 20 liter bucket, sieved through a nested
screen series (smallest mesh size = 6.4 mm), and all live
mussels were removed and identified to species. Sixteen
samples were taken from each of the four sites.
Sedimentation rates were measured by anchoring six
P\'C sediment traps (length : width = 25.4:2.5 cm; two
collection pipes/trap) to steel cables. Sediment traps were
placed at 3 m intervals along a 21 m transect at each of
the four sites. Sediment traps were placed in the river
by divers on JuK 10, 1986 and retrie\ed on JuK 23. 1986.
In the laboratorv, the trapped material was allowed to
C. M. Way et a/., 1989
Page 97
Table 1. A summary of the biological and physical data from the lower Tennessee River, 1986. Means in a given row with the
same letter (a, h. c) are not significaiitK differenl (p < 0.05), Densities are expressed as clams m -.
Collcci
lion site
Inshore
Inshore
Offshore
Offshore
upstream
downstream
upstream
downstream
Total number of mussels
751
604
319
295
Total number of species
17
15
18
17
Total mussel density
187.7 a
151.0 a
79.7 b
73.7 b
(SD)
(16.3)
(27.4)
(12.6)
(5.6)
Total densitv of Ftisconaia
133.3 a
111.0a
41.0b
40.0 b
(SD)
(14.5)
(21.1)
(6.9)
(3.9)
Densitv of Ftisconaia adults
74.0 a
68.0 a
22.2 b
25.2 b
(SD)
(10.4)
(12.3)
(3.7)
(1.9)
Densitv of Ftisconaia juveniles
59.2 a
43.0 b
18.7 c
19.0 c
(SD)
(7.1)
(9.5)
(4.4)
(2.5)
Species diversity (H')
1.112a
1.040 a
1.591 b
1.598 b
Evenness (J)
0.392
0.384
0.550
0.564
Sediment deposition (g cm- day ')
0.055 a
0.060 a
0.031 b
0.033 b
(SD)
(0.003)
(0.003)
(0.002)
(0.002)
Current velocity (cm/sec)
11
19
'% Sand/% gravel at 0-5 cm depth
60/32
35/61
'% Sand, "^ gravel at 5-10 cm depth
35 '20
57/37
Remaining percentage of substrate was made up of silt
settle for 24 hr, decanted, and the remaining sediment
dried for 24 hr at 100 °C and weighed.
Sediment composition was also determined at each of
the four sampling sites. Eight 10 x 15 cm cores (Miller
and Bingham, 1987) were randomly taken by divers at
each site. Cores were brought to the surface and divided
into three depth fractions (0-5, 5-10, and 10-15 cm) and
subsequentlv analyzed for particle size distribution.
RESULTS
Total mussel density was significantly greater at the in-
shore sites, although there w as no substantial difference
in total number of species between the inshore and off-
shore sites (table 1). No significant differences in any of
the remaining measured parameters were observed be-
tween upstream and downstream sites. Fusconaia ebena
was the dominant mussel at both the inshore and offshore
sites representing 72 and 53% of the community, re-
spectively. Total mussel density was two times greater
at inshore compared to offshore sites. This was due main-
ly to F. ebena. which was about three times as dense at
inshore versus offshore sites. The difference in densities
of F. ebena was significant for both large (> 30 mm
shell length) and small (< 30 mm shell length) clams
(table 1). Species diversity (Shannon-Weaver index, H';
Poole, 1974) and evenness (J; Poole, 1974) was greater
at the offshore sites. The greater species diversity and
evenness at the offshore sites was the result of a decrease
in the densities of F. ebena relative to the other species
in the community.
Sedimentation and substrate composition diftered be-
tween the inshore and offshore sites. Sediment deposition
over the 2 week period was significantly greater at the
inshore sites than at the offshore sites. Particle size anal-
ysis of inshore sediments indicated the substrate consisted
mainly of medium sand, whereas offshore sediments con-
tained higher percentages of gravel (table 1). Current
velocities at the offshore sites were approximately twice
those at the inshore sites (table 1).
DISCUSSION
Salmon and Green (1983) reported that there was an
increase in the frequency of occurrence of unionids as-
sociated with slow moving, shallow water with relatively
coarse substrate. Strayer (1983) reported that stream size
and surface geology determined the distribution of
unionids in streams in southeastern Michigan. Although
some unionids appear to be substrate specific, many are
tolerant of a wide range of substrate types (Murray &
Leonard, 1962; Parmalee, 1967; Strayer, 1981). Green
(1971, 1972) found that the distribution of Anodonta
grandis Say, 1829 and Lampsilis radiata (Gmelin, 1791)
in 32 lakes was due more to water chemistry and to
different geological conditions than to sediment char-
acteristics. Previous workers have indicated that sedi-
mentation negatively affects freshwater mussels (Ellis,
1933; Stansbery, 1970); however, their conclusions apply
to abnormal!) high levels of sedimentation (sufficient to
bury mussels) that often result from impoundment, chan-
nel modification, or disposal of dredged materials.
The various parameters that affect the distribution and
abundance of unionids (suitable fish hosts, current ve-
locity, substrate type, stream geomorphology, water
chemistry, etc.) probably have varying levels of impor-
Page 98
THE NAUTILUS. Vol. 103, No. 3
tance depending on the specific site studied. These factors
are responsible for the lack of statistical correlations be-
tween microhabitat use and unionid abundance (Strayer,
1981), but favor statistical correlations between unionid
abundance and geological conditions, water chemistry,
and substrate t\ pe. Our data show that w ithin a mu.ssel
bed in the lower Tennessee River, higher densities of
mussels (especialK' the dominant species, F. ebena) may
be associated with differences in sedimentation rate and
water velocit\ . These ph\ sical effects act upon all species
in the assemblage, but the resulting changes in com-
munity structure are due to a shift in the relative abun-
dance of the dominant species, F. ebena. We hypothesize
that physical factors are of paramount importance in
structuring this mussel community in the lower Tennes-
see River.
ACKNOWLEDGMENTS
These data were obtained as part of the Repair, Evalu-
ation, Maintenance, and Rehabilitation Program, and the
Environmental Impact Research Program, of the United
States .\rmy C^orps of Engineers by the US Army En-
gineer Waterways Experiment Station (WES). The fol-
lowing divers from the Tennessee \'alley Authority col-
lected mussels: Larry Neill, Roger Fuller, William Host
Jr., and Jim Walden. C. Rex Bingham, Ken Conley, Te-
resia Xaimo, WES, and Terry Siemsen, US Army En-
gineer District, Louisville, assisted in the field. C. A.
Miller- Way, WES, pro\ided useful comments on the ideas
presented. Permission was granted b\ the Chief of En-
gineers to publish this information.
LITERATURE CITED
Ellis, M. M. 1933. Erosion silt as a tactor in aquatic environ-
ments. Ecology 17:29-42
Green, R. H. 1971. A multivariate statistical approach to the
Hutchinsonian niche: bivalve molluscs of central Canada.
Ecology 52:543-556.
Green, R H 1972 Distribution and morpliological sariation
of Lampsilis radiata in some central C^anadian lakes: a
multivariate statistical approach. Journal of the Fisheries
Research Board of Canada 29:1565-1570.
Ilynes. H. B. .\ 1960 The biology of polluted waters. Liv-
erpool Universit\ Press, London.
Hynes, H. B. N. 1970. The ecology of running waters. Uni-
versity of Toronto Press, Toronto.
Miller, A. C. and C R. Bingham 1987. A hand-held benthic
core sampler. Journal of Freshwater Ecology 4:77-81.
Murray , H D and .\ B Leonard 1962. Handbook of unionid
mussels in Kansas. Miscellaneous Publication, University
of Kansas Museum of Natural History 28:1-184.
Parmalee, P. W. 1967. The fresh-water mussels of Illinois.
Illinois State Museum Popular Science Series 8:1-108.
Pennak, R W. 1978. Fresh-water invertebrates of the United
Stales. John Wiley and Sons. New York.
Poole, R. VV. 1974. .An introduction to quantitative ecology.
McGraw-Hill, Inc., New York.
Salmon, A. and R. H. Green. 1983. Environmental deter-
minants of unionid clam distribution in the Middle Thames
River, Ontario. Canadian Journal of Zoology 61:832-838.
Sickle, J. B. 1985. Biological assessment of the freshwater
mussels in the Kentucky Dam Tailwaters of the Tennessee
River. Report to the Kentuck\ Division of \\ ater, Frank-
furt, KY.
Stansbery, D. H. 1970. Eastern freshwater mollusks; (I) The
Mississippi and St. Lawrence River Svstems. Malacologia
10:9-22.
Strayer, D. 1981. Notes on the microhabitats of unionid mus-
sels in some Michigan streams. .American Midland Natu-
ralist 106:411-415.
Strayer, D, 1983, The effects of surface geology and stream
size on freshwater mussels (Bivalvia, Unionidae) distri-
bution in southeastern Michigan, U.S.A. Freshwater Bi-
ology 13:253-264.
van der Schalie, H. 1939. .Additional notes on the naiades
(fresh-water mussels) of the lower Tennessee River. .Amer-
ican Midland Naturalist 22:452-457.
THE NAUTILUS 103(3):99-104, 1989
Page 99
On the Distribution of Nautilus pompilius in the Samoas,
Fiji and Tonga
W. B. Saunders
Paul N. Bond
Department of Geolog)
Bryn Mawr College
Bryn Mawr, Penns> Ivania 19010, USA
Lee C. Hastie
16 C'airnie Crescent
Arbroath, Angus DDII 4DU
Scotland
David Itano
Office of Marine and Wildlife
Resources
P.O. Box 3720
Pago Pago, American Samoa 96799
ABSTRACT
Deep-water trapping off American and Western Samoa, Tonga,
and Fiji provides new data on the distribution of Nautilus in
the southwestern Indo-Pacific Thirt\-nine specimens of »V.
pompilius were trapped at 270-31U m depth in American Sa-
moa, extending the kno« n geographic range of Sautilus 1,600
km eastward. The negative results of trapping for Nautilus off
Western Samoa are inconclusive, but the lack of Nautilus in
traps containing diverse, commonly associated organisms at
290-500 m off Tonga indicates Nautilus does not occur there.
Traps set at 220-470 m off Suva, Fiji, yielded 40 -V. pompilius.
Fiji and American Samoa Nautilus exhibit some differences in
shell morphologv, but are similar to other populations of N.
pompilius in most respects.
INTRODUCTION
The easternmost occurrence of living Nautilus has gen-
eralK been regarded as Fiji, from whence the first doc-
umented hve specimen was obtained during trawHng by
the Challenger E.xpedition (Moseley, 1892), and where
a number of studies have been completed in recent years
(e.g., Hayasaka. 1985; Zann, 1984; Muntz & Raj, 1984).
Until now, in spite of numerous recent reports of Nau-
tilus at more westerly locales (see Saunders, 1987, for
review), there has been no knowledge of whether this
organism occurs further east than Fiji. The present re-
port, based on trapping efforts during 1986 in .\merican
Samoa, Western Samoa, Tonga, and Fiji, extends the
range of Nautilus eastward by more than 1,600 km (fig-
ure 1). The find of Nautilus in American Samoa suggests
that its actual range may pro\ e to be considerably greater
than has been surmised on the basis of drifted shells,
because in that area there was neither knowledge of the
living animal nor of the shells — in fact, there is appar-
ently no Samoan word for Nautilus. Nevertheless, ani-
mals were trapped outside Pago Pago, literally within
sight of the Governor's residence. Subsequent trapping
efforts for Nautilus in Western Samoa and Tonga yielded
no Sautilus, but trap yields from south of Suva, Fiji
prov ide new data on depth distribution and on organisms
associated with Nautilus in that region. In a sense, this
report marks the first effort to delimit the geographic
range of living Nautilus. Following is a brief account of
the new occurrence in Samoa, along with observations
on morphological differences, ecological conditions and
associated organisms at the other sites where deep-water
trapping was undertaken.
MATERIALS AND METHODS
Deep-water trapping for Nautilus followed procedures
developed in Palau (Saunders & Spinosa, 1978). It in-
volved setting funnel-ended, rectangular fish traps (1 x
1 X 2 m) baited \\ith skipjack tuna at bottom forereef
sites, with depth and topographic selection determined
by echofinder. The traps were marked at the surface
with buoys, and were left out for 1-2 nights at depths
ranging from 100-500 m. Following is a summary of
locations, and yields:
1. American Samoa: Traps were set three times, during
July, 1986, along Taema Bank, southeast of Pago Pago
Harbor, Tutuila Island (figure 1), for 1-2 nights at depths
of 270, 280, and 310 m. Individual trap yields varied
from 3-29 specimens of Nautilus pompilius plus mis-
cellaneous fishes and invertebrates (table 1). During
March, 1987, additional catches totalling 16 specimens
of Nautilus pompilius were reported by National Marine
Fisheries Service personnel, conducting deep-water trap-
ping at 300-400 m in the same area.
2. Western Samoa: Traps were set three times, ap-
proximately 6 km north of .^pia Harbor, L^polu Island
(figure 1). In this region, the bottom descends gradually
to a depth of ca. 100 m, w here it becomes a sheer, vertical
face that extends to se\ eral thousand meters — well below
Nautilus depth. This configuration made it impossible
to set bottom traps at optimal Nautilus depths. Traps set
at ca. 100 m \ielded no Nautilus, and only a few spec-
imens of teleosts. Because of the proximity to .American
Samoa (approximately 130 km eastward), where Nau-
tilus appears to be abundant, conclusions regarding the
absence of Nautilus in Western Samoa based on the
present work are premature.
Page 100
THE NAUTILUS, \ol. 103, No. 3
Taema Bank
AM. SAMOA
W. SAMOA
' AM. SAMOA
^ 'C^~^ff
Figure 1. Map showing deep-water trapping sites (asterisks)
in ,'^merican and Western Samoa, Tonga, and Fiji. Nautilus
was obtained in Fiji and .\merican Samoa.
3. Tonga: Two traps were set overniglit si.\ times, in-
and just outside Piha Passage, 5-16 km off Nukualofa,
Tongatapu Island (figure 1), during July, 1986 at 280-
500 m depth. ,\lthough no Mautilus were trapped, a
diverse assemblage of organisms was obtained, that are
typically found associated w ith Sautilus at other locales
(table 1); we conclude that \autilus is not present in this
region of Tonga.
4. Fiji: Traps were set overnight fi\e times, 0.8-3.5 km
southwest of Suva at depths of 220-470 m. A total of 40
specimens of \. pompilius w as obtained, along with nu-
merous shrimps and deep water teleosts (table 1).
Reference specimens of Sautilus, as well as associated
organisms, have been reposited at the National Museum
of Natural Histor\ , Smithsonian Institution. Washington,
D.C. (USNM 816658-816659, 816704-186709); the Brit-
ish Museum (Natural Histor> ), London; the J. P. Hayden
Museum, Pago Pago (specs. AS 33, 35); and with the
Office of Fisheries, Nukualofa, Tonga.
RESULTS
The general consensus has been that the genus Nautilus
is morphologically conservative and exhibits little vari-
ation (e.g., Ward, 1984). However, recent multivariate
analyses of a large number of morphological characters
(Swan & Saunders, 1987) shows that the most common
and widespread species. A', pompilius, exhibits consid-
erable morphological differentiation in geographically
isolated populations. In addition, genetic anaKses of the
same populations using electrophoretic surveys of protein
polymorphisms (Woodruff et al., 1987) show that the
species also exhibits a high level of genetic variation.
Comparative studies of other li\ ing Sautilus populations
are currently being undertaken that should permit more
definite taxonomic assessments in the near future. Fol-
lowing is a summar) of some of the differences observed
between the Fijian and Samoan populations.
Table I. Deep-water trap yields of cephalopods, fishes and shrimps in American Samoa, Western Samoa, Tonga and Fiji, based
on trapping conducted in 19Sfr Shrimp identifications follow King (1984).
.\ssoc. species
.\m. Samoa
W Samoa
1 onga
F,,i
Depth (m)
Cephalopods
Nautilus pompilius
Octopus sp.
Fishes
Conger sp.
Muraenesox cinereus
Epinephelus sp.
Etelis carlmnculus
Tridon macropterus
Pristopimoides mtdlidans
SqiHilus sp.
Shrimps
Heterocarpus ensijcr
II gillhoHHS
H. sihogac
Parapandalus serratifrons
Plesionika longirostris
Pi. martin
270-310
x(39)
-100
280-500
220-470
x(40)
W, B. Saunders et al., 1989
Page 101
I. N. pompilius, American Samoa
Shell size, maturity, and sex ratios: The 39 specimens
available range in size from 106.1-180.3 mm shell di-
ameter, but mature specimens range from 163-179.2 mm
diameter (figure 2; table 2). This size range is similar to
that reported for A', pompilius from Tanon Strait, Phil-
ippines, and from Papua New Guinea (Saunders & Davis,
1985; Saunders, 1987; Saunders et al., 19S7). .\s in vir-
tually all other populations of Nautilus for which data
are available, mature males are larger (5%) and heavier
(18%) than females (mean diameter for males 174.5 mm;
mean weight 861.3 g; females 164.9 mm; 703 g), and
males (19.5%) substantiall) outnumber females (see also
Saunders & Spinosa, 1978; Hayasaka, 1983, 1985). Al-
though mature animals typically comprise 75% of Nau-
tilus populations, onl\ 28.2% of the Samoan specimens
are fully mature; this probabK reflects the small sample
size.
Distinguishing characteristics: In basic shell form, the
Samoan Nautilus are indistinguishable from the wideK
distributed species X. pompilius. How ever, several fea-
tures of coloration and sculpture are distinctive, and war-
rant description.
Shell coloration is more prominent than in other pop-
ulations of this species, in that the stripes are wider and
more numerous; i.e., the proportion of striped (brown)
dorsal shell is considerably greater than unstriped (white)
shell (figures 3-7). In addition, the umbilical area is brown,
and the color bands tend to show a zig-zag pattern on
the flanks of the shell. Although numerical analysis of
these features will be required to evaluate the degree of
uniqueness compared to other populations, both stand in
contrast to the general trend toward reduced shell col-
oration, including a white umbilical region, in shells from
the southern part of the range of N. pompilius — notably
in Papua New Guinea and northwestern .Australia.
The shells from Samoa also exhibit delicate but dis-
tinctive longitudinal (concentric) sculpture (figure 5). This
feature is developed more strongh- (and was cited as a
species characteristic) in \. belauensis b\' Saunders (1981),
and it is even more strongly developed in N. scrobicu-
AMERICAN SAMOA
Immature (n=28)
mature (n=l 1)
-I 1 1 h-
115 125 135 145 155
Shell diameter (mm)
FIJI
175 185
Shell diameter (mm)
Figure 2. Frequency distribution of live-caught specimens of
.V, pompilius from American Samoa and Fiji, showing size
range (maximum shell diameter) and proportion of mature
individuals (see Table 2 for additional data).
latus, but it is not t\pical of N. pompilius, N. stenom-
phalus or N. macromphalus.
II. N. pompilius, Fiji
Shell size, maturity, and sex ratios: The 40 specimens
available range from 80.7-155 mm diameter, but 22
Table 2. Morphologic data from live-caught .V. pompilius s. I. from .\merican Samoa and Fiji, arranged by sex, showing range
in variation in mature animals and sexual dimorphism (shell width measured beneath the ocular sinus; total weight, shell plus body
weii;ht in air)
Shell diameter (mm)
Shell width (mm)
Total weight (g)
Sex
Range
Mean
SD
Range
Mean
SD
Range
Mean
SD
II.
Nautilus pompilius, .•\merican Samoa
Females (n = .3) 16.3-166 164.9
Males (n = 8) 168.6-179.2 174.5
Total (n = 11) 163-179.2 171.9
Nautilus pompilius. Fiji
Females (n = 2) 136.5 136.5
Males (n = 20) 138.8-155 146
TotaKn = 22) 136,5-1.55 145.1
1.65 68-72.8 69.8 2.6
4.1 76.7-84.3 81 2.3
5.7 68-84.3 77.9 5.7
0 58.1-58.2 58.2 1
4.2 60.7-74.1 68 3 1
4.9 58.1-74.1 67 1 41
680-740 703.3 32.2
800-925 861.3 51.1
680-925 818.2 86.5
400-410 405 7.1
4.35-670 528 56
400-670 516.8 64.5
Page 102
THE NAUTILUS, Vol. 103, No. 3
n
r i"--
n
* ■¥■
^
10
N
W. B. Saunders et al., 1989
Page 103
mature specimens range from 136.5-155 mm diameter
(figure 2; table 2). Only two specimens are mature fe-
males; they are 136.5 mm diameter, and are smaller
(6.5%) than the mature males (mean 146 mm; n = 20),
and they weigh 23% less (mean 405 g females vs. 528 g
males). The proportion of mature animals (55%) is some-
what less than usual for Nautilus populations, but the
high overall proportion of males (80%) is typical. Con-
siderable additional comparative morphologic data are
available for the Fiji Nautilus, in published accounts by
Ward et al. (1977), Ward & Martin (1980), Zann (1984)
and Ha\asaka (1985).
Distinguishing characteristics: Given the range of vari-
ation in size and color patterns that have now been doc-
umented in Nautilus poinpilius (see Saunders & Davis,
1985; Saunders, 1987; Saunders & Swan, 1987) it is not
surprising that the Fijian population exhibits some mor-
phologic differences compared to other populations for
which data are available. Mature shell size (mean 145.1
mm) is small compared to most populations; mean ma-
ture size of Tanon Straits, Philippines, specimens is 165
mm, and mature size for Papua New Guinea populations
ranges from 144 mm (Lae) to 169 mm (Kavieng; Saun-
ders & Davis, 1985). In addition, one population of even
smaller mature shells (mean 114 mm) is known from the
Sulu Sea (Saunders, 1987).
The coloration of the Fiji specimens is generally coarser,
(i.e., relativeK fewer but broader stripes) than in typical
N. pompilius. and in man\' specimens, color bands to not
extend to the umbilicus, leaving a white umbilical area
(figures 8-10). The latter feature seems also to be more
common in southern Papua New Guinea populations and
in specimens of N. pompilius from the Great Barrier
Reef.
An electrophoretic survey has recently been completed
on the Fiji populations (Woodruff et al., 1987). The re-
sults indicate that the Fiji population is genetically well
differentiated from populations of \. pompilius in Papua
New Guinea and the Great Barrier Reef. This is perhaps
not so surprising, given the geographic distances between
the samples, and analysis of geographicalK intermediate
populations w ill be required to evaluate the significance
of the genetic differentiation recorded. In this regard, it
will be of considerable interest to compare the results of
electrophoretic studies (now underway) of the Samoan
population, which is even further removed geographi-
cally, to the Fiji and other populations.
ACKNOWLEDGMENTS
The efforts of fisheries officers and personnel in .American
and Western Samoa, Tonga, and Fiji were instrumental
in obtaining the results reported here. In particular, we
wish to acknowledge the assistance of Ray Tulafono (Di-
rector), Raymond Buckley, and the fisheries staff at Pago
Pago, American Samoa. In Western Samoa, Lui Bell,
Fisheries Biologist, Apia, provided unlimited assistance.
In Tonga, Viliami and Sarah Longi, and Malcolm
McGregor assisted immeasurably, along with others at
the Office of Fisheries in Nuku alofa (Semisi Fakahu,
Director). Deep-water trapping in Fiji was undertaken
with support of Dr. Peter Hunt, Chief Fisheries Officer,
Tui Lancala and Malakai Tuilou, also of Fisheries; and
we owe particular thanks to Keith Meecham, Programme
Director, South Pacific Regional Fisheries Development
Program, Suva, and to Robert Gillett, Fisheries Devel-
opment Advisor.
The Nautilus project relied heavily on the support and
assistance of Operation Raleigh (U.K.) personnel on board
SES Sir Walter Raleigh. In particular, we wish to ac-
knowledge the efforts of P. David King (Expedition
Leader), Captain Mike Kichenside, First Officer Malcolm
Philips, Assistant Expedition Leader Tony Walton and
venturers Byron White, Roy Jarvis, and Tony Wong;
Matt Richmond assisted w ith shrimp identifications. Sup-
ported b\ National Science Foundation grant BSR 86-
08065.
LITERATURE CITED
Hayasaka, S. (ed.) 1983, Studies on Nautilus pompilius and
its associated fauna from Tanon Strait, the Philippines.
Kagoshima University Research Center of the South Pa-
cific, Occasional Papers, No. 1.
Hayasaka, S., (ed.) 1985. Marine ecological studies on the
habitat of Nautilus pompilius in the environs of Viti Levu,
Fiji. Kagoshima L^niversit) Research Center of the South
Pacific, Occasional Papers, No. 4.
King, M. G. 1984. The species and depth distribution of deep-
water caridean shrimps (Decapoda, Caridea) near some
southwest Pacific islands. Crustaceana 47:174-191.
Moseley, H. N. 1892. Notes by a naturalist. An account of
observations made during the voyage of "H.M.S. Chal-
lenger" around the world in the years 1872-1876. John
Murray, London, 540 p.
Muntz, W. R. A. and U. Raj. 1984. On the visual system of
Nautilus pompilius. Journal of Experimental Biolog) 109:
253-263.
Saunders, \V. B. 1981. The species of living Nautilus and
their distribution. X'eliger 24:8-17.
Saunders, W. B. 1987. The species of Nautilus. In: Saunders,
W. B. and N. H. Landman (eds). Nautilus: the biology
and paleobiolog) of a living fossil. Plenum Press, New
York and London, p. 35-52.
Saunders, W. B. and L. E. Da\is. 1985. A preliminary report
on Nautilus in Papua New Guinea. Science in New Guinea
11:60-69.
Figures 3-10. Nautilus pompilius Linne, 1758. 3-7. Specimens from American Samoa. 8-10. Specimens from Fiji. 3. Living
animal (USNM 816658) held in an aquarium ('/3 x ); 4, 6. 7. Lateral views of mature (4, USNM 816708; 7, USNM 816709) and
immature (6, USNM 816707) shells (V2 x ); 5. Detail of shell surface of specimen in figure 7, showing delicate longitudinal sculpture
(3.0 X). 8-10. Lateral views of immature (8, USNM 816706) and mature shells (9, USNM 816705; 10, USNM 816704) ('2 x ).
Page 104
THE NAUTILUS, Vol. 103, No. 3
Saunders, W B and C," Spinosa 1978. Se.xual dimorphism in
Saulilwi from Patau Paleobiology 4:.'349-358.
Saunders, W B.. Davis, L. E., and R. L. Knight. 1987. Sym-
patric species of S'autilus (.V. pompilius and A', scrobic-
ulattis) in the .-Kdmiralty Islands, Papua New Guinea. The
Nautilus 101:92-99.
Swan, A. R. H, and VV. B. Saunders. 1987. Morphologic vari-
ation in Sauttlus from Papua New Guinea. In: Saunders,
W , B and N H Landman (eds.). Sautilus. the biology
and paleobiolog) of a living fossil. Plenum Press, New
York and London, p. 8.5-103.
Ward, P. D. 1984. Is Sautilus a living fossil'-" In: Eldredge,
N. and S. M. Stanley (eds. ). Living fossils. Springer \'erlag.
New York, p. 247-256.
Ward. P D and .\ W Martin. 1980 Depth distribution of
Sautilus pompilius in Fiji and Sautilus macromphalus
in New (Caledonia. X'eliger 22:259-264.
Ward, P. D., Stone, R., Westermann, G , and .A. Martin 1977
Notes on animal weight, cameral fluids, swimming speed,
and color polymorphism of the cephalopod Sautilus pom-
pilius in the Fiji Islands. Paleobiologx 3:3.'37-388.
W oodruff, D. S., Garpenter, M. P., Saunders, W B., and P D
W ard. 1987. Genetic variation and ph\logen\ in Sau-
tilus. In: Saunders, W. B and N H Landman (eds.).
Sautilus. the biology and paleobiology ol a li\ing fossil.
Plenum Press, New York and London, p. 65-83.
Zann, L. P 1984. The rhythmic activity of Sautilus pom-
pilius with notes on its ecology and behaviour in Fiji.
Veliger 27:19-28.
THE NAUTILUS 103(3):105-108, 1989
Page 105
Repaired Shell Damage in a Complex of Rissoid Gastropods
from the Upper Continental Slope South of New England
Faigel K. Vale
Michael A. Rex
Department of Biologx'
University of Massachusetts at Boston
Boston. MA 02125, USA
Several theories have invoked predation as a potentially
important factor structuring deep-sea benthic commu-
nities (review ed in Jumars and Eckman, 1983; Rex, 1983;
Grassle, 1989), but there is little direct evidence for this.
In coastal faunas, the incidence of repaired shell damage
in snails, when critically and carefully interpreted
(Schoener, 1979; Vermeij,' 1982a; Schindel et ai, 1982),
has been a useful measure of the relative importance of
predation. In this note we analyze patterns of repaired
shell damage in an assemblage of four rissoid gastropods
from the upper continental slope south of New England.
The four species, Frigidoalvania brychia (Verrill, 1884),
Onoba pelagica (Stimpson, 1851), Pusillina harpa (Ver-
rill, 1880), and Pusillina pseudoareolata (Waren, 1974),
are all deposit feeders with nonplanktotrophic devel-
opment and similar ventricose shells. Rex et al. (1988)
recently presented a biometrical analysis of shell form
in the assemblage that revealed strong differentiation
with depth across the upper slope. Predation was sug-
gested, but not explored further, as a possible cause of
this depth-related geographic variation. Our aim here is
to test hypotheses that shell-repair frequency corresponds
to: 1, depth of sampling and hence the observed depth-
related clinal effects, and 2, differences in shell archi-
tecture that in coastal species are associated with the
relative ability to deter predators.
Station data and species lists for the five samples are
provided in table 1 (see Rex et al., 1988, for details of
Table 1. Station data, sample sizes, incidence and statistical analysis of repaired shell damage for rissoid snails from the upper
continental slope south of New England (USA). Chi-square tests compare the frequency of repaired damage (using raw data)
between Frigidoalvania brychia and the other three species combined, and between rugose and smooth forms of F. brychia in
station 87
Fre-
Station
Latitude
Longitude
Depth
Sample
Number
quenc\
Chi-
Signifi-
number
(N)
(W)
(m)
Species
size
broken
broken
square
cance
88
.39°.54,1'
70°37.0'
478
Onoba pelagica
Pusillina pseudoareolata
Pusillina harpa
3
6
11
0
0
0
0
0
0
Frigidoalvania brychia
17
2
0.12
0.719
n.s.
96
39°55.2'
70°39.5'
498
Onoba pelagica
Pusillina pseudoareolata
Pusillina harpa
12
12
79
0
1
7
0
0.08
0,09
Frigidoalvania brychia
122
16
0.13
1,162
n,s.
105
39°56.6'
71°03.6'
530
Onoba pelagica
Pusillina harpa
44
84
4
3
0.09
0.04
Frigidoalvania brychia
155
33
0.21
13,186
P < 0,001
207
39°51..3'
70°54.3'
808
Onoba pelagica
Pusillina pseudoareolata
Pusillina harpa
3
SlOO
SlOO
1
20
12
0,33
0,20
0,12
Frigidoalvania brychia
SlOO
27
0,27
4,217
P < 0,05
87
39°48.7'
70°40.8'
1,102
Frigidoalvania brychia
Rugose form
S200
53
41
8
0,21
0,15
Smooth form
147
33
0,22
0,881
n,s.
S before
sample size
indicates that
a random subsample (rather than all
appropriate
shells at that station)
was scored.
Page 106
THE NAUTILUS, Vol. 103, No. 3
Figure§ 1-12. Specimens of rissoid gastropods collected from the upper continental slope south of New England ^USA): 1.
Frigidoalvania hrychia showing repaired shell damage at the end of the second whorl (station 105, shell height is 3.5 mm). 2.
Onoba pelagica (sta. 207, 2.3 mm). 3. Pusillina harpa (sta. 96, 2.0 mm). 4. Pusillina pseudoareolata (sta. 96, 2.2 mm). 5-8. Rugose
forms of F. brychia (sta. 87; 3.6, 3.3, 3.8, 3.7 mm respectively). 9-12. Smooth forms of F. brychia (sta. 87; 3.7, 3.5, 3.2, 3.5 mm
respectively). See Table 1 for station data.
sampling and a bathymetric map showing station local-
ities). Individuals were scored for repaired shell damage
using the criteria developed by Schindel et al. (1982)
and Vermeij (1982a). We recorded only what Vale and
Rex (1988:65) termed "major" damage: "conspicuous
breaks generally resulting in displacement of subsequent
growth patterns and interruption of sculpture." An ex-
ample of such damage in Frigidoalvania brychia is shown
in figure 1. EssentialK the same criteria have been used
to identify predator-induced shell breakage in numerous
F. K. Valen and M. A. Rex, 1989
Page 107
shallow-water snail faunas (see Vale and Rex, 1988, for
a review). We scored all individuals with at least a half
post-larval whorl, and recorded the number of repaired
breaks for the first, second and third v\horls (shells of
these species seldom have a complete third whorl) to
determine whether there was any variation in repaired
damage with shell size. If the samples were very large
(636-5,318 individuals, see Rex et ai, 1988; table 1), we
scored 100-200 randomK- selected individuals. The in-
cidence of shell repair was calculated as simply the fre-
quency of repaired shells (Raffaelli, 1978). An alternative
index is the average number of scars per shell (Vermeij,
1982b). The measures are nearly identical for the rissoid
complex because onl\- 5% of the shells that were repaired
had more than one break.
The number and frequency of repaired breaks are
gi\en in Table 1. We report data for the entire adult
shell (i.e., whorls 1-3 combined) because no consistent
differences in frequency were associated with shell
growth. The median frequency of repaired damage for
the whole complex in all 5 samples is 0.11 (0.13 if only
large, \ > 40, samples are used). This value is near the
median value obtained for the deep-sea prosobranch fau-
na as a whole (0.15) by Vale and Rex (1988), and falls
within the range of values reported for a wide variety
of shallow-water habitats (see \'ale and Rex, 1988: table
2).
We used chi-square tests to determine whether there
is any association between sampling depths [about 500
m (stations 88, 96, and 105 combined), 800 m, and 1,100
m] and frequenc) of shell repair. The null hypothesis is
that frequencies of repair are equal among snails grouped
by depth. The alternative h\ pothesis is that depth groups
differ in frequency of repair. None of the rissoids show
a significant difference in the incidence of repaired shell
damage among sampling depths (chi-square values range
from 2.30-4.38, 0.20 > P > 0.05, two-tailed test, Siegel,
1956).
We also explored whether the frequency of repaired
damage is associated \\ ith shell architecture. Frigidoal-
vania brychia exhibits more rugose sculpture than the
other three species (figures 1-4). Its heavy shoulder knobs
and strong spiral costae are identical to sculptural fea-
tures that deter predators in shallow- water snails (Palm-
er, 1979). At the 500 m stations all individuals of F.
brychia are heavily armored like the individual shown
in figure 1, but at greater depths variation increases to
include both rugose forms and smoother forms (figures
5-12). Rex et al. (1988) interpreted the increased vari-
ation in deeper populations of F. brychia to be a possible
case of competitive release. It was also suggested that
heavy sculpture on F. brychia might confer greater pro-
tection against predation than the less sculptured shells
of the other species. Frigidoalvania brychia might also
be less vulnerable to predation because of its larger size
(Rex et al, 1988: table 2, and figures 1-12 herein). To
test whether this might be reflected in the frequency of
repaired shell damage, we compared the incidence of
repair between F. brychia and the otlier three species
combined at sites where they are sympatric, and between
rugose (figures 5-8) and smooth (figures 9-12) forms of
F. brychia at the 1,102 m site. The sculptural variation
in F. brychia at 1,102 m is continuous. Following Rex et
al. (1988), we used a qualitative scale to score shells as
either rugose (in the range of figures 5-8) or smooth (in
the range of figures 9-12).
Whether rugose forms should have a higher or lower
frequency of shell repair than smooth forms is difficult
to predict. The relationship of predation intensity to in-
cidence of repair depends on a complex set of life-history
features and population dynamics of both predators and
prev, and evolved responses to predator-prey interactions
(Schoener, 1979; Schindel et al, 1982; Vermeij, 1982a,
1983). Virtually none of this information is known for
either the rissoids or their likeK predators, the decapod
crustaceans and fishes. We can test only in a general way
whether rugose and smooth shell forms experience pre-
dation differently. Two-tailed chi-square tests on the fre-
quency of repair between rugose and smooth forms of
Frigidoalvania brychia and between F. brychia and the
other rissoids do not reveal a consistent pattern (table 1).
Within F. brychia there is no significant difference in
repair frequency between smooth and rugose forms.
Among the species, F. brychia generally shows a higher
frequency of repair, but the difference is strongK- sig-
nificant in onK one of the 500 m stations (sta. 105), and
weakly significant at 808 m (sta. 207).
Our results show that the rissoid assemblage is sub-
jected to a comparatively high level of potentially lethal
predation. However, there is no clear correspondence of
the incidence of repaired shell damage to the depth-
related patterns of geographic variation observed by Rex
et al. (1988). Nor is there convincing evidence that dif-
ferences in shell size and architecture relate to predation.
These findings are consistent with the hypothesis of Ver-
meij (1978) and Vale and Rex (1988) that, while crushing
predation imposes a certain level of mortality on deep-
sea snails, relationships between gastropod prey and
crushing predators are more generalized and poorly co-
evolved than they appear to be in their shallow-water
counterparts.
We thank Ron Etter and .Andrea Rex for reading the
manuscript. Mary Smith photographed the specimens in
figures 1-12. The gastropods were collected by vessels of
the Woods Hole Oceanographic Institution and were
made available to us by Howard Sanders.
LITERATURE CITED
Grassle, J. F. 1989. Species diversity in deep-sea communities.
Trends in Ecology and E\olution 4:12-15.
Jumars, P. A. and J. E. Eckman. 1983. Spatial structure within
deep-sea benthic communities. In. Rowe, G. T. (ed). Deep-
sea biology. Wiley, New York, p. 399-451.
Palmer, .\. R. 1979. Fish predation and the evolution of
gastropod shell sculpture: experimental and geographic
evidence. Evolution ■33:697-713.
Raffaelli, D. G. 1978. The relationship between shell injuries,
Page 108
THE NAUTILUS, Vol. 103, No. 3
shell thickness and habitat characteristics of the intertidal
snail Littorina rujis Maton. Journal of Molluscan Studies
44:166-170.
Rex, M. \. 1983. Geographic patterns of species diversity in
the deep-sea benthos In: Rowe, G. T. (ed.). Deep-sea
biology \Vile\. .New York, p. 45.'3-472.
Rex, M. .\ , M Campbell Watts, R J Etter, and S. O'Neill
1988. Character \ariation in a complex of rissoid gastro-
pods from the upper continental slope of the western North
Atlantic. Malacologia 29:325-340.
Schindel. D. E., G. J. \'ermeij, and E. Zipser. 1982. Fre-
cjuencies of repaired shell fractures among Pennsylvanian
gastropods of north-central Texas. Journal of Paleontology
56:729-740.
Schoener, T. W. 1979. Inferring the properties of predation
and other injurs -producing agents from injury frequen-
cies. Ecology 60; 1 1 1 0- 11 15.
Siegel, S. 1956. Nonparametric statistics for the behavioral
sciences. McGraw-Hill, New York, 312 p.
\'ale, F. K. and M. A. Rex. 1988. Repaired shell damage in
deep-sea prosobranch gastropods from the western North
Atlantic Malacologia 28:6.5-79.
N'ermeij, G J. 1978. Biogeography and adaptation. Har\ard
Universit) Press, Cambridge, M.\, 332 p.
X'ermeij, G. J. 1982a. En\ ironmental change and the evolu-
tionary history of the periwinkle [Littorina littorea) in
North .•Vmerica. Evolution 36:561-580.
Vermeij, G. J. 1982b. Gastropod shell form, breakage and
repair in relation to predation by the crab Calappa. Mal-
acologia 23:1-12.
Vermeij, G. J. 1983. Shell-breaking predation through time.
In: Tevesz. M.J.S and McCall, P L. (eds.). Biotic inter-
actions in Recent and fossil benthic communities. Plenum,
New York, p. 649-669.
THE NAUTILUS 103(3):109-112, 1989
Page 109
Distribution of Melampus bidentatus (Say) and
Succinea wilsoni (Lea) within a Tidal Marsh in
Eastern Connecticut
Bradford H. Burnhani
Paul E. Fell
Department of Zoology
Connecticut College
New London, CT 06320, USA
Succinea ivihoni Lea 1864 and Melampus bidentatus
Say 1822 are halophilic plumonate snails which occa-
sionalK share the same microhabitat (Fell & Williams,
1985). Melampus occurs primariK in the higher salinity-
regions of salt marshes, its population density being gen-
erally low in brackish areas (Parker, 1976; Fell & Wil-
liams, 1985). Succinea is found in freshwater marshes
and the less saline regions of salt marshes (Grimm, 1975).
The present report describes the distribution of Melam-
pus and Succinea within a single marsh system in Con-
necticut,
MATERIALS AND METHODS
This study was conducted on the Paffard Marsh which
is located in Stonington, Connecticut, and is situated
north of Route 1 along the upper extent of a tidal creek
(Oxecosset Brook) that empties into Fishers Island Sound
at the eastern end of Long Island Sound (figure 1). The
marsh is cut b\ a series of mosquito ditches that drain
into the creek. Throughout much of the marsh, most of
the interditch area is covered by stunted Spartina alter-
niflora Loisel. 1807, Tall Spartina alterniflora grows along
the banks of the ditches; and a belt of Spartina patens
(Ait) Muhl 1817 frequently occurs between the ditches
and the interditch Spartina alterniflora. Distichlis spi-
cata (L,) Greene 1887 and Juncus gerardi Loisel. 1809
are found in patches within the marsh, being most abun-
dant near the upland border. Phragmites australis (Cav.)
Trin ex Steud. 1820 and Eleocharis rostellata Torr. 1843
also occur along the edge of the marsh. At the upper
ends of the two major branches of the tidal creek, there
are large stands of Typha angustifolia L. 1753. During
mid-JuK- the salinit> at the upper end of the western
branch of the tidal creek was 1-2 parts per thousand
(ppt) and that of the creek at Route 1 was 27 ppt.
The study was conducted from 6 June through 22 July
1987, The densities of the snails at 55 stations were de-
termined using a 50 cm square wooden frame, 9 cm
high, which was tossed onto the marsh in areas chosen
for study. The vegetation within the frame was clipped
at the surface of the peat and all of the snails were
collected and counted. The shell lengths of the snails
were measured to the nearest 0.5 mm under a dissecting
microscope using a mm ruler. The salinity of the soil
water was determined at 44 stations by squeezing water
from a piece of peat, filtering the water through What-
man No, 1 filter paper, and measuring the salinity with
a Goldberg refractometer.
Spatial overlap between Melampus bidentatus and
Succinea wilsoni was calculated using the equation:
C = 1 - >/2[y(PAi - PBi)] (Schoener, 1970)
Where C is the spatial overlap, PAi is the percentage of
the total number of species A {Succinea wilsoni) found
in a particular microhabitat category (i), and PBi is the
percentage of the total number of species B (Melampus
bidentatus) occurring in the same microhabitat category
(i). Each of the 55 quadrats was treated as a separate
microhabitat category. A value for C of 0 indicates that
there is no spatial overlap betw een the two species, while
a value of 1 indicates that there is complete overlap.
Voucher specimens of Melampus bidentatus (USNM
#858073) and Succinea wilsoni (USNM #858074) col-
lected on the Paffard Marsh have been deposited at the
National Museum of Natural History, Smithsonian In-
stitution,
RESULTS
Melampus occurred at greatest densities where the soil
water salinity ranged between 20 ppt and 30 ppt (figure
2) and in regions dominated b\' stunted Spartina alter-
niflora and Distichlis spicata (figure 4), The density of
Melampus was weakl) correlated with soil water salinity
(r = 0,420), Succinea was most abundant at stations where
the soil water salinity varied from 15 ppt to 22 ppt (figure
3) and where the vegetation consisted of Eleocharis or
Typha (figure 4). It exhibited the highest densities at the
upper ends of the two major branches of the tidal creek
Page 110
THE NAUTILUS, Vol. 103, \o. 3
Figure 1. Map of the Paffard Marsh, in Stonington, Connecticut, showing the 55 stud)' sites. Squares represent sites that contained
25 or more Melampus bidentatus per 0.25 sq. meter; triangles show sites that contained 25 or more Succinea wilsoni per 0.25 sq.
meter; and circles represent sites where only a few snails were found. One site contained 25 or more of both species of snail per
0.25 sq. meter. The inset (lower right) shows the location of the Paffard Marsh (arrow) on the Connecticut shore
350
in
CM
(A
3
a.
E
M
»
s
250 -
150
50
0
B °
a a n B B B a a a 3
1 0
20
— I —
30
350
o
a>
c
o
o
3
250-
150
50
0
° G
qqBI^Iqbo
o a a
1 0
2 0
3 0
Salinity (ppt)
Figure 2. Densities of Mclanipus bidentatus in relation to soil-
water salinities on the Paffard Marsh in Stonington, Connect-
icut.
Salinity (ppt)
Figure ,3. Densities of Succinea wilsoni in relation to soil-
water salinities on the Paffard Marsh in Stonington, Connect-
icut.
B. H. Burnham and P. E. Fell, 1989
Page 111
in
in
CM
(0
c
V)
6
c
>
<
(5)
Q Melampus bidentatus EBB
00 -
B Succinea wilsoni nli
80-
1
(9) (5) 1
60 -
40 -
20-
n -
(5
)
1
i
n
1
-
(7
(19)
(5)
1
Table 1. Abundance, no. per 0.25 m- (mean ± S.D., range),
ot Melampus bidentatus (Say) and Succinea wilsoni (Lea) at
55 sampling sites on the Paffard Marsh in Connecticut.
Snail abundance
Number of
quadrats
Many Melampus (83 ± 44, 33-155) only 10
Many Melampus (63 ± 39, 32-126);
few .Succinea (1.4 ± 0.9, 1-3) 5
Man\ Succinea (101 ± 77, 29-306) only 11
Many Succinea (40 ± 15, 27-55);
few Melampus (10 ± 9, 4-20) 3
Few Melampus (6 and 15) only 2
Few Succinea (9 ± 7, 1-21) only 8
Few Melampus (5.4 ± 5.7, 1-18);
few Succinea (4.5 ± 3.2, 1-12) 13
Many Melampus (27); many Succinea (69) 1
No snails 2
sl.S.a w SI b a m [i ..' jL'-r,.;s s pa! bieoch lypha
Vegetation
Figure 4. .\yerage densities of Melan^pus bidentatus and Suc-
cinea wilsoni in different \ egetation types on the Paffard Marsh
in Stonington, Connecticut. The number of quadrats studied
in each yegetation type is indicated in parentheses, st. S.a. w
= stunted Spartina alterniflora with standing water; st. S.a. m
= stunted Spartina alterniflora with moist soil; Dist. = Dis-
tichlis spicata. Juncus = Juncus gerardi: S. pat. = Spartina
patens, Eleoch. = Eleocharis rostellata, Typha = Typha an-
gustifolia.
(figure 1). Density oi Succinea was negatively correlated
with soil water salinity (r = —0.534).
Spatial overlap between Melampus and Succinea was
calculated to be 0.07 indicating little spatial overlap be-
tween the two species. Melampus and Succinea occurred
together in 22 of the 55 quadrats that were examined;
however, in only one quadrat were both species present
in large numbers and in 36% of these quadrats one species
or the other predominated (Table 1). The snails tended
to occur together in regions of the marsh covered by
Distichlis. Juncus and/or Spartina patens.
Succinea ranged in size from 1 mm to 8 mm, but most
of the snails were small The largest size class was 2-3
mm. Melampus ranged in size from 4 mm to 12 mm,
with 9-10 mm snails constituting the largest size class.
Melampus egg masses were observed occasionally from
6 July through 21 July 1987. The egg masses were found
in areas of the marsh covered by stunted Spartina al-
terniflora, Spartina patens and Distichlis and where soil
salinities ranged from 17 ppt to 29 ppt. No Succinea eggs
were observed in the field. However, adult Succinea were
kept in the laboratory and observed for egg laying. Be-
ginning on 6 June, ten adult Succinea averaging 7 mm
in length were kept in a plastic container with wet paper
towels and fed iceberg lettuce. Egg la\ ing was observed
on 22 June. The eggs occurred singly or in clusters of up
to five capsules. They were 1 mm in diameter, with the
early embryo being about 0.1 mm in diameter and the
remainder of the capsule being filled with albumen. Al-
though no eggs hatched, the embryos developed for up
to two weeks. At the end of this period, the young snails
had filled the entire egg capsule.
DISCUSSION
The distribution of Melampus bidentatus observed in
this study is consistent with the pattern described in
previous reports (Leathem & Mauer, 1975; Parker, 1976;
Fell et a/., 1982; Fell & Williams, 1985). This snail, which
is restricted to the high marsh, extends far up estuaries
into regions of low salinity; but its population density is
low where salinities fall below about 10 ppt (Parker,
1976; Fell & Williams, 1985). Succinea wilsoni was found
primarily in the more brackish regions of the marsh as
predicted by earlier studies. Grimm (1975) states that
this snail usually occurs in the freshest zone of saltmarshes
but that it occasionalK ma> be found in shaded fresh
water swamps or in more saline environments. Melam-
pus and Succinea were found occurring together in 40%
of the quadrats examined, but in only one of the quadrats
were both of them present in large numbers (>25 per
0.25 sq. meter). Although these snails occupy some of
the same microhabitats, they evidently exhibit little spa-
tial overlap. There appears to be only one previous report
of the co-occurrence of Melampus and Succinea in salt
marshes (Fell & Williams, 1985). However, such an as-
sociation of these snails is relatively common in brackish
marshes in Connecticut (unpublished observations). Fur-
ther studies of Melampus and Sitccinea in other brackish
marshes should provide important information concern-
ing the factors that determine the distributions of these
snails.
ACKNOWLEDGEMENTS
We are indebted to the Mashantucket Land Trust for
permission to conduct the study in Paffard Marsh. This
work was supported by Research Experience for Un-
Page 112
THE NAUTILUS, Vol. 103, No. 3
dergraduates Grant No. BBS-8713347 from the National
Science Foundation to Connecticut College.
LITERATURE CITED
Fell, P E., N. C. Olmstead, E. Carlson, W. Jacob, D. Hitchcock,
and G. Silber. 19S2. Distribution aiul abundance of mac-
roin\ertebrateson certain (Joiinectieut tidal marshes, with
an emphasis on the dominant molluscs. Estuaries 5;2'34-
239.
Fell, P. E. and ]. H Williams. 1985. Distribution of the snail
Melampus hidentatus, and the mussel, Geukensia demis-
sa, along the Petaguanset Estuary (Connecticut) in relation
to salinity and other tidal marsh invertebrates. The Nau-
tilus 99:21-28.
Grimm, F. W. 1975. .\ review of Succinea ailsoni a coastal
marsh snail of eastern North .\merica The Nautilus 89:
39-43.
Leathern, VV. and D. .\laurer. 1975. The distribution and
ecolog\ of common marine and estuaririe gastropods in
the Delaware Ba\ area. The .Nautilus 89:73-79.
Parker. X. H. 1976. The distribution, growth and life history
of Melampus hidentatus (Gastropoda: Pulmonata) in the
Delaware Ba> region. .Masters thesis. University of Dela-
ware, 65 pp.
Schoener, T. \^^ 1970. Nonsynchronous spatial overlap of
lizards in patchy habitats. Ecology 51:408-418.
THE NAUTILUS 103(3):113-115, 1989
Page 113
The "Gray Catalogues" [Mollusca] of the British Museum
Alan R. Kabat
Museum of Comparative Zoc
Harvard Universits
Cambridge, MA 02138 USA
logy
John Edward Gray (1800-75) was the longtime Keeper
of Zoology at the British Museum (Montagu House,
Bloomsbury); in 1824 he was appointed as an assistant
and became a full keeper in 1840 (following the death
of J. C. Children). Gray played an important role in the
development of systematic zoology in England, and his
research greatly enhanced the value of the zoological
collections now housed in the Natural History Museum
[formerly known as the "British Museum (Natural His-
tory)"; i.e.. South Kensington]; further biographical de-
tail is in Gunther (1975, 1980) and Smith (1906).
Gray authored over 1,160 publications on all the major
groups of animals of which a tabulation is provided in
Gray (1875); this list, however, contains numerous small
errors and omissions. Gray edited an extensive series of
catalogues of the natural history collections contained in
the British Museum. These catalogues covered the whole
spectrum of systematic zoology and are of great impor-
tance as they contain generic diagnoses and descriptions
of new taxa. The first such, the "Synopsis of the Contents
of the British Museum" (from 1808 to 1856, in 63 edi-
tions), were rather cursory overviews of the entire col-
lection; the more specific "Lists ..." and "Catalogues
..." are unquestionably of greater importance today.
Gunther (1912) discussed the history, rationale, and
limitations (specifically, the few illustrations provided)
of these various publications. Needless to say, Gray did
not write all of these catalogues, although he did provide
prefaces for many of those authored by his colleagues.
Sherborn (1926a, 1934) provided a valuable, albeit ab-
breviated, collation of these publications, with the dates
of publication determined by when "they were laid upon
the table of the Trustees ..." (Sherborn, 1926a:271).
These dates show some slight variance from those pro-
vided in the "Catalogue of Books, Manuscripts, Maps,
and Drawings of the British Museum (Natural History)"
(Woodward, 1903b).
In the field of systematic malacology, the "Gray Cat-
alogues" are still most useful in providing a guide to the
taxa of Gray, Baird, Deshayes, d Orbigny and Pfeiffer,
as well as of the pioneering collections from Cuba, the
Canaries, South America and Mazatlan, and of the var-
ious molluscan taxa monographed. Nonetheless, it must
be admitted that some of the taxa listed were nude names
(lacking a description) and were probably subsequently
(re-)described by Gray or another author. In particular,
many of the generic names tabulated in the various edi-
tions of the "Synopsis of the Contents of the British Mu-
seum ..." were not made available until Gray's 1847 "A
List of the Genera of Recent Mollusca . . . ," as discussed
by Iredale (1913); all of Gray's works should be carefully
consulted with respect to these generic names. A number
of family-group names were also made available in these
catalogues.
Due to the slight variations in previous citations of the
dates of publication of these catalogues, some confusion
has inevitably arisen over the years. Some authors, ap-
parently unaware of Sherborn's determinations, have
based the dates upon those printed on the title page or
following the editorial preface. However, the actual date
that should be used is, in some cases, the following year.
For example, Ruhoff (1980) not only provided several
incorrect dates, but also listed Gray as the author when
he was merely the editor of a colleague's work.
Herein is presented as complete a collation as possible
of the 22 catalogues of the British Museum that contain
treatments of the Mollusca. They are listed by author,
in chronological order (based on the dates of Sherborn,
1926a, 1934), with the editor (if difi^erent) and "date of
publication in text' noted at the end of each citation.
The publisher is the British Museum; the printer is also
given. As some readers may be aware. Carpenter's 1857
"Catalogue of the Collection of Mazatlan Shells ..." was
simultaneously printed under a separate title, on differ-
ent paper of a larger size. The two printings differ only
in their title, prefatory material, and dimensions. All the
other British Museum catalogues are approximately 16.0
cm X 10.3 cm; the "Catalogue of the Reigen Collection
of Mazatlan Mollusca ..." is 17.5 cm x 10.2 cm (sizes
may vary due to trimming and binding).
LIST OF CATALOGUES
Baird, William. 1850 [12 June]. Nomenclature of Molluscous
Animals and Shells in the Collection of the British Museum.
Part I. Cvclophoridae. Spottiswoodes and Shaw, London,
69 p. [Preface by J. E. Gray, "March, 1850."]
Carpenter, Philip Pearsall. 1857 [1 .■\ugust]. Catalogue of the
Collection of Mazatlan Shells, in the British Museum: Col-
lected by Frederick Reigen. Oberlin Press, Warrington,
Page 114
THE NAUTILUS, Vol. 103, No. 3
xii + .552 p (Some copies have the pagination of intro-
ductory material as "i-iv, ix-xvi."]
Carpenter, P. P. 1857 [1 .August]. Catalogue of the Reigen
Collection of Mazatian Mollusca, in the British Museum.
Oberlin Press, Warrington, viii + xii + 552 p. [Preface,
J. E. Gray, "April 22nd, 1857."] [The title page gives the
date as "1855-7"; this refers to the printing of the sheets
but not their distribution, i.e., "publication" (Iredale, 1916:
36).] I have seen five copies of this and the preceding title,
each with slight variations as to the prefatory material;
only one copy had both Carpenter s and Gray's prefaces
(both prefaces are dated "April 22nd, 1857"). [Plates pub-
lished b\ Brann (1966); text reprinted, 1967, Paleontolog-
ical Research Institution, Ithaca.]
Deshayes, Gerard Paul 1853 [25 June]. Catalogue of the Con-
chifera or Bivalve Shells in the Collection of the British
Museum. Part I. X'eneridae, CXprinidae and Glauco-
nomidae. Taylor and Francis, London, [ii] -I- 216 p. [In-
troduction b\ J. E , Gray, "June 27, 1853"; note that
Sherboni, 1934, gave an earlier date; one must be in error]
[Part II. 1855]
Deshayes, G. P. 1855 [12 Ma\]. Catalogue of the Conchifera
or Bivalve Shells in the Collection of the British Museum.
Part II. Petricoladae (concluded); Corbiculadae. Taylor
and Francis, London, p. 217-292. [Title page, "1854."]
[Part I, 1853]
Gray, John Edward 184()a. Mollusks, In: Synopsis of the
Contents of the British Museum, 41st ed. G. VV'oodfall,
London, p. 82-84. [ii] + 302 p. ["27 June 1840."]
Gray, J. E. 1840b, Mollusks. In: Synopsis of the Contents of
the British Museum, 42nd ed. G. Woodfall and Son, Lon-
don, [iv] -I- 370 p. [In three issues: (I) p. 105-152 (16 Oct.
1840); (2) p. 86-89 & p. 106-156 (4 Nov. 1840); (3) p.
106-156 (1841); see Sherborn, I926b:99.]
Gray, J. E 1841 Mollusks. In: Synopsis of the Contents of
the British Museum. 43rd ed. G. VV'oodfall and Son, Lon-
don, p. 78-130. 159-162. [ii] + 382 p. ["June 17, 1841."]
Gray, J. E. 1842. Mollusks. In: Synopsis of the Contents of
the British Museum, 44th ed. G. Woodfall and Son, Lon-
don, p. 48-92. [iv] -I- 308 p. ("21 May 1942."]
Gray, J. E. 1849 [30 June]. Catalogue of the Mollusca in the
Collection of the British Museum. Part I. Cephalopoda
Antepedia. Spottiswoodes and Shaw, London, viii + 164
p. [Preface, "I2th February, 1849,"]
Gray, J. E. 1850a [9 February], Catalogue of the Mollusca in
the Collection of the British Museum, Part II, Pteropoda,
Edward Newman, London, iv + 45 p, [Preface, "11th
January, 1850"]
Gray, J. E. 1850b (6 July] Catalogue of the Bivalve Mollusca
in the Collection of the British Museum. Part I. Placen-
tadae and .-Vnomiadae. Edward .Newman, London, 22 p.
[No preface.]
Gray, J, E, 1851 [22 November], List of the Specimens of
British .-Vnimals in the Collection of the British Museum,
Part \'I1, Mollusca .Acephala and Brachiopoda Richard
Taylor, London, iv -I- 167 p, [Preface undated] [Reprint
titled as "List of British Mollusca and shells, with synon-
yma, in the collection of the British Museum, Part I,"]
Gray, J. E. 1854a [9 December], List of the Shells of the
Canaries in the Collection of the British Museum, Col-
lected by MM, Webb and Berllielot Described and Fig-
ured by Prof, .\lcide d'Orbigny in the "Histoire Naturelle
des lies Canaries. Taylor and Francis, London, 32 p,
[Preface, "1 Sept, 1854,"]
Gray, J. E. 1854b [9 December]. List ol the Shells of Cuba
in the Collection of the British Museum. Collected by M.
Ramon de la Sagra. Described b\ Prof, .■\lcide D'Orbigny
ui the "Histoire de I'lle de Cuba. " Taylor and Francis,
London, [ii] -I- 48 p. [Preface, "Sept. 1, 1854 ."]
Gray, J. E. 1855a (13 January]. List of the Shells of South
.■Vmerica in the Collection of the British Museum. Col-
lected and described b\ M. .-Vlcide d Orbigny , in the "Voy-
age dans r.\merique Meridionale " Taylor and Francis,
London, [ii] + 89 p. [Preface, "I Oct, 1854."]
Gray, J. E. 1855b [13 January], List of Mollusca and Shells
in the Collection of the British Museum, collected and
described by MM, Eydoux and Souleyet, in the "Voyage
autour du Monde," execute pendant les annees 1836 et
1937, sur la Corvette 'La Bonite," and in the "Histoire
naturelle des Mollusques Pteropodes, " par MM. P.-C.-A.-L.
Rang et Souleyet Taylor and Francis, London, 27 p, [Pref-
ace, "January II, 1855. ']
Gray, J, E, 1855c [8 December], List of Mollusca in the Col-
lection of the British Museum, Part I, V'olutidae, Tavlor
and Francis, London, [ii] + 23 p, [Preface, "5 Nov. 1855. "]
[Part II, 1865.]
Gray, J. E. 1857 [9 May ]. Guide to the Systematic Distribution
of Mollusca in the British Museum. Part I [Gastropoda].
Taylor and Francis, London, xii -I- 230 p , 121 text-figs.
[Preface, "Dec. 10, 1856,"']
Gray, J E, 1865 [11 March] List of the Mollusca in the
Collection of the British Museum, Part II Olividae, Taylor
and Francis, London, 41 p. [Preface. J, E, Grav, "Dec.
1864,""] [Part I, 1855]
Pfeiffer, Ludwig Georg Carl [= '"Louis "], 1853 [12 February].
Catalogue of Phaneropneumona, or Terrestrial Opercu-
lated Mollusca, in the Collection of the British Museum.
Woodfall and Kinder, London, [ii] + 324 p, [Introduction
by J. E, Gray, ""I6th September, 1852.""]
Pfeiffer, L. 1855 [14 .-Vpril], Catalogue of Puhnonata or .\\i-
Breathing Mollusca in the Collection of the British Mu-
seum. Part I, Taslor and Francis, London, [ii] -I- 192 p.,
5 text-figs. [Preface, J. E. Gray, '"29 March, 1855"; see
Woodward, 1903a:261,]
Pfeiffer, L, 1857 [10 October] Catalogue of Auriculidae, Pro-
serpinidae, and Truncatellidae in the Collection of the
British Museum Taylor and Francis, London, [ii] + 150
p., text-figs. [Preface, J. E. Gray, undated]
ACKNOWLEDGEMENTS
Kenneth J. Boss, Eugene V. Coan and Richard E. Petit
provided helpful reviews of the manuscript. M. G. Ha-
rasewych supplied a photocopy of Gray (1851) from the
Smithsonian Institution Library.
LITERATURE CITED
Brann, D. C, 1966, Illustrations to ""Catalogue of the Collection
of Mazatian Shells"" b\ Philip P Carpenter. Paleontological
Research Institution, Ithaca, 111 p [inci, pis, 1-60],
Gray, J. E. 1847. .\ list of the genera of the Recent Mollusca,
their synony ma and t\ pes. Proceedings of the Zoological
Society of London 1847:129-219,
Gray, J. E. 1875, List of the books, memoirs, and miscella-
neous papers of Dr, John Edward Gray, FR.S., with a few
historical notes, Taylor and Francis, London, 58 p.
Giinther, A. 1912. General history of the Department of Zo-
A. R. Rabat, 1989
Page 115
olog) from 1856 to 1895. The history of the colloftions
contained in the natural hislor\ departments of the British
Museum. \'o\ II Appendix British Museum, London, x
+ 109 p.
Gunther, A. E. 1975. A century of zoology at the British
Museum through the Hves of two keepers, 1815-1914.
Dawsons, London, 533 p.
Gunther, .\. E. 1980. The founders of science at the British
Museum. Halesworth Press, Suffolk, x + 219 p.
Iredale, T. 1913. A collation of the moiiuscan parts of the
S\nopses of the Contents of the British Museum, 1838-
1845. Proceedings of the Malacological Society of London
10l4):294-309.
Iredale, T. 1916. On some new and old moUuscan generic
names. Proceedings of the Malacological Society of Lon-
don 12(l):27-37.
Ruhoff, F. A. 1980. Index to the species of Mollusca intro-
duced from 1850 to 1870. Smithsonian Contributions to
Zoolog) 294:iv + 640 p.
Sherborn, C. D. 1926a. Dates of publication of earl\' Cata-
logues of Natural History issued by the British Museum.
The .\nnals and Magazine of Natural History (ser. 9) 17(98):
271-272.
Sherborn, C. D. 1926b. Note on the 42nd edition of the
Synopsis of the Contents of the British Museum, 1849.
Proceedings of the Malacological Society of London 17(2-
3):99.
Sherborn, C. D. 1934. Dates of publication of Catalogues of
Natural History (post 1850) issued by the British Museum.
The .\nnals and Magazine of Natural History (ser. 10)
13(74):308-312.
Smith, E. A. 1906. Mollusca In: The history of the collections
contained in the natural histor\ departments of the British
Museum, \'ol. IL Separate historical accounts of the several
collections included in the Department of Zoology British
Museum, London, p. 701-730. 782 p.
Woodward, B. B. 1903a. Note on the dates of publication of
J. E. Gray's "Catalogue of Pulmonata ... in the . . . British
Museum. Pt. I," 12mo, 1855; and of A. Moquin-Tandon's
"Histoire Naturelle des Mollusques Terrestres et Fluvia-
tiles de France, " 2 vols. 8vo, 1855. Proceedings of the
Malacological Society of London 5(4):261.
Woodward, B. B. 1903b. Catalogue of Books, Manuscripts,
Maps, and Draw ings of the British Museum (Natural His-
tory), Vol. I. .A-D. Hazell, VN'atsonand Viney, Ld., London,
viii + 500 p.
THE NAUTILUS 103(3): 116, 1989
Page 116
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THEt^NAUTILUS
CONTENTS
Volume 103, Number 4
April 18, 1990
ISSN 0028-1344
Harald A. Rehder
Clarification of the identity of the snail
groenlandicus (Gmelin, 1791) (Gastropoda
Mar4a|5(^$ne Biological Laboratory
' rochidae) LIBRARY
MAY 2 m\}
Two new species of Chicoreus subgenus Stratus
(Gastropoda: Muricidae) from northeastern Brazil I J4
I Woods Hole, Mass.
Emily H. Yokes
\ .,.. ,
William k. Emerson
^ alter E. Sage, III
Distorsio ridens (Reeve, 1844): a synonym of Distorsio
clathrata (Lamarck, 1816). (Gastropoda: Personidae)
131
kurt J. Jirka
Richard J. Neves
Freshwater mussel fauna (Bivalvia: Unionidae) of the New
River Gorge National River, West Virginia
136
Eva Pip
Copper, lead and cadmium concentrations in a sample of
Lake Winnipeg Anodonta grandis
140
kurt Auffenberg
Roger W. Portell
A new fossil land snail (Gastropoda: Pulmonata:
Polygyridae) from the Middle Miocene of northern Florida
143
THE NAUTILI'S 103(4): 117-123, 1990
Page 117
Clarification of the Identity of the Snail
Margarites groenlandicus (Gmelin, 1791)
(Gastropoda: Trochidae)
Harald A. Rehder
Department of Invertebrate Zoology
National Museum of Natural Hi.stor\'
Smithsonian Institution
Washington, DC 20560, USA
ABSTRACT
The boreal Atlantic trochidA/argan7esgroen/an(f!C!W (Gmelin,
1791) is generally regarded as a species ranging from Massa-
chusetts. Scotland and Norway north to polar waters, and vary-
ing in sculpture from smooth to strongK lirate. A critical ex-
amination of many specimens and of the putative syntypes of
Trochus groenlandicus Chemnitz, ITcSl, the basis of Gmelin s
name, revealed that two species are involved in what has been
known as Margarites groenlandicus: (1) the typical A/, groen-
landicus, with a usually larger, smooth, more broadly conical
shell, found only in the colder Arctic and polar waters, and (2)
M. striatus (Leach, 1819), with a smaller, more elevated, spi-
rally lirate shell with a narrower umbilicus, found in northern
Europe and Iceland in the eastern Atlantic, and on the western
.•Mlantic coast from southern Massachusetts northward to south-
ern Baffin Land and Disko in western Greenland. Biometric
and geographic data are adduced to support the distinctiveness
of these two taxa
INTRODUCTION
In the process of preparing a field guide to North Amer-
ican marine shells, I found that there is confusion over
the true identity of Margarites groenlandicus (Gmehn,
1791), a species found in the cold waters of the North
Atlantic.
In "American Seashells" (Abbott, 1974:36), Margarites
groenlandicus is described as having "whorls with about
a dozen smooth spiral Hrations or almost entirely smooth
(form urnlfilicalis Broderip and Sowerby, 1829) ', and its
geographic range is given as "Arctic seas to Massachusetts
Bay." Examination of material in the collection of the
National Museum of Natural History revealed that two
distinct taxa are involved, and that the smooth form
occurs no farther south than northernmost Labrador and
the southern tip of Greenland. A study of material on
which Chemnitz's Trochus groenlandicus was probably
based shows that this name should be applied to the
smooth, broadly umbilicate, .\rctic form.
The true taxonomy of Margarites groenlandicus and
Margarites striatus (Leach, 1819), the species with which
it has been confused, is reviewed in the following syn-
onymies and discussions.
SYSTEMATICS
Margarites groenlandicus (Gmelin, 1791)
(figures 1-7, 13)
Trochus Gronlandicus umhilicatus anfractibus . . . etc. —
Chemnitz. 1781:108. pi. 171, fig. 1671 (type locality, north-
ern Greenland). [Non-binomial].
Trochus groenlandicus Gmehn, 1791:3574; Dillwyn, 1817:771.
Margarita umbilicalis Broderip and Sowerby, 1829:379; Sow-
erbv, 1838a:26 (tvpe locality, Melville Island); Sowerby,
1838b:pl. 133, fig! 5; Reeve,' 1855:393; Sowerby II, 1878:
pi. 1, fig. 1; Posselt, 1895:90.
Margarita sulcata Sowerbv, 1838a:26 (tvpe locality. Melville
Island); Sowerby, 1838b:pl. 132, fig.' 1.
Margarita groenlandica, Sowerbv, 1838a:25; Sowerbv, 1838b:
pi. 133. fig. 10; Morch. 1857:89; Sowerbv II, 1878:pl. 1,
fig. 6: Odhner, 1912:56-62 (in part), pi. 4, figs. 15, 16, 18,
19; Thorson, 1944:15-20
Margarita undulata laevior Moller, 1842:81 (type localit\,
Greenland).
Trochus umbilicalis, Philippi, 1852:245. pi. 37, fig. 2; Jeffreys,
1877:237.
Trochus gronlandicus, Philippi, 1852:247, pi. 337, fig. 5.
Trochus rossi, Philippi, 1852:288, pi. 43. fig. 9 [new name for
Margarita sidcata Sowerby, 1838a. not Trochus sulcatus
Lamarck, 1804, etc.].
Margarites umbilicalis, Pilsbry, 1890:288, pi. 3, figs. 61, 62,
64, pi. 64, figs. 39-41; Baker, 1919:503-504; Johnson, 1934:
72.
Margarites umbilicalis spiralis. Baker, 1919:503-504 (type lo-
cality, Etah, CJreenland).
Margarites groenlandicus umbilicus, Odhner, 1915:149; .Ab-
bott, 1954:108, fig. 31d; Galkin, 19.55:82-83, fig. 25; Ab-
bott, 1974:.36. fig. 215.
Margarita groenlandica, var. laevigata Odhner. 1915:149 (type
localit). Isfjorden. West Spitsbergen).
Diagnosis: Shell reaching 24.85 mm in width, 16.85
mm in height, glossy, broadly conical, occasionally some-
Page 118
THE NAUTILUS, Vol. 103, No. 4
Figure 1. Lectotype of Margarites groenlandicus (Gmelin,
1791) Diameter 13.75 mm Zoologisk Museum Copenhagen,
Denmark.
what depressed; suture deeply impressed; color light olive
gray to light grayish brown (Kelly & Judd, 1965); early
whorls usually strongly and spirally lirate, but may be
weakly lirate, with shallow grooves, or smooth; later whorls
smooth or with fine, impressed, spiral lines; umbilicus
rather broad, deep; rarely are specimens found that are
dull and strongly lirate on all whorls, but otherwise typ-
ical in form and umbilical characters.
Discussion: Chemnitz (1781:109) described a species he
called "Der Gronlandische Krausel, Trochus Gronlandi-
cus iimhilicatus, anfractibus rotundatis, intus margari-
taceus". It is figured rather poorly on plate 171, figure
1671.
Chemnitz described the shell as thin, pearly under a
thin, flesh-colored layer, the six rounded whorls with very
delicate spiral lines, and with a broad, deep umbilicus.
The figure depicts a shell about 17mm wide, broadly
conical and rather elevated, with numerous fine, inter-
rupted spiral lines on the rounded whorls. The locality
is given as "from the most distant shores of cold, raw
Greenland", and the specimens are designated as "ex
museo nostra". The brief description and poor figure
have led to a misunderstanding of the identity of Chem-
nitz's species.
Chemnitz was never in Greenland, and according to
J. Knudsen (in litt.) he probably received these specimens
from Otto Fabricius, a missionary in Greenland from
1768 to 1773, and from 1774 to 1779 a pastor in southern
Norway, a country at that time united with Denmark.
He returned to Denmark and in 1783 received a post in
Copenhagen, where he spent the rest of his life. While
residing in Norway Fabricius worked on his "Fauna
Groenlandica", published in 1781, and it seems very
likely that during this time he was in correspondence
^^^-i^-e^X^fCih^
Zoologigk Museum. Kebenhavn.
nan
■ lo:i
Figure 2. Labels present with syntypes of Margarites groen-
landicus (Gmelin). Label in handwriting of O. A. Morcli.
with Chemnitz, a relationship that undoubtedly became
closer when he returned to Copenhagen.
Chemnitz's name was validated by Gmelin as Trochus
groenlandicus, and listed under this name b\ Dillwyn
in his "Descriptive Catalogue" (1817). Neither Chemnitz
nor Gmelin gave any measurements but Dillwyn gives
the dimensions as "about seven lines long and eight
broad". This is equivalent to 14.8 mm height and 17 mm
width, almost exactly the dimensions of Chemnitz's fig-
ure.
Through the kindness of Dr. Jorgen Knudsen I was
able to examine two lots of specimens labeled Trochus
groenlandicus and Turbo groenlandicus from the col-
lections of the Zoologisk Museum in Copenhagen. One
lot, comprising ten specimens, has a laliel written, ac-
cording to Knudsen, about 40 years ago which has the
word "type" on it. .Accompanying it is an older label,
written by Morch, which states that the specimens came
from Fabricius. Near the top of the label is the name
"Turbo groenlandicus nob. " in a slightly different hand-
writing, and below it is a reference to (Chemnitz's de-
scription and figure (figure 2).
Since this lot appears to consist of specimens received
from Fabricius, and ma>' very well have been seen by
Chemnitz, I feel justified in selecting one of these spec-
imens (figure 1) as the lectotype.
Examination of this material shows that Margarites
groenlandicus (Gmelin) is the earliest valid name for the
species called Margarites umlnlicalis Broderip & Sow-
H. A. Rehder, 1990
Page 119
Figures 3-7. Shells of Margarites groenlandicus (Gmelin), showing apertural, dorsal, and ventral views. 1.5 x 3. Sabine Id., E.
Greenland (USNM 219180). 4. Clavering Id., E. Greenland (USNM 406156). 5. Cumberland Gulf, Baffin Land (USNM 219181).
6. Coburg Id., S of Ellesmere Land (USNM 466586). 7. Cape Sabine, Ellesmere Land (USNM 126754),
erby by Pilsbry and Margarites groenlandicus form ttm-
bilicalis by Abbott (1974).
This species is restricted to the cold Arctic waters from
Victoria Island, northern Canada, eastward to Franz Jo-
seph Land and south to the northern tip of Labrador and
to the southern point of Greenland (figure 6). A more
detailed discussion of the geographic range of this species
is given below.
Margarites striatus (Leach, 1819)
(figures 8-13)
Trochus cincerarius Fabricius, 1780:391 (Greenland) [not T.
cinerarius Linne, 1758].
Margarita striata Leach, 1819:464 (type locality, Baffin Bay);
Gray, 1826:567.
Turbo carneus Lowe, 1825:107-108, pi. 5, figs. 12-13 (type
locality, Argyllshire, Scotland); Gray, 1826:567.
Trochus margaritus Gray, 1826:567 [new name for Turbo car-
neus Lowe, 1825].
Margarita carnea, Sowerby, 1838a:25, Sowerby, 1838b: 1, pi,
133, fig. 9.
Margarita undulata Sowerby, 1838a:26 (type locality, Arctic
Seas); Sowerby, 1838b:l', pi. 132, fig. 4; Gould, 1841:254,
fig. 162 = 172; Reeve, 1842:169, pi. 221, fig. 4; M6ller,
1842:81; Binney, 1870:280, fig. 341; Sowerby H, 1878:pl.
1, fig. 2; Pilsbry, 1890:290, pi. 39, figs. 36-39, pi. 64, figs.
42-44.
Turbo incarnatus Couthouy, 1838:98, pi. 3, fig. 13 (type lo-
cality, Phillips Beach, Massachusetts).
Margarita undulata trochijorntis Moller, 1842:81 [new name
for Trochus cinerarius Fabricius, 1780, not Linne 1758].
Trochus leachii Philippi, 1852:247, pi. 37, fig. 6 [new name for
Margarita striata Leach, 1819, not Trochus straitus Linne
1767].
Trochus fabricii Philippi, 1852:284, pi. 42, fig. 2 [new name
for Trochus cinerarius Fabricius, 1780, not Linne, 1758].
Trochus undulatus. Forbes and Hanley, 1853:528, pi. 68, figs.
1, 2, pi. 73, figs, 5, 6.
Margarita groenlandica var. undulata, Morch, 1857:89.
Margarita cinerea, Morch, 1857:89 [not Turbo cinerea Couth-
ouy 1838].
Trochus groenlandicus, Jeffreys, 1865:298-299, pi. 61, fig. 5
[not Trochus groenlandicus Gmelin, 1791].
Margarita incarnata. Sowerby II, 1878: pi. 3, fig. 18.
Margarita groenlandica. Sars,' 1878:133-134; Posselt, 1895:80;
Hagg, 1905:9-13; Odhner, 1912:56-62 [in part], pi 4, figs.
4-14, 17, 20-27; Thorson 1941:13-14 [not Trochus groen-
landicus Gmelin 1791].
Margarites groenlandicus, Johnson, 1915:88; Winkworth, 1932:
Page 120
THE NAUTILUS, Vol. 103, No. 4
Figures 8- 12. Shells of Margarites strialus (Leach), showing apertural, dorsal, and ventral views. 1.5 x 8. Aiigmagssalik, Greenland
(USNM 466647). 9. Spitsbergen (USNM 181706). 10. Spitsbergen (USNM 181736). 1 1. Schooner Cover, Labrador (USNM 604949):
12. Seal Harbor, Maine (USNM 438215).
220; Johnson, 1934:72; Abbott, 1954:108; Galkin, 1955:80-
82, fig. 24; Abbott, 1974:36; Sneli, 1975:36, fig. 50; Fretter
and Graham, 197742, fig 28 [not Trochus groenlandicus
Gmelin. 17911
Margarites undulata (Sowerbv) = groenlandica, Nordsiek, 1968:
17, pi, 3, fig, 09,01,
Margarites striatus. Rehder, 1981:370, fig. 300.
Diagnosis: Shell reaching 15.5 mm in width, 14.75 mm
in height, dull, moderately thick-shelled, broadly to el-
evated-conical; suture not impressed; color light grayish
yellowish brown to light brown (Kelly & Judd, 1965);
protoconch smooth, early post-nuclear whorls with spiral
grooves or ridges rarely smoothish, later whorls strongly
ridged, usually with short, low broad, wave-like axial
ridges below the suture; body whorl slightly angled at
the periphery; base somewhat flattened, with fine, low,
spiral cords; umbilicus narrow, funnel-shaped, deep; ap-
erture showing low, spiral ridges within the outer lip,
corresponding to the external spiral .sculpture.
Discussion: Margarites striatus is distinguished from
M. groenlandicus by its generally heavier and more el-
evated shell, its smaller size, stronger spiral .sculpture, a
suture that is not impressed, and usually by the presence
of broad, axial ridges below the suture, which are never
found in M. groenlandicus. The base of M. striatus is
flattened, with numerous fine spiral lirae, and the um-
bilicus is narrower and obscurely angled at the edge.
Meaurements of width, height, and the height /width
ratio of 50 specimens of each species, randomK chosen
from lots collected from throughout the range of each
species, are given in condensed form in Table 1. These
figures clearly show that M. striatus has a generally
smaller, more elevated shell. The largest specimens of
M. groenlandicus are less elevated than the smaller or
average-sized specimens of the same species. The larger
shells of M. striatus tend to be slightly less elevated than
those of average size.
Misidentification of these two species seems to have
started with Jeffreys (1865:298), who gave the spirally
Urate specimens found in western Scotland and the Ork-
ney and Shetland Islands the name Trochus groenlan-
dicus Chemnitz.
Before Jeffreys (1865), Margarites striatus had gen-
erally been known as Margarita (or Trochus) undulata
Sowerby, as the synonymy given above shows. Pilsbry
(1890:290-291) continued to call this species, found from
Massachusetts to northern Labrador, southern Green-
H. A. Rehder, 1990
Page 121
Figure 13. Map of the North Polar regions, northeastern North
America, and northern Europe, showing distribution of Mar-
garites groenlandicus and Margarites striatiis.
Table 1 . Summary of measurements of 50 specimens of each
species. Measurements in mm
groenlandicus striatus
Largest shell
Diameter (d)
Height (h)
h/d
24.85
16.85
0.68
15.85
14.75
0.93
.Smallest Shell
Diameter (d)
Height (h)
h/d
7.90
6.65
0.84
6.90
6.00
0.97
Average
Diameter (d)
Height (h)
h/d
14.54
10.92
0.75
9.43
8.48
0.90
Mean
Diameter (d)
Height (h)
h/d
13.82
10.75
0.78
9.15
8.37
0.91
5 largest shells h/d
0,72
(JS6
land, and northern Europe, Margarita undulata, and
included in the synonymy of this species Chemnitz's
Trochus groenlandicus umbilicatus, although he did not
accept the name because it was nonbinomial. He over-
looked Gmelin's validation of the Chemnitz name, and
did not follow Jeffreys' use of groenlandicus because it
was preoccupied by Trochus gronlandicus Philippi, 1852,
which is the same, of course, as Chemnitz's species.
The species with larger, more depressed shells with a
wider umbilicus, rounded whorls, and with the last whorl
smooth or with fine, incised lines was called Trochus
umbilicalis Broderip and Sowerby by Jeffreys (1877:237),
who stated that, in his opinion, it was distinct from what
he called M. groenlandicus. Pilsbry (1890:288) followed
Jeffreys in this usage, and this name has since then been
used for this Arctic species, either as a distinct species or
as a subspecies, variety, or form. Calkin, for instance, in
his otherwise excellent treatment of boreal and arctic
Trochidae (Calkin, 1955:80-83) used Margarites groen-
landica groenlandica for the typical boreal subspecies,
and M. groenlandica umbilicalis for the arctic subspe-
cies.
Although Calkin (1955:80-83) and Abbott (1974:36)
have treated these two taxa as subspecies of one species,
I prefer to consider them as separate species. The shell
characters of each are quite distinctive, and in the few
places where their distributional ranges overlap there is
no evidence of hybridization or blurring of shell char-
acter differences. Future biochemical studies may sug-
gest a closer relationship.
The species called Margarites groenlandicus umbili-
calis (Broderip & Sowerby, 1829) by most recent authors
must bear the name Margarites groenlandicus (Gmelin,
1791), and the species called M. groenlandicus groen-
landicus (Cmelin, 1791), should bear the name Mar-
garites striatus (Leach, 1819). Leach's diagnoses is very
brief, but Gray (1826:567) stated that he compared
Leach's type with the description and figure of Turbo
carneus Lowe, 1825, and verified their identity. Lowe's
species was based on specimens from northern Scotland.
The localities of almost 200 lots of both M. groenlan-
dicus and striatus were plotted on a map of the North
Atlantic (figure 13). The distributional pattern indicates
that each species has a fairly discrete range. Margarites
groenlandicus occurs only in the colder Arctic waters,
from southwest of Victoria Island, Northwest Territories,
Canada, eastward to northern Spitsbergen and Franz
Joseph Land, and south to the northern tip of Labrador
and the southern tip of Greenland. Margarites striatus
is found from southern Massachusetts to northern Lab-
rador along the west coast of Greenland only to slightly
north of Disko and in Iceland, the Faeroes and Shetland
Islands, northern Scotland, and the Norwegian coast from
near Bergen to the Kola Peninsula, and in western and
southern Spitsbergen.
It appears that in only two places do the ranges of the
two species overlap: at the northern end of Labrador and
in the central part of the west coast of Greenland. This
distribution map (figure 13) agrees well with the map
given by Calkin (1955:80-81), e.xcept that he included
additional records from the Arctic north of Norway and
the U.S.S.R., extending the range of M. striatus to the
east coast of Novaya Zemlya Islands, and, to the north,
the range of M. groenlandicus reaches to the Laptev Sea,
just beyond the Taymyr Peninsula.
The presence of M. striatus on the shores of Iceland,
Norway, Spitsbergen and Novaya Zemlya, so much far-
ther north than in the western Atlantic, can be explained
by the effects of the Gulf Stream and North Atlantic
Current, and its extension, the Norwegian Coast Current.
On the west coast of Greenland the warm West Green-
Page 122
THE NAUTILUS, Vol. 103, No. 4
land Current, and extension of the westward-turning
gyre of the North .Atlantic Current permitted the estab-
lishment of M. striatum as far north as Disko. The absence
of M. striatus on the east coast of Greenland is due to
the influence of the cold East Greenland Current coming
from the Arctic.
ACKNOWLEDGMENTS
I am grateful to Dr. Jorgen Knudsen of the Zoologisk
Museum, Copenhagen, Denmark, for making it possible
for me to examine what is undoubtedly type material of
Trochus groenlandicus Gmelin. My sincere appreciation
to the late Dr. Joseph Rosewater and Dr. R. Tucker
Abbott for reviewing this manuscript and making helpful
comments. Thanks are due to Mrs. Caroline Cast, De-
partment of Invertebrate Zoology, National Museum of
Natural History, for assistance in preparing the distri-
butional map, and to Victor E. Krantz of the Smithsonian
Photographic Service for preparing photographs.
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principally to the genus Chiton, observed on the coast of
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THE NAUTILUS 103(4):124-130, 1990
Page 124
Two New Species of Chicoreus Subgenus Siratus
(Gastropoda: Muricidae) From Northeastern Brazil
Emily II. Yokes
Geology Department
Tulane University
New Orleans, LA 70118-5698, USA
ABSTRACT
Two new species of muricid gastropods are described herein
from the vicinity of Salvador, Bahia, Brazil. Both forms have
been known for some time but have been misidentihed in the
hterature. Chicoreus (Siratus) carnUjnac has been identified as
"Murex chrysostoma Sowerby," but differs in having a deflect-
ed siphonal canal. Chicoreus (Siratus) coltrorum, has been
identified as "Murex consuela (Verrill)," but differs in having
a paucispiral protoconch.
Key words: Muricidae; Gastropoda; Chicoreus. Siratus; Bra-
zil
INTRODUCTION
Most authors (e.g., Abbott, 1974; Rios, 1970, 1975, 1980)
have identified moliusks from eastern Brazil by the names
of well known Caribbean species. In some cases, these
identifications are proving to be incorrect. In this paper
two previously misidentihed species of muricid gastro-
pods are described.
One of these, described herein as Chicoreus (Siratus)
carolynae. is of particular interest as it demonstrates the
close relationship, not well understood heretofore, be-
tween members of Haustellum Schumacher, 1817, and
Siratus Jousseaume, 1880. When I first began studying
the Muricidae of the western Atlantic (Yokes, 1963) I
assigned all species with three spinose varices and a long
siphonal canal to Murex sensu stricto. Within this group,
I separated two subgroups: the "Indo-Pacific" form, with
a straight siphonal canal; and the "Western Atlantic"
form, in which the canal was deflected dorsally (see figure
lb). Further work on the family (Yokes, 1965) convinced
me that the latter group should be taken out of Murex
and transferred to the subgenus Siratus. which 1 felt was
better placed within the genus Chicoreus, leaving only
those species with a long straight canal in the genus
Murex.
More recent work on the genus Murex in the Indo-
west Pacific (Ponder and Yokes, 1988) disclosed that
there are certain morphological differences between Mu-
rex and Haustellum, including the presence of a labral
tooth in species of Murex but not in Haustellum, the
presence of rugae on the parietal lip in species of Haus-
tellum but not in Murex, as well as differences in early
teieoconch ornamentation, color pattern, and other fea-
tures. This convinced me that not only the three spinose
varices present in both Murex and Siratus but also the
long straight canal in Murex and Haustellum are a result
of convergence. The ultimate conclusion brought about
by work on the Indo-Pacific species of Murex and Haus-
tellum was the somewhat disturbing realization that there
are no species of Murex in the Western Hemisphere (with
one possible exception — Murex surinamensis Okutani,
1982) and that all the western Atlantic "Indo-Pacific"
forms of Murex (sensu Yokes, 1963) are actually to be
placed in the genus Haustellum.
This shift in nomenclature was corroborated by work
done on the fossil muricids of the Dominican Republic
(Yokes, 1989), where one sees two species that are so
similar as to be very difficult to distinguish — except that
one (Murex nwssorius Sowerby, 1841) has a straight si-
phonal canal and is assigned to Haustellum and the other
(Murex domingensis Sowerby, 1850) has a deflected ca-
nal and is assigned to Siratus. I believe that the similar-
ities seen in these two species transcend convergence and
reflect close phylogenetic affinity.
On the basis of the geologic history of the two groups,
it seems probable that the New World species, which are
taxonomically separated into tw o distinct genera — Haus-
tellum and Chicoreus. are much more closeK related to
each other than either is to true Murex. Nowhere is this
more obvious than in the species described below, which
has heretofore been identified as Haustellum chrysosto-
ma, or a variety thereof, but which is here placed in the
subgenus Siratus. Perhaps the strongest similarity be-
tween H. chrysostoma and C. (S.) carolynae, n. sp., is
the color pattern, in which the Siratus species repeats
the typical brown and white striped pattern of many
species of Haustellum.
This color pattern is just one of the man\ characters
that is used to separate Haustellum from Murex. The
true species of Murex are usualh monochromatic cream
or tan in color, have a distinct labral tooth and a smooth
parietal lip. Species of Haustellum, and of Siratus usually
have brown and white spiral bands, no labral tooth, and
E. H. Yokes, 1990
Page 125
Figures 1-6. Chicoreus (Siratus) carolynae, n. sp. ( x I'/j). 1. USNM 860504 (holotype); height 52.8 mm, diameter 24.4 mm, Ilha
de Itaparica, Bahia. 2. USNM 860505 (paratype 1); height 56.0 mm, diameter 24 5 mm, Ilha de Itaparica, Bahia. 3. 4. USNM
860505 (paratype 2): height 53 8 mm. diameter 27.0 mm (figure 4, x 10), Ilha de Itaparica, Bahia. 5. MORG 20.748 (paratype
3); height 49 6 mm. diameter 25.3 mm, Itapua. Bahia. 6. MORG 22.129 (paratype 4); height 47 6 mm, diameter 23.2 mm. Yacht
Club, Salvador, Bahia; 10 m. All specimens collected dead, all except that in figure 6 at low tide. Specimens in figures 1-4 whitened
to show details of ornamentation.
a rugose parietal lip. In addition, in those species of
Haustellum with relatively well-developed varical spines
{e.g., Haustellum kiiensis [Kira, 1959]) there is a ten-
dency to develop webbing between the spines. This is
carried to its fullest extreme in Siratus. The onK differ-
ence between Haustellum and Siratus is the presence of
a straight vs. a deflected siphonal canal. In other words,
what I was attempting to distinguish in 1963 was not
two groups of Murex but Haustellum and Siratusl
In the second species, described below as C. (S.) col-
Page 126
THE NAUTILUS, Vol. 103, No. 4
trorum, we have a more straightforward case of two
superficially similar forms, with the Brazilian species
having pre\iously been misiderUihed as C. (S. ) consuela.
These two species differ primariK by the nature of the
protoconch. .\ similar case has also been discovered in
the two similar appearing, and often synonymized, species
Chicoreus (Phyllonotus) pomum (Gmelin, 1791) and C.
(P.) onilatus (Reeve, 1845) that ma\ be unequivocally
distinguished by different types of protoconchs (Houart,
1987). Throughout the Muricidae the nature of the pro-
toconch is extremely stable, and is the best means of
distinguishing similar appearing species (Ponder and
Yokes, 1988, p. 3).
SYSTEMATIC DESCRIPTIONS
Class Gastropoda
Family Muricidae Rafinesque, 1815
Subfamily Muricinae Rafinesque, 1815
Genus Chicoreus Montfort, 1810
Chicoreus Montfort, 1810, Coriciiyl. Syst,, v. 2, p. (ill
Type species: Murex ramosus Linn., 1758, by original
desig.
Subgenus Stratus Jousseaume, 1880
Stratus Jousseaume, 1880, Le Naturaiiste, .■\nnee 2, no. 42, p.
335.
Type species: "Purpura sirat" Adanson (= Murex sen-
egalensis Gmelin, 1791), by original desig.
Chtcoreus (Stratus) carolynae, n. sp.
Figures 1-6
Murex chrysostoma Sowerb), var.? Kios, 1970, Coastal Brazil-
ian Seashells, p. 77, pi. 21; Rios, 1975, Brazilian Marine
Mollusks Icon., p. 84, pi. 24, fig. 339.
Murex chrysostoma Sowerby. Rios, 1985, Seashells of Brazil,
p. 81, pi 29, fig. 354 (not of Sowerby).
Description: Shell with seven teleoconch whorls. Pro-
toconch of one and one-half rounded whorls with a small
keel adjacent to the suture, ending at a sharp vari.x. Spiral
ornamentation on early teleoconch whorls of three to five
indistinct cords, best seen where crossed by axial ridges.
Number of axial cords increasing gradually. Body whorl
ornamented by six to eight cords, alternating with weaker
secondary cords. In some cases, weaker tertiary threads
present. Cord at shoulder somewhat stronger than others.
Six to eight relatively weak cords on siphonal canal, some
alternating with tertiary threads. Axial ornamentation on
earliest teleoconch whorls of twelve rounded ridges per
whorl. Beginning at fourth teleoconch whorl every fourth
ridge strengthened into a small rounded varix. Remain-
ing ridges persisting as intervarical nodes. Number of
intervarical nodes between each pair of varices some-
times decreased to only two on latest teleoconch whorls
but most commonly three present on all whorls. Three
varices per whorl beginning on approximately sixth te-
leoconch whorl and persisting to adult stage. Raised ridges
formed at intersection of spiral cords and varices. Small
spines developed on varices only at shoulder and juncture
of bod\ whorl and siphonal canal. Rarely a small flange
along the outer margin of the anterior portion of the
varix. Aperture elongate-oval, parietal lip appressed at
posterior end, free-standing at anterior end, with one
strong anal tooth at posterior end and several rugae over
entire length, but stronger on anterior half. Margin of
outer lip scalloped by spiral cords, with a notch at in-
tersection of each cord and edge of lip. Deeper notches
corresponding to stronger cords. Anterior half of margin
extended more adaperturally than posterior half. At
meeting of spiral cords and axial ridges small elongate
welts raised on top of ridges, giving entire surface of
intervarical area a nodulose appearance. Siphonal canal
long, narrow, almost sealed, open only by a narrow slit.
Siphonal canal deflected dorsally, with terminations of
former canals remaining as a series of spurs surrounding
base of body whorl. Color ranging from white to tan
with brown spiral bands. Brown band at shoulder and
ba.se of body whorl darker. On margin of outer lip a
dark spot of brown staining each notch formed by spiral
cords. Operculum reddish-brown, typically muricine with
a terminal nucleus.
Holotype: USNM 860504; height 52.8 mm. diameter
24.4 mm (figure 1).
Paratype I : USNM 860505; height 56.0 mm, diameter
24.5 mm; type locality (figure 2).
Paratype 2: USNM 860505; height 53.8 mm, diameter
27.0 mm; type locality (figures 3, 4).
Paratype 3: MORG No. 20.748; height 49.9 mm, di-
ameter 25.3 mm; Itapua, Bahia, low tide. Coll. L. C.
Araujo, 1975 (figure 5).
Paratype 4: MORG No. 22.129; height 47.5 mm, di-
ameter 23.2 mm; Yacht Club, Salvador, Bahia, 10 m
depth. Coll. Bernardo Linhares, 1982 (figure 6).
Paratype 5: MORG No. 8016; height 45.3 mm, diam-
eter 22.9 mm; Bahia, 13 m depth. Coll. B. Tursch, 1962
(Rios, 1970, pi. 21, M. chrysostoma var.? — apertural view
only; 1975, pi. 25, fig. 339; 1985, pi. 29, fig. 354).
Other material studied: Five unfigured paratypes:
MORG No. 8016; height 59.8 mm, diameter 28.7 mm;
Bahia, 13 m. MORG No. 20.748; specimen a, height 47.0
mm, diameter 23.4 mm; specimen h. height 40.4 mm,
diameter 22.3 mm; specimen c, height 36.9 mm, di-
ameter 19.8 mm; Itapua, Bahia, at low tide. MORG No.
22.129; height 53.3 mm, diameter 26.5 mm; Yacht Club,
Salvador, Bahia 10 m. Also 14 additional specimens col-
lected by Carolyn Voss, from type locality.
Type locality: Ilha de Itaparica, Bahia, at low tide.
Discussion: This species has been figured by Rios as
Murex chrysostonia Sowerby, 1834, var.? (1970, 1975)
and later as Murex chrysostoma. In the first two editions
he correctly identified true Haustellum chrysostoma
E. H. Yokes, 1990
Page 127
(1970, pi. 20; 1975, pi. 24, fig. 338) but in the third edition
(1985) he changed this identification to Murex tnessorius
Sowerby, 1841 (1985, p. 81, pi. 29, fig. 355) and changed
the former "Murex chrysostoma var.?" to Murex chry-
sostoma (1985, p. 81, pi. 29, fig. 354), indicating the
reason for the change was that M. chrysostutna nia\ be
"distinguished from messorius by the radular teeth."
This statement is based upon a mi.\-up of illustrations in
an otherwise excellent study of the muricid species found
on the coast of Venezuela by Gonzalez and Flores (1972).
In this work they illustrate (figure 9-c) a thaidine radula
(probably Thais haemastoma) as "Murex chrysosto-
mus." It appears very different from the radula of
Murex messorius. There is no difference between the
radulae of M. messorius and M. chrysostoma, both are
typically muricine. Gonzalez and Flores have also illus-
trated as the radula of Murex donmoorei BuUis, 1964
(their figure 9-b), another thaidine radula. It would ap-
pear that two illustrations have been switched, their fig-
ures 9 and 2, for the latter shows as rachidian teeth of
Thais deltoidea, T. rustica. and T. haemastoma flori-
dana, three illustrations of muricine radulae (presumably
Murex cf. messorius, M. donmoorei, and M. chrysosto-
ma, as indicated for figure 9).
Although confounded with Haustcllum chrysostoma
by Rios, the two forms bear only a general resemblance
to each other, in that both have three essentially non-
spinose varices, a long siphonal canal, and brown color
bands. The closest relationship is actually with the mem-
bers of the group of C. (S.) motacilla (Gmelin, 1791), C.
(S. ) cailleti (Petit de la Saussaye, 1856), and C. (S. ) cailleti
kugleri (Clench and Perez Farfante, 1945). These three
closely related forms have been well figured by Clench
and Perez Farfante (1945: pi. 9, figs. 1, 2, C. cailleti form
kugleri; figs. 3, 4, typical C. cailleti; and figs. 5, 6, C.
perelegans Vokes, 1965 [new name for Murex elegans
Sowerby non Donovan], not C. cailleti, as indicated; and
pi. 11, C. motacilla). This new species differs from C.
motacilla in being more slender, with a narrower si-
phonal canal and with usually three weaker intervarical
nodes between each pair of varices, in contrast to the
invariably two, strong nodes seen in C. motacilla. In this
respect, C. carolynae more nearly resembles C. (S.) cail-
leti form kugleri, which in the early whorls usually has
four intervarical nodes between each pair of varices. This
number decreases to only two in the adult stage, as in
typical C. cailleti and C. motacilla. Again, the differ-
ences between C. carolynae and C. kugleri are the more
slender body and less deflected siphonal canal in C. car-
olynae.
This new^ species is known to occur only in the State
of Bahia, Brazil, in shallow water. In addition to the type
material, Rios has reported it from Ponta Jaburu, in 7
fms (10 m), and Porto da Barra, noting that it lives on
sandy bottoms. He suggests that it is "perhaps a shallow
water form" of H. chrysostoma (1970, p. 77). In the
latest edition (1985, p. 81) he states that the species is
dredged from 18 to 90 m on sandy bottoms, but this may
reflect confusion with true H. chrysostoma. All type ma-
terial was collected in water shallower than 10 m, but
most of it was beach material inhabited by hermit-crabs,
and conset]ueiitly the living depth is unknown.
Chicoreus (Siratus) coltrorum, n. sp.
Figures 7-13
Murex {Murex) pulcher Adams Clench, 1959, Johnsonia, v. 3,
no. .39. p. 333.
[?] Murex (Murex) consuelae Verrill. Bullis, 1964, Tulane Stud.
Zoology, V. 11. no. 4. p. 103.
Murex (Murex) pulcher .\(\iims. Rios, 1970, Coastal Brazilian
Seashells, p. 77, pi. 20.
Siratus consuela (Verrill), Rios, 197.5, Brazilian Marine Mol-
liisks Icon., p. 84, pi. 24, fig. 342.
Murex (Murex) consuelae (Vokes) [sic]. Rios, 1985, Seashells
of Brazil, p. 82, pi. 29, fig. 357.
Descriplion: Shell with seven teleoconch whorls. Pro-
toconch of one and one-half bulbous whorls, ending at
a small, sharp varix. Suture deeply impressed. Spiral or-
namentation on earliest teleoconch whorls of three small
cords. A fourth cord and intermediate secondary cords
gradually appearing on shoulder ramp. On body whorl
eight major cords present, plus an additional four or five
on siphonal canal, each pair separated by a secondary
thread. Axial ornamentation on earliest teleoconch whorls
of about 12 small ridges, forming nodes at intersection
with spiral cords. On approximately fourth teleoconch
whorl every fourth ridge enlarging into a small, rounded
varix. Other three remaining as intervarical ridges be-
tween each pair. These persisting up to body whorl, most
adapertural ridge sometimes weaker than other two. On
about fifth teleoconch whorl small open spines developed
on shoulder, at juncture of spiral cord and varices. In
intervarical area, at intersection of spiral cords and axial
ridges, small elongated nodes produced, two nearest to
spiral cord at shoulder often fused into one larger node.
Raised welts at crossing of spiral cords over varices, cor-
responding in size to strength of cord. A small flange
sometimes produced on anterior portion of varices but
not extending onto siphonal canal. The latter, instead,
with usually two small open spines. Aperture elongate-
oval. Parietal lip free-standing at anterior end, appressed
at posterior end with a large anal tooth. Several rugae
along entire length of parietal lip but stronger on its
anterior half. Margin of outer lip crenulated by termi-
nation of spiral cords, inner side of outer lip with a series
of elongate, often paired lirae. Siphonal canal long, al-
most straight, distal end deflected dorsally. Siphonal ca-
nal almost sealed, open only by a narrow slit. Color white
to orange to tan with two darker brown spiral bands, one
at periphery and one at base of body whorl. Operculum
unknown.
Holotype: MORG No. 20.749; height 54.0 mm, diam-
eter 22.4 mm (figure 7).
Paratype 1: MORG No. 20.749A; height 45.2 mm, di-
ameter 21.1 mm; type locality (figure 8).
Paratype 2: MORG No. 15.203; height 28.7 mm, di-
ameter 14.1 mm; off Recife, Pernambuco, 100 m. R/V
Page 128
THE NAUTILUS, Vol. 103, No. 4
\
i:
Figures 7-11. Chicoreus (Siratus) coltroruni. n .sp. 7. MORG 20.749 (holotype); height 54.0 mm, diameter 22.4 mm (x I'A),
Ilha tie Itaparica, Bahia 8. MORC 20.749.\ (paratype 1); height 45 2 mm, diameter 21.1 mm (x IV;). Ilha de Itaparica, Bahia. 9,
10. MOHG 15.203 (paratype 2); height 28.7 mm. diameter 14.1 mm (figures 9a, 9b, x 2; figure 10, x 10), Recife, Pernambuco;
100 m. II. USNM 860506 (paratype 3); lieight 30,6 mm, diameter 14.2 uun (x 2), Ilha de Itaparica, Bahia. Except for specimen
in figures 9, 10, all specimens taken dead at low tide. Figures 12, 13. Chicoreus (Sirutus) consuela (Verrill) \'okes Coll; height
46.3 mm, diameter 20.0 mm (figure 12, x IV2; figure 13, x 10), Soufriere, Dominica, West Indies; 30-40 fms (54-73 m) in hsh-
trap). All specimens except specimen in figure 1 1 whitened to show details of ornamentation.
Almirante Saldanha, 1968; specimen figured by Rios,
1970, pi. 20 (back view) (figures 9, 10).
Paralype 3: USNM 860506; height 30.6 mm, diameter
14.2 mm; type locality (figure 11).
Paralype 4: MORG No. 15.082; height 29 mm, diam-
eter 15 mm; off Recife, Pernambuco; specimen figure
by Rios: 1970, pi. 20 (apertural view); 1975, pi. 24, fig.
342; 1985, pi. 29, fig. 357.
Other material studied: Six unfigured paratypes. MORG
No. 20.749; specimen a, height 53.0 mm, diameter 24.9
mm; specimen b, height 43.0 mm, diameter 20.7 mm;
type locality. MORG No. 1 1.234; .specimen a, height 38.2
mm, diameter 18.0 mm; specimen b, height 34.4 mm,
diameter 15.2 mm; Porto da Barra, Salvador, Bahia.
MORG No. 19.315; specimen a. height 40.9 mm, di-
ameter 18.6 mm; specimen b, height 36.5 mm, diameter
16.4 mm; type locality. Plus eight additional specimens
E. H. Yokes, 1990
Page 129
collected by Jose and Marcos Coltro, from type locality,
and one collected by them from C.uarapari, Espirito San-
to.
Type locality: lliia dc Itaparica, Bahia, at low tide.
Discussion: As is obvious from the synonymy given
above, this species has been considered to be the form
originally described as Murex piilcher A. Adams, 1853.
This taxon is preoccupied b\ Murex pulcher Sowerby,
1813, and DeFrance, 1827, and the next available name
is Murex consuela Verrill, 1950. The Brazilian species is
a similar-appearing form that differs, however, in one
critical feature — the protoconch is markedly different.
Both C. consuela and C. coltrorum, n, sp., are pre-
sumed to be descendants of C. (S.)eumekes Yokes, 1989,
described from the Mio-Pliocene Gurabo Formation of
the Dominican Republic. This fossil species has a pro-
toconch consisting of three and one-half whorls; C. con-
suela has a protoconch of two and one-quarter whorls
(see figure 12) and the Brazilian species has a protoconch
of one and one-half whorls (see figure 10). Inasmuch as
the usual trend in the Muricidae is to decrease the num-
ber of whorls through time, implying a change in re-
productive strategy from planktotrophic to lecitrophic
to direct development, this decrease would suggest that
the Brazilian species is the most recent offshoot from the
ancestral line.
With the exception of the protoconch, the differences
between C. consuela and C. coltrorum are minimal. One
other noticeable difference is the suture, which is more
impressed in C. coltrorum, causing the individual whorls
to appear more rounded and distinct than in C. consuela.
In C. coltrorum the siphonal canal is narrower, less de-
flected dorsally, and does not develop the flange that is
seen in C. consuela. The development of the varices on
the early whorls in C. coltrorum differs in that no varices
appear until the fourth teleoconch whorl and no spines
are produced on these varices until the fifth whorl. In
contrast, in C. consuela the pattern develops one whorl
earlier, i.e., varices on third teleoconch whorl and spines
on fourth whorl.
In the Pleistocene and Recent faunas of the Caribbean
C. consuela is widespread, occurring in the Pleistocene
Moin Formation of Costa Rica, and in the Recent fauna
from off the coast of Texas (Houston Mus. Nat. Sci., from
Flower Garden Banks) to Curafao (de Jong and Coo-
mans, 1988, p. 71), but most commonly in the Lesser
Antilles, usually in depths of 70 to 100 m.
The new species seems to be restricted to the coast of
northeastern Brazil from the islands off Rio Grande do
Norte as far south as Espirito Santo. Rios (1975, p. 84)
has reported it from Atol das Rocas, Fernando de No-
ronha, off Recife, Pernambuco, and off Yitoria, Espirito
Santo, in depths from 35 to 100 m. Except for Rios'
figured specimen (paratype 2) from 100 m, most of the
type lot was collected at low tide, and consists of shells
occupied by hermit-crabs.
Clench (1959, p. 333) reported "Murex (Murex) pul-
cher" from about 30 km off northernmost Bahia, in 40
fms (= 73 m). His specimen (MCZ 164967) unfortunately
lacks the critical protoconch, but otherwi.se is identical
to Rios' figured specimen (paratype 2, herein), and, given
the locality there seems little doubt about the reference
to the new species.
BuUis (1964, p. 103) has also reported "Murex (Murex)
consuelae" from about 300 km east of Ilha de Maraca,
Amapa, just north of the mouth of the Amazon, in 53
fms (= 97 m). This specimen cannot be located in the
U.S. National Museum and given its locality, north of
the Amazon, is only questionably included.
ACKNOWLEDGEMENTS
I am extremely grateful to Mrs. Carolyn Yoss, Ham-
mond, Louisiana, and the brothers Jose and Marcus Col-
tro, Sao Paulo, Brazil, all amateur collectors who pro-
vided most of the material studied here. Dr. E. C. Rios,
doyen of Brazilian malacology, also generously loaned
me relevant material from the collections of the Museu
Oceanografico, Funda9ao Universidade do Rio Grande,
Brazil (MORG). Drs. Ruth Turner and Silvard Kool, of
the Museum of Comparative Zoology (MCZ), Harvard
University, kindly provided the material upon which
Clench's record of C. consuela was based.
ABBREYIATIONS OF REPOSITORY
COLLECTIONS
MCZ — Museum of Comparative Zoology, Harvard
University, Cambridge, Massachusetts, USA.
MORG — Museu Oceanografico, Funda^ao Universidade
do Rio Grande, Rio Grande do Sul, Brazil.
USNM — U.S. National Museum of Natural History,
Washington, D.C., USA.
LITERATURE CITED
Abbott, R. T. 1974. American seashells, 2nd ed. Van Nostrand
Reinhold Co., New York. 663 p., 24 color pis., 6405 text-
figs.
Bullis, H. R, Jr. 1964. Muricidae (Gastropoda) from the north-
east coast of South .America, with descriptions of four new
species. Tulane Studies in Zoology 11(4):99-107, 1 pi., 2
tables.
Clench, W. J. 1959. The genus Murex in the western Atlantic.
Johnsonia 3(39):331-333, pi. 174.
Clench, W. J. and I. Perez Farfante. 1945. The genus Murex
in the western Atlantic. Johnsonia l(17):l-58, pis. 1-29.
Gonzalez, A. R. and C. Flores. 1972. Nota sobre los generos
Thais Roeding, Purpura Brugiere [sic] y Murex Linnaeus
(Neogastropoda: Muricidae) en las aguas costeras de Ven-
ezuela. Boletin del Institute Oceanografico de la Univer-
sidad del Oriente ll(2):67-82, 11 text-figs.
Houart, R. 1987. Rehabilitation de Chicoreiis (Phyllonotus)
oculatus (Reeve, 1845) (Gastropoda; Muricidae). Apex 2(1):
7-10, 1 pi.
Jong, K. D. de and H. E. Coomans. 1988. Marine gastropods
from Curasao, Aruba, and Bonaire. Studies on the Fauna
of Curasao and other Caribbean Islands 69(214):1-261,
pis. 1-47.
Page 130
THE NAUTILUS, Vol. 103, No. 4
Ponder, W. F. and E. H. Yokes. 1988. A revision of the Indo-
Wesl Pacific fossil and Recent s[X"cies of Murex s s. and
Haustellum (Miiricidae: Gastropoda: Mollusca). Records
of the .Vnstraiian Museum. Supplement 8:1-160, 89 text-
figs., .57 tables
Rios, E. C. 1970. Goastal Brazilian .Seashells. Museu Ocean-
ografico, Rio Grande. 2.5.5 p., (iO pis., 4 maps.
Rios, E. C. 1975. Brazilian Mollusks Iconography. Museu
Oceanografico, Rio Grande. S^\ p., 91 pis.
Rios, E. C. 1985. Seashells of Brazil. Funda^ao Universidade
do Rio Grande, Museu Oceanografico, Rio Grande. .329
p., 102 pis.
V'okes, E H 196.3. Cenozoic Muricidae of the western .At-
lantic region. Part I — Murex sensu stricto Tulaiie Studies
in Geology 1(3)93-123. pis. 1-4.
X'okes, F. H. 1965 Onozoic Muricidae of the western .At-
lantic region. Part 11 — Chicoreus sensu stricto and Chi-
coreus (Siratus). Tulane Studies in Geology 3(4):181-204,
pis. 1-3, 2 text-figs.
\Okes, E. H. 1989. Neogene Paleontology in the northern
Dominican Republic. 8. The famiK Muricidae (Mollusca:
Gastropoda I Bulletins of .American Paleontology 97(332):
5-94, pis. 1-12, 21 text-figs., 3 tables.
THE NAUTILUS 103{4):131-135, 1990
Page 131
Distorsio ridens (Reeve, 1844): A Synonym of
Distorsio clathrata (Lamarck, 1816)
(Gastropoda: Personidae)
William K. Emerson
Walter E. Sage, III
DepartiiKMit of Invertebrates
American Mu.seum of Natural History
New York, NY 10024-5192, USA
ABSTRACT
A reevaiuation of the identity of Distorsio ridens (Reeve, 1844),
suggests that this taxon was based on specimens of two New
World species having erroneous locahty data, and we conclude
ttiat this taxon is a junior subjective synon> m of Distorsio clath-
rata (Lamarck, 1816), We correct the erroneous t\pe localit\
of D. ridens and select Key West, Florida as the type localit\
INTRODUCTION
We were recently asked to identify specimens of two
species of Distorsio obtained by commercial fishermen
trawling off Somalia in deep water. As a result of this
study, we found that a reexamination of the status of
Distorsio ridens (Reeve, 1844) was in order before we
could proceed with our review of the Somalian speci-
mens. This paper is a result of our investigation.
It should be noted in passing, that the genus Distorsio
and related genera were recently removed from Ranel-
lidae Gray, 1854 (= Cymatiidae Iredale, 1913) and
awarded familial recognition, Personidae Gray, 1854, as
a sixth family of Tonnoidea by Beu (1988).
HISTORICAL REVIEW OF
DISTORSIO RIDENS
Triton ridens Reeve (1844a, pi. 12, Triton sp. 46) was
briefly described in both Latin and English texts and was
illustrated by a colored lithograph executed by G. B.
Sowerby, 2nd. Only an apertural view was presented of
the figured specimen, which was said to be from the
Philippine Islands and collected by Hugh Cutning. Reeve
compared T. ridens with "Murex cancellinus de Roissy",
(1805:56, 57; see Emerson & Puffer, 1953:97), which is
now recognized as a junior synonym of the Indo-Pacific
Distorsio reticularis (Linne, 1758); see Beu (1987:314).
Some seven months later, the same description, together
with the remarks in a slightly altered form, appeared
without an illustration in the Proceedings of the Zoolog-
ical Society of London (Reeve, 1844b:115). This ta.xon.
therefore, dates from the original description in Con-
chologia Iconica (Reeve, 1844a).
Unfortunately, subsequent workers have confused
Reeve's taxon with other earlier or later described con-
geners from several biogeographical faunal provinces.
The resulting taxonomic vicissitudes have caused a gen-
eral disagreement on the identity of D. ridens. For ex-
ample. Reeve's figured specimen was reproduced by
Tryon (1880:35, pi. 17, fig. 177) as "Distorsio cancellinus
Roissy" and by Wagner and Abbott (1978:12-802, fig.
13-116) as "Distorsio reticulata Roding". The question-
able identity of Distorsio ridens has resulted in this name
being misapplied to D. smithi von Maltzan, 1844, from
West Africa, by Nickles (1950:86, fig. 133), to D. per-
distorta Fulton, 1938, from Japan, by Oyama (1958: pi.
1, figs. 7, 8), to D. reticulata (Roding, 1798) [= D. re-
ticularis Linne, 1758] from the Philippines, by Emerson
and Puffer (1953:103); Puffer (1953:114); Springsteen
(1984:5; 1985:3); Springsteen and Leobrera (1986:117),
and to D. decussata Valenciennes, 1832, from the eastern
Pacific, by Beu (1985:62) and Parth (1989:54).
Lewis (1972:47, 48, figs. 45-48) discussed at length the
status of Distorsio ridens and accorded this taxon full
specific recognition on the basis of the data then available
to him. For D. ridens, he selected and illustrated a lec-
totype, provided photographs of a paralectotype, and
found this taxon to be separable from D. reticularis, D.
perdistorta and D. decussata. He did not compare D.
ridens with D. clathrata (Lamarck) from the western
Atlantic. Wolfe (1976:12) and Springsteen (1981:8) also
considered D. ridens a distinct species.
REEVALUATION OF THE IDENTITY OF
DISTORSIO RIDENS
In our view, two factors have been largely responsible
for the misinterpretation of D. ridens. First, was the lack
of a dorsal view of the specimen figured by Reeve (1844a).
As a result, the nature of the sculpture on the dorsum
could not be determined with certaintv from the illus-
W. K. Emerson and W. E. Sage, III, 1990
Page 133
Figures 14, 15. Distorsio dathrata (Lamarck, 1816), aperture
enlarged to show bifid tooth of the primary columellar pica.
Figure 14. Specimen illustrated in figure 4. Figure 15. Lec-
tot\ pe illustrated in figure 2.
tration or from the brief description. Second, is the gen-
eral acceptance of the validity of the type locality, the
Philippine Islands. This has led most reviewers to limit
comparisons with Indo-Pacific species. We shall expand
on these subjects.
Lewis (1972:48) searched the collection of the British
Museum (Natural History) for potential syntypes of D.
ridens. He could not locate the specimen figured by
Reeve (1844a: pi. 12, sp. 46). Therefore, he selected a
specimen in the collection of the American Museum of
Natural History (AMNH) which he stated ". . . so very
closely matches Reeve's figure that I feel it is reasonable
to designate it as the lectotype" (Lewis, 1972:48, fig. 48).
The specimen (AMNH 6369), labeled "Philippines", was
from the William A. Haines Collection, received by the
Museum in 1879. Haines (1822-80), who was a member
of the AMNH Board of Trustees in the decade before
his death, was a colleague of the New York conchologist,
John C. Jay (1808-91). He could have obtained the spec-
imen from Jav or perhaps from England directly from
Lovell A. Reeve (1814-65) or Hugh Cuming (1791-1865),
who is known to have been a dealer in shells (c/. Lewis,
1972:48). The source of Haines' specimen, however, is
not known, for the AMNH catalog entry lacks such data.
We agree with Lewis' conclusion (1972:48) that the
specimen designated by him as the lectotype of D. ridens
closely resembles the morphological features of the spec-
imen figured by Reeve (1844a); see figures 1-3. Lewis
noted some differences in the artist's depiction of the
apertural morphology. He attributed to artistic exagger-
ations the distorted representation of the second row of
plicae on the lower left parietal shield and the extrava-
gant manner shown for the character of the groove on
the lower shield at the point of entry into the aperture.
These apparent morphological discrepancies may reflect
a composite drawing based on more than one specimen,
most likely representing different species in the type lot.
Reeve's drawing, however, illustrates well the major col-
umellar plica as a bifid tooth, a character not uncom-
monly found in specimens of D. dathrata (cf. figures 1,
14, 15). The wide groove shown entering the aperture
below the parietal shield is, however, more reminiscent
of D. decussata (cf. figures 1-3 with figures 10, 12). It
is our conclusion that the specimen of D. ridens illus-
trated by Reeve (1844a; figure 1, herein), together with
the specimen designated the lectotype (AMNH 6369) by
Lewis (figures 2, 3, herein), and the paralectotype figured
by Lewis (1972: figs. 45, 46), are all referable to D.
dathrata (Lamarck, 1816: pi. 413, figs. 4a, 4b, Liste p.
4).
At this point, it should be mentioned that a more
inflated, lower spired, and shorter canalled morph of D.
dathrata was named D. robinsoni by Petuch (1987:64,
65, pi. 11, figs. 3, 4), on the basis of a few specimens
from Honduras and Brazil. An examination of a large
series of specimens in the AMNH collection from North
Carolina to the Gulf of Mexico and in the Caribbean
region, including Honduras, indicates that D. robinsoni
is an infrasubspecific form that appears in the samples
of the typical form of D. dathrata {cf. figures 4, 5 with
figures 8, 9). Also, see comments on D. robinsoni Petuch
by Parth (1989:52, fig. 1, right) and Manoja (1989:29).
We subsequently were able to examine personally the
three paralectotypes of D. ridens remaining in the British
Museum (Natural History; Registry number 1967630,
Acquisition number 1829, "H. Cuming Collection, Phil-
ippines "). Two of these specimens are referable to D.
dathrata; the smaller one measures 70.8 mm in height
(figures 6, 7), and the larger specimen (Lewis, 1972: figs.
45, 46) measures 78.6 mm in height (apical whorls are
lost). The third paralectotype is referable to D. decussata
(figures 10, 1 1 ). It measures 62.2 mm in height, essentially
the same as the height of Reeve's (1844a) figured spec-
imen, which measures 63.1 mm in height. The specimen
designated the lectotype by Lewis (1972:48; fig. 48) is
also close to the height of Reeve's figured specimen,
Figures 1-9. Distorsio dathrata (Lamarck, 1816). Figure 1. Copy of original illustration of Triton ridens Reeve (1844a, Triton
sp. 46). Figures 2, 3. Lectotype of Distorsio ridens (AMNH 6369), W. A. Haines Collection. Figures 4, 5. A specimen with
orange-pigmented shield and outer lip as in the lectotype, trawled off Punta Patuca, Atlantic Honduras, in 18 to 27 meters (AMNH
238556), e.\ E. Garcia Collection. Figures 6, 7. Paralectotype of D. ridens [BM(NH) 1967630]. Figures 8, 9. Specimen of morph
robinsoni Petuch, 1987, from same lot as the specimen illustrated by figures 4, 5. Figures 10- 13. Distorsio decussata (Valenciennes,
1832). FigureslO.il. Paralectotype of D. ridens [BM(NH) 1967630]. Figures 12, 13. Paralectotype of D. ridens (MCZH
186600). All xl.
Page 134
THE NAUTILUS, Vol. 103, No. 4
measuring 64.3 mm (tip of the siphonal canal is broken)
vs. 63.1 mm. We suspect that Reeve's figured specimen
may be a composite drawing based on specimens of D.
clathrata and D. decussata in the type lot. However, we
cannot be certain that the three paralectotypes in the
British Museum (Natural HistorN ) actualK- were the spec-
imens available to Reeve at the time he described D.
ridens. One of the label scraps accompanying the type
specimens reads: "St. Johns [?= Saint Johns, Antigua,
17°06"N, 61°5r'W], Mr. Hartvig". Thus, at least one of
the three specimens forming the paralectotype lot in the
British Museum (Natural History) may have been added
after the ta.xon was described. Furthermore, there is a
paralectotype in the Museum of C;omparative Zoology,
Harvard Universit\ (186600) labeled "Triton ridens
Reeve, Philippine Islands, H. Cuming, C. B. .\damsColl.,
exch. Amherst College, 1942, Ace. 1173 ". This specimen,
which measures 37.1 mm in height, is referable to D.
decussata (see figures 12, 13), as Lewis (1972:48) has
pointed out.
It should be noted that the ta.xa, D. decussata (Valenci-
ennes, 1832) and D. clathrata (Lamarck, 1816), were not
recognized by Reeve (1844a,b). He does not mention
Valenciennes" ta.xon and he refers "Triton clathratus,
Lamarck" to the synonymy of "Murex cancellinus De
Roissy", which is a junior synonym of Distorsio reticu-
laris (Linne, 1758; see Beu, 1987:314). Reeve apparently
confused Lamarck's D. clathrata with D. reticularis from
the Indo-Pacific and did not recognize the presence of
D. clathrata in the Western Atlantic.
Therefore, it is clear that the type locality of D. ridens,
"Philippine Islands, Cuming ", is in error. Although Hugh
Cuming is known to have collected extensively in the
Philippine Islands (1836-40) and on the west coast of
South America (1828-30), he apparently did not collect
in the Caribbean region (Dance, 1986:111-131). He most
certainly, early in his career, however, had access to
common marine shells of the western Atlantic by ex-
changes. (He is pictured with a Caribbean shell in a
photograph taken ca. 1861, Dance, 1986: pi. 25). As oth-
ers have pointed out (Clench, 1945; Dance, 1986), much
of the material in the Cuming collections was accom-
panied by mislocalized data. This would seem to be the
case for D. ridens, as all of the type specimens represent
New World species. We here correct the type locality of
D. ridens and select Key West, Florida as the type lo-
cality.
In summary, we must address the question: should D.
ridens be placed in the synonymy of D. clathrata from
the western Atlantic or D. decussata from the eastern
Pacific? As we have noted. Reeve's (1844a) figured spec-
imen more closely resembles the apertural characters of
D. clathrata than those of D. decussata, although the
drawing may represent a composite based on specimens
of both species. Furthermore the description (Reeve,
1844a,b) states that the ". . . transverse ridges [are] du-
plicate . . .", suggesting the presence of a double row of
spiral cords at the periphery of the whorls, a morpho-
logical feature common to D. decussata. On the other
hand. Reeve described the cancellated sculpture of D.
ridens as being wider and more prominent than that
found in D. reticularis. These are sculptural features
characteristic of D. clathrata. This statement serves to
contradict his description of duplicate transverse ridges.
.Again, these descriptive conflicts reinforce our conclusion
that two different species formed the basis for the de-
scription of D. ridens. Moreover, Reeve remarked, ". . .
the orange-stained colouring of the enamelled disc is
particularly characteristic " of D. ridens (see figure 4).
This pigmentation is not infrequentK found in D. clath-
rata, but is not known in D. decussata. Furthermore the
dorsal sculpture of the lectotype of D. ridens is consistent
with that of D. clathrata and not "duplicate " as in D.
decussata.
Inasmuch as none of the existing types of D. ridens
can be attributed without doubt as being the specimens
used by Reeve to describe this taxon, we believe Reeve's
figured specimen (figure 1, herein) should be the primary
source for the identitx of this species. .Additionally, we
consider the lectotype virtually identical to Reeve's fig-
ured specimen. Therefore, we must conclude that Dis-
torsio ridens (Reeve, 1844) is best placed in the synon-
\my of D. clathrata (Lamarck, 1816).
ACKNOWLEDGMENTS
We are indebted to a number of people for providing
us with specimens and/or data needed for this review.
Types and other specimens were kindly lent by Kathie
M. Way of the British Museum (Natural Histor\). by
Gary Rosenberg and M. Andria Garback of the Phila-
delphia Academy of Natural Sciences, and by Silvard
Kool of the Museum of Comparative Zoology, Harvard
University. Emilio Garcia of Lafayette, Louisiana gen-
erously donated critical specimens. Our AMNH col-
leagues, Andrew S. Modell and Stephanie Crooms, re-
spectively, contributed the photography and the word-
processing of the manuscript. We are most grateful to
Alan Beu of the New Zealand Geological Survey and
Hal Lewis of the Philadelphia .\cadem\- of Natural Sci-
ences for critically reviewing the manuscript. Dr. Beu
entirely agrees with our conclusions, whereas Mr. Lewis
still holds to his view that D. ridens is a probable Indo-
West Pacific faunal constituent that is a unique species
of which additional specimens have yet to be discovered.
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THE NAUTILUS 103(4): 136-139, 1990
Page 136
Freshwater Mussel Fauna (Bivalvia: Unionidae) of the
New River Gorge National River, West Virginia
kurt J. Jirka
Iclitluological Associates, Inc.
301 Forest Drive
Ithaca, New York 14853, USA
Richard J. Neves
Virginia Cooperative Fish and
Wildlife Research I'nit'
U.S. Fi.sh and Wildlife Service
Virginia PoKtechnic Institute
and State University
Blacksburg, Virginia 24061, USA
ABSTRACT
An extensive freshwater mussel survey of the New River Gorge
National River, a national park in West Virginia, during the
summers of 1984 and 1983 yielded seven species of living
mussels and empt\ valves of an eighth species. Actinonaias
ligamenlina dominated the mussel fauna in all beds, consti-
tuting 94''c of the mussels counted. Mussels were abundant
throughout most of the stud) area; the estimated total exceeded
1.5 million individuals. Most mussel beds occurred in the up-
stream third of the park. Lack of suitable habitat appears to
preclude the establisfiment of mussel beds in the extreme lower
portions of this river reach. The mussel fauna within the park
differed significantly in species composition and richness from
that reported in upstream reaches of the New River and the
upper Kanawha River, about 19 km downstream from the park
boundary. Of particular note was the dominance of A. liga-
mentina in the park and its scarcity or absence elsewhere in
the drainage.
INTRODUCTION
The freshwater mussel fauna (Unionidae) of the New
River has been sampled periodically throughout this cen-
tury, beginning with a brief survey by Ortmann (1913).
Since then, several other mussel surveys have been con-
ducted in various areas in the New River drainage (Jirka
& Neves, 1985), but none have focused on the stretch of
river designated as the New River Gorge National River
(NRGNR) in West Virginia (figure 1). This national park
includes the 84-km section of river from Hinton, West
Virginia, at New River Mile (RM) 63 to a point just
downstream from Fayetteviile, West Virginia (RM 11).
The lack of faunal surveys in this river reach is mainly
' The Virginia Unit is jointly supported by the L nited States
Fish and Wildlife Service, the X'irginia Department of Game
and Inland Fisheries, and Virginia Polytechnic Institute and
State University.
due to its extensive white-water rapids and limited ac-
cessibility. Surveys that have included sites in the park
were cursory, and most of the New River Gorge has
remained virtually unexplored biologically.
We present the results of an extensive mussel survey
conducted from June 1984 to September 1985 through-
out the stretch of New River within NRGNR. Objectives
of the survey were to identify locations supporting con-
centrations of mussels and to determine species com-
position and mussel abundance within NRGNR. Of par-
ticular interest to personnel at the park was the possible
presence of the federally endangered pink mucket,
Lampsilis abrupta (Say, 1831), since live specimens of
this species were collected below Kanawha Falls, 2 km
downstream of the mouth of the New River and about
19 km downstream from NRGNR (Stansberv, 1980;
Clarke, 1982; Taylor, 1983).
METHODS
In June 1984, we conducted a preliminar) reconnais-
sance of the river by inflatable raft to locate important
mussel beds in NRGNR. River reaches having high con-
centrations of mussels or potentially suitable mussel hab-
itat were identified for later qualitative sampling. Qual-
itative sampling consisted of searching a total of 44 sites
for mussels by snorkeling, handpicking, using water-
scopes, and sorting middens of muskrats (Ondatra zi-
bethicus) to determine species present. Most areas were
examined for 0.5-1.0 hour by three or four individuals
using snorkels; mussels were identified, counted, and re-
turned to the substratum.
On the basis of the qualitative sampling, we chose
eleven mussel beds for quantitative sampling to estimate
mussel abundance. Criteria used in selecting these sites
were species diversity, overall mussel abundance,
uniqueness of habitat type, and location within NRGNR.
We attempted to locate study sites throughout the length
of the park to record changes in species composition and
abundance in different reaches of the river (figure 1).
K. J. Jirka and R. J. Neves, 1990
Page 137
Park Bound*
F*YETTE VILLE
WEST
VIRGINIA
Study Area
Lick Creek
auret Creek
Figure 1. Location of major mussel beds (1-11) in the New
River Gorge National River. West Virginia.
We sampled mussels quantitatively by snorkeling along
transects at each site. Transects were established using a
stratified random sampling design to ensure that the dif-
ferent habitats within a bed were sampled in proportion
to their areas. Some areas of certain beds could not be
effectively sampled because depths were unsuitable
(<0.25 or >2.0 m) or time constraints prevented sam-
pling of exceptionally large beds. However, areas chosen
for sampling were representative of most of the bed.
Sampling consisted initially of laying a plastic tape of
known length (usually 50 m) along the river bottom
parallel to the current. Densities of mussels (no./m-) along
the transect were determined by identifying and count-
ing all mussels within 0.5 m of each side of the tape and
dividing the total by the length of the tape. Two snor-
kelers were used on all transects, each counting the mus-
sels on one side of the tape. The presence of empty valves
of each species was also noted. The length and number
of transects on each bed varied with the dimensions of
the bed; at least six transects, each 50 m long, were used
on most beds. Densities from transects in the same bed
were averaged to estimate mean mussel density for the
bed. This mean density was multiplied by the area of
the bed to obtain a minimal estimate of adult mussel
abundance at each site. Nomenclature of mussels is ac-
cording to Turgeon et al. (1988).
RESULTS
Faunal Description
Dense mussel assemblages were found throughout most
of the park (figure 1). In addition to these major mussel
beds, numerous smaller beds and scattered individuals
were found in areas between these larger concentrations.
Live mussels were found in all reaches of the river from
Hinton (RM 63) downstream to Sewell (RM 19). Most
mussel beds were in the upstream third of the river reach
in NRGNR, and their abundance decreased markedly
downstream from Glade Creek (RM 45). No live mussels
or valves were found downstream from a large pool at
Sewell.
Seven species of mussels were collected alive during
the survey (table 1), together with two empty valves of
an eighth species, Lasmigona subviridis (Conrad, 1835).
Actmonaias ligamentina (Lamarck, 1819) was by far
the dominant species, composing about 94% of the total
mussels collected and present at all locations having mus-
sels. Cyclonaias tuberculata (Rafinesque, 1820) account-
ed for 4% and Elliptio dilatata (Rafinesque, 1820) for
1% of the fauna; these species were in all of the major
beds in the park. Tritogonia verrucosa (Rafinesque, 1820)
was relatively common upstream from Sandstone Falls
(RM 55), composing over 2% of the mussels found in this
reach. Since no live specimens of this species were col-
lected below RM 52.5, it made up less than 1% of the
total mussels in the park. Lampsilis ovata (Sa\, 1817),
L. fasciola (Rafinesque, 1820), and Alasmidonta mar-
ginata (Say, 1817) were collected infrequently and are
considered uncommon throughout the river below Blue-
stone Dam. No specimens or valves of Lampsilis abrupta
were collected in NRGNR.
Mussel Densities
Naiad densities in the beds ranged from 2.5 to 13.7/m^
(table 2). The density was highest in bed 9 and the lowest
in bed 11. No discernible pattern of overall mussel den-
sity was detected within the park, though some species
showed evidence of shifting abundance in certain reaches
of the river. Densities of Actinonaias ligamentina were
relatively high in all beds, and freshly dead animals or
relic shells (shells with a dull, chalky nacre from speci-
mens presumed dead for at least a year) of this species
were present at every site. The abundance of Cyclonaias
tuberculata was generally low in beds upstream from
Sandstone Falls and was highest in beds 6 to 8. Densities
of Elliptio dilatata were highest in the river upstream
from Sandstone Falls and showed a general trend of
decreasing numbers downstream. Relic valves of this
species were seen in all beds. Tritogonia verrucosa was
fairlv common in most beds above Sandstone Falls but
Page 138
THE NAUTILUS, Vol. 103, No. 4
Table 1 . Species composition and relative abundance of freshwater mussels in the New River Gorge National River. West Virginia.
Species
Common name
Hflative
abundance (%)
Actintmaias hgamentina (Lamarck, 1819)
Cyclunaias tuln'rculata (Rafinesque, 1820)
Elliptiu dilalala (Rafinesque. 1820)
Triiogonia verrucosa (Rafinesque, 1820)
Lampsilis fasciola (Rafinesque, 1820)
I.anipsilis ovata (Sa\. 1817)
Ahistnidanta rnarginata (Sa\, 1818)
Mucket
Purple wartyback
Spike
Pistolgrip
VVav\-rayed lampmussel
Pocketbook
Elktoe
94
4
1
<1
<1
<1
• I
was collected alive oriK in bed 6. among the beds below
the falls. However, a tew relic shells of this species oc-
curred in every bed. Lampsilis fasciola, L. ovata, and
Alasmidonta marginata were patchily distributed and
were uncommon throughout NRGNR. X'alves of Lamp-
silis fasciola and L. ovata were often found in beds where
no live individuals of these species were collected.
Mussel Abundance
Estimates of total numbers of mussels in the major beds
sampled ranged from 5,475 in bed 2 to 392,175 in bed
8 (table 2). The estimated total number of mussels in all
11 beds combined was about 1.2 million, of which 94%
were Actinonaias Hgamentina. These estimates do not
include juveniles, the smaller mussel beds, or numerous
pockets of mussels that were not counted during the
quantitative portion of the survey. Records from the
qualitative survey indicated the existence of at least 16
minor mussel beds and two additional major beds within
the park. A conservative estimate of 3,000 mu.ssels per
minor bed and 150,000 mussels per major bed would put
the estimated minimal immber of adult mussels within
beds of NRGNR at more than 1.5 million animals. Mus-
sels were most abundant between Sandstone Falls (RM
55) and Piney Creek (RM 38).
(Say, 1829); Toxolasma parvus (Barnes, 1823); Villosa
iris (Lea, 1829); and Quadrula quadrula (Rafinesque,
1820). In 1984, an empty valve of Anodonta grandis
was found immediately below Bluestone Dam b\ one of
us (R.J.N.), and another valve was collected near the
lower end of Brooks Island, in NRGNR by W. A. Tolin
(USFWS, pers. comm). Toxolasma parvus has been found
only near the mouth of the New River at Gauley Bridge.
West Virginia (D. H. Stansbery, Ohio State Museum,
unpublished records). The record of Quadrula quadrula
in the river near Sandstone Falls (Bates, 1979) is ques-
tionable since this species is unreported for the New River
drainage. Villosa iris has not been found in the main
stem of the New River, but one specimen was recently
reported from the Bluestone River, a major tributary of
the New River, by Tolin (1985).
Most of the mussel species collected appeared to have
health) populations in at least a portion of the park, and
those in the ri\er that are considered rare were generally
widespread. Possibly Lasmigona subviridis. of which only
two empty valves were collected (at RM 57 and RM 19),
also has a small population within NRGNR, and the same
ma\ be true of Anodonta grandis. The distributions of
two species, Tritogonia verrucosa and Lampsilis ovata,
were unexpected, particularly in the numbers and lo-
DISCUSSION
The NRGNR supports an abundant mussel fauna that is
relativeK low in diversity. Large beds of naiades are
found from its upstream boundary, downstream to Sew-
ell in the lower gorge. Mu.s.sel colonization of the river
downstream from Sewell appears to be precluded by a
scarcity of suitable habitat and possibly by a lack of
suitable fish hosts for certain mussel species.
The mussel fauna of the park varies considerably from
the fauna found elsewhere in the New River — most no-
tably, in the dominance of Actinonaias Hgamentina
within NRGNR. The dominant species in the river im-
mediately above Bluestone Lake, only 35 km upstream
from NRGNR, are Cyclonaias tuberculata and Lamp-
silis ovata (Tolin, 1985). Further upstream in Virginia,
Tritogonia verrucosa and Elliptio dilatata are also com-
mon (Dillon, 1977). Species reported from the New River
drainage by Jirka and Neves (1985) that were not re-
corded in the present survey were Anodonta grandis
Table 2. Mussel density and abundance in eleven major mussel
beds within the New River Gorge National River, West Vir-
ginia.
Bed'
Mussel
Area
Transect
densitv
Population
No.
(m-)
area (m-)
(no./m-)
estimate
1
4,200
335
3.6
15.120
2
1,165
217
4.7
5,475
3
9,350
300
5.4
50.490
4
18.750
350
3.7
69,375
5
14,700
350
6.8
99,960
6
39,600
300
7.6
300,960
7
1,600
241
3.7
5.920
8
37,350
300
10.5
392.175
9
14.850
300
13.7
203.445
10
9.100
300
7.0
63.700
11
5,500
300
2.5
13,750
Total
1,220.370
' See Figure
1 for local
;i(iii--
K. J. Jirka and R. J. Neves, 1990
Page 139
cations of empty valves collected. Although a healthy
population of Tritogonia verrucosa lives above Sand-
stone Falls, no live specimens of this species were seen
below bed 6, about 4 km below the falls. However, nu-
merous relic shells were found in all major beds down-
stream from Brooks Island, suggesting that T. verrucosa
may have once been relatively common throughout the
river below Sandstone Falls but have since declined in
abundance. Similarly, although live Larnpsilis ovata were
collected from several sites throughout the park, many
areas contained numerous relic valves but no living spec-
imens, indicating that this species may have been more
common in the river in the recent past than during our
surve\.
The relatively low diversity of the mussel fauna in
NRGNR, and in the New River as a whole, is notable
when compared with the 34 species of naiades reported
in the Kanawha River (Stansbery, 1980; Clarke, 1982;
Taylor, 1983), which is joined by the New River about
19 km downstream from NRGNR. Many factors have
been suggested as potential causes of the scarcity of mus-
sel species in the New, but none have been substantiated.
Most of these factors relate to the river's geologic and
climatic past, the presence of physical barriers to dis-
persal, lack of a diverse fish fauna in the river, or stream
captures h\ other drainages (Ross & Perkins, 1959; Neves,
1983; Jirka & Neves, 1985). Physical barriers to upstream
dispersal of fish and mussels, particularly Kanawha Falls,
have probably had the greatest influence on mussel di-
versitv in the New River since 34 species occur imme-
diately below the falls (Taylor, 1983).
The overall abundance of naiades in NRGNR is among
the highest in West Virginia. The NRGNR supports a
mussel assemblage of at least 1.5 million individuals —
an abundance at least comparable to that in the upper
Kanawha River and considerably greater than that in
any reach of the upper New River in Virginia. The
abundance of mussels in NRGNR is strong evidence that
it is not a lack of suitable habitat or poor environmental
quality that has limited the number of species in this
river, but rather, impediments to mussel colonization of
suitable habitats throughout much of NRGNR.
The most striking aspect of the mussel fauna of the
NRGNR is the e-xtreme dominance of Actinonaias li-
gamentina which constituted 94% of the mussel fauna
and exceeded 1,4 million individuals. In contrast, Taylor
(1983) reported that this species represented an average
of only 5% of the mussels at 14 stations sampled in the
upper Kanawha River. Also notable is the scarcity of A.
ligamentina in the New River immediately above Blue-
stone Dam, only 3 km upstream from NRGNR, and the
absence of this species in the New River above Round-
bottom Creek, about 40 km upstream from NRGNR.
Why A. ligamentina has flourished within NRGNR while
being only a minor or missing component of the mussel
fauna elsewhere in the New Kanawha drainage is not
known. An explanation for this distribution pattern would
probably add insight into the factors influencing faunal
diversity and composition within NRGNR and the entire
New River.
ACKNOWLEDGEMENTS
We thank Bret Preston, Del Lobb, Kurt Buhlmann and
Richard Fades for assisting with this survey, and Dr.
David Stansbery for permission to cite collection records
of the Ohio State University Museum of Zoology. The
Virginia Cooperative Fish and Wildlife Research Unit,
the National Park Service, and the Department of Fish-
eries and Wildlife Sciences at Virginia Polytechnic In-
stitute and State University provided financial support
for this project.
LITERATURE CITED
Bates, J. M. 1979. Mussel investigations, state of West Vir-
ginia. West Virginia Project .3-97-R, 91 p.
Clarke, A. H. 19.S2. Survey of the freshwater mussels of the
upper Kanawha River (RM 91-9.5), Fayette County, West
X'irginia, with special reference to Epiohlasma torulosa
torulosa (Rafinesque) and Larnpsilis abrupia (Say) ( =
Larnpsilis orhiculata (Hildreth), of authors) Final Report
to U.S. Fish and Wildlife Service, 104 p.
Dillon, R. T., Jr. 1977. Factors in the distributional ecology
of upper New River mollusks (Virginia/North Carolina).
Unpublished Honors thesis. Biology Department, N'irginia
Polvtechnic Institute and State University, Blacksburg,
59 p.
Jirka, K. J. and R. J. Neves. 198.5. A review of the mussel
fauna of the New River. Proceedings of the New River
Svmposium 4:27 -.36.
Markham, S. L., C. H. Hocutt, and J. R. Stauffer, Jr 1980.
The crayfish (Decapoda: Astacidae and Cambaridae) and
the freshwater mussels (Mollusca: Pelecypoda) of the lower
New River, Virginia and West Virginia. Natural History
Miscellanea No. 208, 1 1 p.
Neves, R. J. 1983. Distributional history of the fish and mussel
fauna in the Kanawha River drainage. Proceedings of the
New River Symposium 2:47-67.
Ortmann, A. E. 1913. The Alleghenian Divide and its influ-
ence upon the freshwater fauna. Proceedings of the Amer-
ican Philosophical Society 52:287-390.
Ross, R. D. and B. D. Perkins. 1959. Drainage evolution and
distribution problems of the fishes of the New (upper Kana-
wha) River system in X'irginia. Part 3. Records of fishes of
the New River. Technical Bulletin, Virginia .Agricultural
Experiment Station No. 145. 35 p.
Stansbery. D. H 1980. The naiad mollusks of the Kanawha
River below Kanaw ha Falls with special attention to en-
dangered species (Bivalvia: Unionidae). Unpublished Re-
port, Ohio State University Museum of Zoology. Colum-
bus, 16 p.
Taylor, R. W. 1983. A survey of the freshwater mussels of
the Kanawha River from riverhead (Gauley Bridge, WV)
to river mouth (Point Pleasant, W\). Final Report to U.S.
Army Corps of Engineers, Huntington District, 61 p.
Tolin. W. A. 1985. Survey of the freshwater mussels of the
New River (VVylie Islands to Bluestone Lake), lower Blue-
stone River, and lower Indian Creek, Summers County,
West N'irginia. Proceedings of the New River Symposium
4:19-26.
Turgeon, D. D., A. E. Began, E. V. Coan, W. K. Emerson, W.
G. Lyons, W. L. Pratt, C. F. E. Roper, A. Scheltema, F.
G. Thompson, and J. D. Williams. 1988. Common and
scientific names of aquatic invertebrates from the United
States and Canada: mollusks. American Fisheries Society
Special Publication 16, 277 p.
THE NAUTILUS 103(4): 140-142, 1990
Page 140
Copper, Lead and Cadmium Concentrations in a Sample of
Lake Winnipeg Anodonta grandis
Eva Pip
Department of Biology
University of Winnipeg
Winnipeg, Manitoba
Canada R3B 2E9
ABSTRACT
A sample of Anodonta grandh from soutliern Lake Winnipeg
was analyzed for tissue content of cadmium, copper and lead.
Copper and lead concentrations in tissues were each signifi-
cant!) inversely correlated witli shell weight and shell length;
copper was also significantly inversely correlated with tissue
dr\ weight. However total body content of cadmium, copper
and lead was each positive!) correlated with shell weight, shell
lengtli and tissue weight Thus, while older individuals carried
greater total body burdens for all 3 metals, younger individuals
showed higher levels of copper and lead per unit body weight.
Key words: Copper, Lead, Cadmium, Anodonta
INTRODUCTION
Unionid clams form an important component of the ben-
thic communities of many freshwater lakes. These mol-
lusks have much longer lifespans than most other fresh-
water invertebrates. Thus they are exposed to their
environment, and particularly to toxicants such as heavy
metals, for a longer period, pre.senting a greater potential
for tissue accumulation. Although concentrations of heavy
metals have been studied in freshwater insects {e.g.,
Brown, 1977; Burrows & Whitton, 1983), zooplankton
(e.g., Mathis & Kevern, 1975) and gastropods as well as
other macroinvertebrates (e.g.. Gale et ai, 1973; Leland
& McNurney, 1974; Namminga et ai, 1974; Enk &
Mathis, 1977; Mathis et ai, 1979; Newman & Mcintosh,
1982), corresponding data concerning imionids are dif-
ficult to find in the literature. Some experimental studies
have been conducted on the toxicity of metals such as
cadmium (Lukacsovics & Salanki, 1964; Radhakrish-
naiah, 1988) and copper (Imiay, 1971) to freshwater clams,
but field studies are rare. Mathis and Cummings (1971,
1973) reported on metal levels in Illinois River clams and
found that whole-animal tissue concentrations exceeded
the concentrations found in the water. Anderson (1977)
included a few unionid individuals in a survey of macro-
invertebrates from the Fox River.
In the present study cadmium, copper and lead tissue
contents of a sample of Anodonta grandis Say, 1829 from
Lake Winnipeg were compared with shell length, shell
weight and animal weight.
MATERIALS AND METHODS
A random sample of 33 A. grandis was collected on
September 28, 1986 at Sandy Hook, on the western shore
of Lake Winnipeg's south basin (50°32'N, 96°.59'W). The
clams were taken to the laboratory, where they were
kept in an aquarium for 3 days in order to evacuate the
gut. The clams were then frozen, freeze-dried and the
tissues of each individual ground to a powder.
Each tissue sample was divided for assay into 3 equal
aliquots, each consisting of up to 0.5 g of dry powder.
Each aliquot was digested by adding 7.5 ml of concen-
trated nitric acid (HNOj) and 1.5 ml 70% perchloric acid
(HCIO4). The mixture was heated but not allowed to boil
for 1 hr. Ten ml of 1% nitric acid were then added. The
solution was filtered through Whatman No. 541 hard-
ened ashless lead-free filter paper to remove insoluble
material. The filtrate was diluted to a volume of 50 ml
with 1% nitric acid and aspirated into a 1L151 atomic
absorption spectrophotometer (Instrumentation Labo-
ratory Inc., Wilmington, MA, USA). The method of stan-
dard additions was used to compensate for matrix ab-
sorption effects (e.g., Newman & Mcintosh, 1982).
Procedural controls, consisting of all reagents and steps
in the procedure less the sample, were run with each
series. All glassware was acid-washed prior to use.
Sediment samples consisting of the top 3 cm layer of
sediment were freeze-dried. One gram samples were
analyzed in triplicate, each extracted with the same acid
mixture as above (Bolter et ai, 1975). Unfiltered water
samples 1 L in volume were frozen and freeze-dried.
The residue was resuspended in a measured volume of
10% nitric acid, heated for 1 hr and aspirated for analysis.
The critical significance level for all statistical tests was
p < 0.05.
RESULTS
Shell weight and shell length were significantly corre-
lated (r = 0.97, p < 0.001) (both variables log trans-
Pip, 1990
Page 141
Table 1. Suinnuir\ nl p;iiaiiicti'rs tor the Aitodonta finindis
sample l,n = :W. Xalut's in parentheses are standard errors.
Mini-
X
nmm
Ma.xinumi
Shell « eight (g)
4.1 (0.5)
1.2
15.6
Shell length (mm)
49.5(1.3)
36.4
71.9
Aiiinuil tissue dr\
weight (g)
0.79(0.07)
0.22
1.55
Cadmium (fjg g dr\
tissue weight)
;3.0(<0.5)
<1.0
10
Copper (Mg g dry
tissue weight)
45.3(3.1)
5.0
80
Lead (Mg/g dr>-
tissue weight)
97.8(7.1)
<4
>150
that appro.\iniatei\ iialf ol the variance in the metal
content in A. p,randiH could be accounted for by dry
weight of the animal: 41% for copper (R- = 0.41, p <
0.001) and 52% for lead (R- = 0.52, p < 0.001). Shell
weight also entered the equations, but with a negative
beta value in both cases, increasing the R- values to 0.51
and 0.65, respectively (p < 0.001). For cadmium, dry
tissue weight was the only variable entered into the equa-
tion (R- = 0.55, p < 0.001) (both independent and de-
pendent variables log transformed).
Metal concentrations in the water and the coarse sand
sediments were below the detection limits for the pro-
cedure used (i.e., < 1 Mg for cadmium, <2 fig for copper
and <4 ng for lead per gram dry sample, or per liter of
water).
formed). Dry weight of tissue was significantly correlated
with shell weight (r = 0.81, p < 0.001) and shell length
(r = 0.85, p < 0.001) (all variables log transformed).
Concentrations of cadmium, copper and lead in the
tissues are summarized in Table 1. Copper content was
highly significantly negatively correlated with shell weight
(r = -0.60, p < 0.001) and shell length (r = -0.58, p
< 0.001) (untransformed variables). Logarithmic trans-
formations of all variables improved both correlation
coefficients to —0.63 (p < 0.001). Lead concentration
was also negatively correlated with shell weight and shell
length (r = -0.39, p = 0.017 and r = -0.40, p = 0.015,
respectively) (transformed variables). Correlations with
cadmium concentrations were not significant.
Correlations between metal concentrations and the dry
weight of the animal (excluding shell) were significant
and inverse for copper (r = —0.44, p = 0.006), but not
significant for cadmium or lead. Concentrations among
the three metals were correlated only between copper
and lead (r = 0.71, p < 0.001) (transformed variables).
The total body burden (excluding shell) of each metal
was significantly positively correlated with dry weight
of animal for all three metals [cadmium r = 0.47, p =
0.003; copper r = 0.64, p < 0.001; lead r = 0.77, p <
0.001 (untransformed variables)]. For cadmium, r im-
proved to 0.75 (p < 0.001) when cadmium content was
log transformed.
Total body content of copper (r = 0.38, p = 0.015)
and lead (r = 0.40, p = 0.013) were correlated with log
shell length; for log shell weight these correlations were
also significant (r = 0.33, p = 0.031 for copper, and r =
0.37, p = 0.022 for lead). For cadmium, correlations
increased for shell weight (r = 0.58, p = 0.001) and shell
length (r = 0.65, p < 0.001) when metal content was
log transformed also.
Total cadmium content was significantly correlated
with copper (r = 0.74, p < 0.001) and with lead (r =
0.54, p = 0.003). Copper and lead levels were correlated
as well (r = 0.82, p < 0.001) (all transformed).
Stepwise multiple regression with shell length, shell
weight and dry animal weight in the regression block
(all log transformed) and total body burden of each metal
as the independent variable (not transformed) indicated
DISCUSSION
The values obtained for A. grandis in the present study
were comparable to those reported by Anderson (1977)
for mussels from the Fox River, Illinois. The results showed
that as clams increase in size and weight, their total body
burdens of metals increase. However when metal con-
centrations were examined in tissues, concentrations of
copper and lead decreased as clam size increased, sug-
gesting that smaller individuals take up this metal at a
greater rate (or dispose of it more slowly) than do larger
clams of the same species. While size also probably re-
flected age of the mussels, A. grandis is known to exhibit
considerable variation in size at a given age within the
same body of water (Hanson et a/,, 1988).
LITERATURE CITED
.Anderson, R. V. 1977. Concentration of cadmium, copper,
lead,- and zinc in thirty-five genera of freshwater macroin-
vertebrates from the Fox River, Illinois and Wi.sconsin.
Bulletin of Environmental Contamination and Toxicology
18:345-349,
Bolter, E., T. Butz, and J. F. .-^rseneau 1975. Mobilization of
heavy metals by organic acids in the soils of a lead mining
and smelting district, hi: Trace substances in environ-
mental health — IX. L niversitv of Missouri, Columbia, p.
107-112.
Brown, B, E. 1977. Effects of mine drainage on the River
Hayle, Cornwall. A) Factors affecting concentrations of
copper, zinc and iron in water, sediments and dominant
invertebrate fauna. Hydrobiologia 52:221-233.
Burrows, I. G. and B. A, Whitton. 1983. Heavy metals in
water, sediments and invertebrates from a metal-contam-
inated river free of organic pollution. Hvdrobiologia 106:
263-273.
Enk, M. D. and B. J. Mathis. 1977. Distribution of cadmium
and lead in a stream ecosvstem. Hvdrobiologia 52:15.3-
158.
Gale, N. L., B. G. Wixson, M. G. Hardie, and J. C. Jennett.
1973. Aquatic organisms and heavy metals in Missouri's
New Lead Belt. Water Resources Bulletin 9:673-688.
Hanson, J. M., W. C. Mackay. and E E Prepas. 1988. The
effects of water depth and density on the growth of a
unionid clam. Freshwater Biology 19:345-355.
Imlay, M. J. 1971. Bioassay tests with naiads. In: Jorgensen,
Page 142
THE NAUTILUS, Vol. 103, No. 4
S. E. and R. W. Sharp (eds.). Proceedings of a Symposium
on Rare and Endangered Mollusks (Naiadsi of the U.S.
US Department of the Interior, Fisli and \\ ildlife Service,
Bureau of Sport Fisheries and W'ildhfe, p 38-41
Leland, H \' and J. M McNurney 1974. Lead transport in
a river ecosystem. Proceedings of the International Con-
ference on Transport of Persistent (Chemicals in .\quatic
Ecosystems, Ottawa. 111:17-23.
Lukacsovics, F. and J Salanki. 1964 Data to the chemical
sensitivity of freshwater mussel (Anodonta cijgnea L.).
.-Vnnals of the Institute of Biology (Tihany), Hungarian
.Academy of Sciences 352.5-34.
Vtathis, B J and T F Cummings. 1971. Distribution of
selected metals in bottom sediments, water, clams, tubi-
ficid annelids and fishes of the middle Illinois River. Uni-
versity of Illinois Water Resources Research Center, Re-
search Report No. 41. 45 p.
Mathis, B. J. and T. F. Cummings. 1973. Selected metals in
sediments, water, and biota in the Illinois River Journal
of the Water Pollution Control Federation 45:1573-1.583.
Mathis, B. J., T. F. Cummings, M. Cower, M. Ta\ lor, and C.
King. 1979 D\ namics of manganese, cadmium, and lead
in e\[)erimental [)ower plant ponds Hxdrobiologia 67:197-
206
Mathis, B. J. and \ R Kevern. 1975 Distribution ot mercury,
cadmium, lead and thallium in a eutrophic lake. Hydro-
biologia 46:207-222.
-Nanmiinga, H. E., J. E. Scott, and S. L. Burks. 1974. Distri-
bution of copper, lead, and zinc in selected components
of a pond ecos\stem Proceedings of the Oklahoma .Acad-
em\ of Sciences 5462-64.
Newman, M C^ and \. W Mcintosh. 1982. The influence of
lead in components of a freshwater ecosystem on mollus-
can tissue lead concentrations .•\quatic Toxicolog\ 2:1-19.
Radhakrishnaiah, K. 1988. Effect of cadmium on the fresh-
water mussel, Lamellidens marginalis (Lamarck) — a
physiological approach Journal of Environmental Biology
9 (1 supplement):73-78.
THE NAUTILUS 103(4): 143-148, 1990
Page 143
A New Fossil Land Snail (Gastropoda: Pulmonata: Polygyridae)
from the Middle Miocene of Northern Florida
Kurt AufTenberg
Roger W. Portell
Florida Museum of Natural History-
University of Florida
Gainesville, FL 32611, USA
ABSTRACT
A new fossil land snail, Praficalella prisca n sp., is described
from the C'harlton Member of the Coosavvhatchie Formation
(middle Miocene) at Brooks Sink, Bradford (>ounty, Florida.
This new species is assigned to Praticolella s.s. Martens, 1892
based on sculpture, palatal lip expansion and a narrow con-
striction behind the lip. This subgenus is presently confined to
central and southern Texas and Mexico under environmental
conditions known to have occurred in northern Florida during
the Miocene. Praticolella prisca n. sp. appears to be the earliest
know n member of the genus.
Key words: Gastropoda; Polygyridae; Praticolella. Miocene;
Florida.
INTRODUCTION
Collections of invertebrate fossils containing a new species
of terrestrial gastropod, Praticolella prisca n. sp., were
made by the authors in the mid 1980's from Brooks Sink,
a large, nearly circular, vertical-walled sink hole located
16.89 kilometers west of Starke, Bradford County, Flor-
ida (figure 1 ). Brooks Sink provides one of the best natural
exposures of Hawthorne Group sediments in Florida.
Exposed in approximately 23.0 meters of section are, in
ascending order, the Marks Head Formation, the undif-
ferentiated Coosawhatchie Formation, and the Charlton
Member of the Coosawhatchie Formation (figure 2).
The age of the Charlton Member (formerly Charlton
Formation) was considered to be Pliocene by Veatch and
Stephenson (1911) and Cooke (1943, 1945). Based upon
ostracods from this unit, identified by Harbans Puri as
middle Miocene and/or upper Miocene, Pirkle (1956)
reported the age to be older. More recently, Jones and
Portell (1988) recognized the middle Miocene clypeas-
teroid echinoid, Abertella aberti (Conrad, 1842) from
this unit. Huddleston (1988) assigned an age of middle
Miocene to the Charlton Member in Georgia based on
molluscan faunas, stratigraphic relationships, and the oc-
currence of several age-diagnostic planktonic foraminif-
era. Jones and Portell (1988) reported over 30 fossil in-
vertebrate taxa within the Charlton Member of the
Coosawhatchie Formation at Brooks Sink including Pra-
ticolella sp., the taxon described below.
MATERIALS AND METHODS
No fossilized shell material was recovered. The speci-
mens representing this taxon are preserved only as in-
ternal and external molds comprised of fine-grained do-
lostone. We use the term external mold to indicate the
impression in the matrix of the outer surface of the shell.
The matrix containing the external mold UF 14397 was
reduced and the specimen sonicated for observation of
the shell ultrastructure under a scanning electron mi-
croscope. Some of the specimens are incomplete in cer-
tain aspects. Nevertheless, standard shell parameters were
measured with vernier calipers whenever possible. All
specimens are reposited in the Florida Museum of Nat-
ural History, Invertebrate Paleontology Division, Uni-
versity of Florida, Gainesville, Florida 32611.
DESCRIPTION
Family Polygyridae Pilsbry, 1930
Subfamily Polygyrinae Pilsbry, 1895
Genus Praticolella Martens, 1892
Praticolella prisca new species
(figures 3-10, table 1)
Adult shell large (width 10.0-13.9 mm, height 6.7-10.0
mm); helicoid, depressed-globose, 0.64-0.72 times as high
as wide; spire moderately elevated, convex in outline
(figures 3, 6, 7); base round and inflated; the 4.6-5.0
convex whorls slowly increasing in size (figure 4); sutures
impressed; body whorl large, slightly flattened at suture,
round at periphery and below; deflection variable, slight-
ly upward or downward in the final 0.25 whorl, but
always descending very slightly before constricting and
inflecting; narrowly constricted posterior to palatal lip,
more deeply constricted along base (figures 4, 6); um-
bilicus narrow, tubular, about 0.10 the diameter of shell
and partially covered by reflected columellar lip (figures
5, 8); the 1.7 protoconch whorls are smooth except for a
Page 144
THE NAUTILUS, Vol. 103, No. 4
Table I. Pratuulclla prisra new species. Linear shell mea-
surements (mm I and whorl counts of holot\pe and 9 paratvpes.
Most of the s[>ecimens are incomplete in some aspects, hence
the different N for the measured parameters.
Figure I . Location of Brooks Sink in Bradford Countv, Florida
(SW/,, SW'/i, sec. 12, T7S, R20E, Brooker 7,5 Minute Quad-
rangle).
few a.\ial striations on the last 0.20 turn (figure 10); sub-
-serjiient whorls are smooth except for a few a.xial stria-
tions on the last 0.20 turn (figure 10); subsequent whorls
and base with fine, irregular, oblique axial striations,
strongest at the suture (figure 9); spiral sculpture absent;
aperture lunate and relatively large, 0.85 times as high
as wide; parietal and palatal barriers absent; palatal lip
broadly expanded, thickened within (figure 7); in lateral
view lip moderately reflected along periphery and base,
expanded forward above, unreflected (figure 6).
Etymology: The species name prisca is derived from
the Latin priscus, meaning ancient.
Type locality: Brooks Sink, 16.89 kilometers west of
Starke, Bradford County, Florida (SW'/4, SWA, sec. 12,
T7S, R20E, Brooker 7.5 Minute Quadrangle), Coosa-
whatchie Formation, Charlton Member, approximately
8.0 meters below surface.
Holotype: UF 14446, 12.5 mm width, 8.9 mm height,
internal mold, protoconch missing.
Paratypes: Width 10.0-13.9 mm, height 6.7-10.0 mm;
UF 14395, 5 adults, internal molds; UF 14397, 1 adult,
external moid and rubber peel; UF 14406, 1 adult, ex-
ternal mold; UF 14407, 1 adult, external mold and rubber
peel; UF 14408, 1 adult, internal mold; UF 14450, 1
partial adult body whorl, internal mold.
Other material examined: UF 28983, 9 fragments, in-
ternal molds; UF 14405, 1 .subadult, external moid and
rubber peel; UF 14433, 1 adult, external mold and rubber
peel; UF 14443, 1 adult, internal mold; UF 22347, 4
adults and 3 fragments, internal molds; UF 22348, 6
juveniles, internal molds, obtusely anguiate at periphery
and have indications of growth stoppage at 3.5-4.0 whorls.
COMPARATIVE REMARKS
The external mold UF 14405 (figures 7, 8) is tentatively
a.ssigned to this new species. It is slightly smaller and
more depressed (10.5 mm wide, 6.7 mm height) than
most of the other specimens. The palatal lip is simple.
Character
N
Range
X
SD
.Shell width
10
10.0-13.9
12.0
1.2
Shell height
5
6.7-10.0
8.5
1.1
Height/width
5
0.64-0.72
0.68
0.03
Aperture height
2
5.5-6.2
5.9
0.4
Aperture width
2
6.4-7.2
6.8
0.4
.\p. height, Ap. w
idth
2
0.85
0.85
0.0
Umbilicus width
7
0.8-1.5
1.1
0.2
Umb. width/shell
width
•7
0,07-0 11
0.09
0.02
Whorls
4
4 75-5 5
5.2
0 3
except basally where it is barely reflected. The body
whorl is slightly constricted on the basal portion. How-
ever, this specimen is identical to the larger, more globose
specimens in other character-states such as sculpture,
spire whorlation and umbilicus. It is our opinion that this
is a small subadult of Praticolella prisca n. sp. and does
not represent a second undescribed taxon.
The classification of pulmonate land snails is based
primarily on features of the soft anatomy making as-
signment of fossil forms difficult. Within the Stylom-
matophora several families have evolved strikingly sim-
ilar shell forms. Convergences in helicoid families such
as the Camaenidae, Polygyridae and Helminthoglypti-
dae are frustratingly commonplace. However, careful
analysis of various morphological shell character-states
can elucidate differences and general trends within and
between families. The combination of sculpture, palatal
lip expansion and the narrow constriction of the body
whorl suggest that this species is best assigned to the
polygyrid genus Praticolella Martens, 1892.
Praticolella Martens, 1892 is characterized in shell
form by its relatively small size, depressed-globose shape,
rounded periphery, narrow umbilicus and rounded base.
The aperture is lunate and the lip is slightly expanded
or reflected and thickened within. The sculpture is of
fine, irregularly spaced axial striations (Pilsbry, 1940).
Praticolella is divided into three subgenera based on
genitalic anatomy and sculpture of the protoconch (Pils-
bry, 1940). The subgenus Filapex Pilsbry, 1940 has dis-
tinct spiral sculpture on the protoconch. Farragutia Va-
natta, 1915 and Praticolella s.s. Martens, 1892 have
smooth protoconchs. Farragutia is relatively smaller than
Praticolella s.s., is slightly more depressed and has a less
rounded base. Praticolella prisca n. sp. is clearly referred
to Praticolella s.s. A close relationship to any extant species
cannot be made.
The polygyrid genera Mesodon s.s. Rafinesque, 1821,
Neohelix s.s. Ihering, 1892 and Praticolella Martens, 1892
are quite similar in general shell shape. Mesodon s.s. and
Neohelix s.s. generalK have distinct, regular axial striae
and engraved spiral lines, while Praticolella has a
smoothish shell sculptured only by weak, irregular axial
K. Auffenberg and R. W. Portell, 1990
Page 145
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LEGEND
I I SANO
(i???^ DOLOMITIC CUAYEY SAND
pag SANDY CLAY
DOLOMITIC CLAY
^^ SANDY LIMESTONE
^^ SANDY SHELLY DOLOMITIC LIMESTONE
ff=r^ DOLOMITE
|:|??| SANDY DOLOMITE
{^ SANDY DOLOMITE W/CLAV CUASTS
^g SANDY SHELLY DOLOMITE
I P I PHOSPHORITE
BOTTOM OF SINK
Figure 2. Geologic section at Brooks Sink modified from Scott (1982). Asterisk indicates zone where Praticolella prisca new species
growth striae. The maximum shell size of Praticolella
and the minimum size of Mesodon s.s. overlap, while
Neohelix s.s. is much larger. The relative palatal lip width
and degree of reflection of Praticolella rarely attains that
of Mesodon s.s. The possibilit\ that this new species is a
small unsculptured Mesodon s.s. cannot be dismissed.
Praticolella prisca n. sp. superficially resembles the
Miocene camaenids Pleurodonte crusta (Dall, 1890) and
Pleurodontites diespiter (Dall, 1890). These species.
however, have papillose sculpture, shells which are larger
and more depressed than Praticolella prisca n. sp. and
body whorls which descend greatly over the last V4 turn.
DISCUSSION
The method of preservation of fossil mollusks has an
important bearing on the interpretation of certain mor-
phological characters. Internal molds are particularly dif-
Page 146
THE NAUTILUS, Vol. 103, No. 4
Figures 3-6. Praticolella prisca new species. Holotype, UF 14446, apertural, dorsal, basal, and lateral views, x6.4.
ficiilt to interpret. For example, if a shell with an inter-
nally thickened palatal lip is filled with and encased in
sediment and the shell dissolves, the thickened portion
of the lip may appear as a constriction on the outside of
the body whorl. A true constriction may be greatly en-
hanced by this process. Praticolella prisca may in fact
be only slightly constricted behind the lip or not con-
stricted at all, only thickened internally. With more ad-
equate material the true nature of this character-state
will be better known.
To our knowledge Praticolella prisca n. sp. represents
the oldest known member of this genus. Fossil forms of
Praticolella s.s. Martens, 1892 from probably the late
Pleistocene epoch of Texas are attributed to extant species
(Hubricht, 1983, 1985). This subgenus is presently con-
fined in its native distribution to savannah and xeric
habitats in southern and central Texas and Mexico. A
similar savannah-like environment is known to have oc-
curred in northern and central Florida during the Mio-
cene (Webb, in press). This ecological association is com-
patible with the taxonomic assignment of Praticolella
prisca n. sp.
Thirteen genera (23 species, 1 subspecies) of land snails
have been recorded from the Miocene sediments of Flor-
ida (Dall, 1890, 1915; Mansfield, 1937). Eleven of these
genera are extant. If current taxonomic assignments are
correct, eight genera (61%) have Caribbean affinities,
four genera (31%) probably emigrated from areas west
K. Auffenberg and R. W. Portell, 1990
Page 147
Figures 7-10. Praticolella prisca new species. 7, 8. UF 14405, apertural and basal views of rubber peel of external mold of
probable subadult (see Comparative Remarks), x6.6. 9, 10. Paratype, UF 14397, scanning electron micrograph of protoconch of
external mold. 9. x28. 10. x47.
of Florida, and one genus (8%) came from tfie north.
The buiimuiid genus Hyperaulax Pilsbry, 1897, which
has an enigmatic distribution of several species from the
Miocene of Florida and a single extant species on Fer-
nando de Noronha Island off Brazil, is considered here
to be part of the Caribbean component of the Miocene
fauna. Many of the Miocene terrestrial vertebrates of
Florida are believed to have been associated with both
mesic and xeric tropical habitats (Auffenberg, 1963; Webb
1978, in press) and some appear to be closely related to
groups now found in dry regions of Central America and
Mexico. It is possible that some of these Florida Miocene
land snails attributed to genera presently found only in
the Caribbean region may be more appropriately as-
signed to genera occurring in mesic or xeric areas of
Central America and Mexico and thus entered Florida
via a dry tropical corridor along the Gulf of Mexico.
The fifty genera of native land snails presently found
in Florida reveal a different zoogeographic pattern than
is suggested by the fossil record. Of this current land
snail fauna, sixteen genera (32%) are well-represented in
the Caribbean region, six genera (12%) probably entered
Florida from the West, while twenty-eight (56%) came
from the North. This present zoogeographic pattern re-
flects the strong post-Miocene trend toward cooler, wetter
climates, allowing immigration of the northern taxa into
Florida. However, the timing of this immigration can
not be determined from the fossil record.
Page 148
THE NAUTILUS, \ol. 103, No. 4
ACKNOWLEDGMENTS
We are grateful to Glen Harris and Joe Gissy of the
Container Corporation of America for access to Brooks
Sink. We also thank Fred G. Thompson and Douglas S.
Jones of the Florida Museum of Natural Histor\ for re-
vicwiiig the manuscript. Kenneth C. Emberton of the
Philadelphia Academy of Natural Sciences also reviewed
the manuscript and shared his knowledge of the Poly-
gyridae. We also acknowledge Walter Auffenberg. S.
David Webb, and Gary S, Morgan of the Florida Museum
of Natural History for their insight into the vertebrate
paleontology of Florida. Micrographs were taken on a
Hitachi 5-415A scanning electron microscope, Depart-
ment of Zoology, University of Florida. This paper rep-
resents University of Florida Contribution to Paleobiol-
ogy 348.
LITERATURE CITED
Auffenberg, W. 1963. The fossil snakes of Florida Tulane
Studies in Zoology 10(3): 131-216.
Cooke, C. W. 1943. Geology of the Coastal Plain of Georgia.
U.S. Geological Survey Bulletin 941:1-121
Cooke, C. W 194.5 Geology of Florida. Florida Geological
Survey Bulletin 29:1-339.
Dall, W. H. 1890. 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. Part 1. Transactions of the Wagner Free Institute
of Science of Philadelphia 3:1-200.
Dall, VV H 191.5 Monograph of the molluscan fauna of the
Orthaulax pugnax Zone of the Oiigocene of Tampa, Flor-
ida Bulletin of the L nited States National Museum 90:1-
167.
Hubricht. L. 1983. The genus Praticolella in Texas (Poly-
g\ridae: Puimonata) \'eliger 25(3):244-250
Hubricht, L. 1985. The distributions of the native land mol-
lusks of the Eastern United States. Fieldiana, Zoology, new
series, 24:1-191.
Hiiddieston, P. F. 1988. A revision of the lithostratigraphic
units of the Coastal Plain of Georgia, the Miocene through
Hoiocene Georgia Geological Survey Bulletin 104 1-162
Jones, D. S and R W Portell 1988 Fossil in\ertebrates from
Brooks Sink, Bradford County. Florida. In: Southeastern
Geological Society Annual Field Trip Guidebook, p. 41-
52.
Mansfield, W. C. 1937. Mollusks of the Tampa and Suwannee
Limestones of Florida. Florida Geological Survev Bulletin
15:1-334.
Pilsbry, H. A. 1940. Land mollusca of North .\merica (.North
of Mexico), Vol. 1, pt. 2. The .-Xcademy of Natural Sciences
of Philadelphia, Monograph 3:574-994
Pirkle, E. C. 1956 The Hawthorne and .-Vlachua Formations
of Alachua County, Florida Florida Academy of Sciences
Quartedy Journal 19(4)197-240.
Scott, T. M. 1982. A comparison of the cotype localities and
cores of the Miocene Hawthorn Formation in Florida. In:
Scott, T. M. and S. B. Upchurch (eds. ). Miocene of the
Southeastern United States. Florida Geological Survev Spe-
cial Publication 25:237-246.
V'eatch, J. O and L \V. Stephenson. 191 1. Preliminary report
on the geology of the Coastal Plain of Georgia. Georgia
Geological Survey Bulletin 26:1-466.
Webb, S. D. 1978. The history of savanna vertebrates in the
New World. Part II: South America and the Great Inter-
change. .Annual Review of Ecolog\ and S\ stematics 9:393-
426.
Webb, S. D. In press. A biogeographic history of Florida In:
Ewel, J. and R. Myers (eds). Ecosystems of t'lorida. .Vc-
ademic Press.
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THE NAUTILUS
Supplement 1
(Issued with Volume 103)
March 6, 1990
ISSN -0028- 1344
A quarterly devoted
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[ Marine Biological Laboratory
LIBRARY
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EDITOR-IN-CHIEF
Dr. M. G. Harasewych
Division of Mollusks
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History
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Division of Mollusks
National Museum of
Natural History
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National Museum of
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Wlarine Biological Laboratory ►
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MAR 2 0 1990 \
4
'"--ds Hole, Mass. I
Catalogue of the Superfamily Cancellarioidea Forbes and
Hanley, 1851 (Gastropoda: Prosobranchia)
Richard E. Petit
and
M. G. Harasewych
Department of Invertebrate Zoology
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560 USA
THE NAUTILUS, Supplement 1: 1-69, 1990
Page 1
Catalogue of the Superfamily Cancellarioidea Forbes and
Hanley, 1851 (Gastropoda: Prosobranchia)
Richard E. Petit
M.G. Harasewych
.department of Invertebrate Zoology
National Museum of Natural History
Smithsonian Institution
Washmgton, DC 20560 USA
ABSTRACT
This catalogue of cancellarioidean taxa is composed of three
sections. The first lists alphabetically 124 genus-group taxa
originally proposed or later included in the superfamily. The
second section lists alphabetically over 1,800 species-group
taxa erected in or subsequently referred to genera now
considered to belong in Cancellarioidea. The third section
consists of a bibliography of works on the systematics of
cancellarioideans referred to in this paper.
key words: Cancellarioidea, genus-list, species-list, bibliogra-
phy.
INTRODUCTION
The Cancellarioidea comprise a highly-specialized, poorly
known, and taxonomically enigmatic group of marine
neogastropods that inhabit subtidal to bathyal sand and
mud bottoms of tropical and temperate latitudes. Al-
though most taxa at all levels within the superfamily
were described on the basis of shell characters, it was
a series of adaptations to suctorial feeding that resulted
in a succession of adaptive radiations beginning during
the late Mesozoic that gave rise to the Cancellarioidea.
Diagnostic synapomorphies of the superfamily include:
an anteriorly tubular, posteriorly saddle-shaped jaw that
spans the dorsal surface of the buccal mass; a uniserial
radula composed of extremely elongated (LAV>15),
ribbon-like, tricuspid teeth with secondary dentition on
one or more cusps [there are several aradulate taxa]; an
extremely short anterior esophagus, with the valve of
Leiblein and buccal ganglia situated at the posterior
margin of the buccal mass; and the lack of a gland of
Leiblein. Although the superfamily forms a mono-
phyletic and morphologically compact natural assem-
blage, its relationships to other prosobranchs is less than
certain. The group had, at various times, been included
in the Toxoglossa (Troschel, 1865; Tryon, 1882; Fis-
cher, 1883), the Volutoidea (Thiele, 1929; Wenz, 1943;
Taylor and Sohl. 1962) and its own order, Nemato-
glossa (Olsson, 1970). Most subsequent authors follow
Ponder (1973) in considering the Cancellarioidea a
superfamily within Neogastropoda. Taylor et al. (1980)
speculated that the Cancellarioidea are an early offshoot
from the Mesozoic Purpurinidae.
In the Tenth Edition of Sysiema Naturae Linne
(1758) described a single species of cancellariid, in-
cluding it in the genus Mure.x. In 1767, he transferred
that species to Valuta, and described a second species,
also as Valuta. By the end of the 18th century, twelve
nomina had been proposed for cancellarioideans. allo-
cated to the genera Mure.x, Voluta, Tritonium, Buc-
cinum and Cantharus. To date, there have been about
1,800 species-group taxa proposed in the Cancellarioi-
dea, many as 'varieties' of Tertiary species.
Lamarck (1799) was the first to recognize that the
then known cancellariids comprised a distinct group and
erected the genus Cancellaria. The second genus group
name to be proposed for a cancellariid was Trigona
Perry, 1811 (preoccupied and replaced by the equivalent
Trigonostoma Blainville, 1827). The third genus-level
name to be introduced was Admete (Kroyer in Moller,
1842). A decade or so later, the flood-gates opened and
many new genera and subgenera were introduced.
Marks (1949) published a list of supra-specific taxa in
Cancellariidae. citing 58 nominal taxa. We now list 124
genus-group taxa of which seven are emmendations,
errors, or nomina mida; seven were originally proposed
in Cancellariidae, but belong elsewhere; seven are
homonyms; and three are objective synonyms. One
hundred validly proposed taxa remain. Genus-group taxa
proposed for cancellariids have not been treated uni-
formly in the literature, as authors have differed widely
in their use of available taxa.
As noted by some authors, many of the nominal
genera and subgenera of cancellariids seem to be
unnecessary. At the same time, there are numerous
species that do not readily fit any of the named taxa.
Dall (1909a:30) wrote: "T may add that what, on a
cursory survey, appears to be a totally unnecessary
number of names has been applied to subdivisions of
the genus Cancellaria; but time fails me at present to
Page 2
THE NAUTILUS, Supplement 1
review them in detail. Most of them are based on slight
modification of form which gradually merge into one
another when a full series of specimens is considered".
This did not, however, preclude Dall from subsequently
proposing several new genus-group taxa in the family.
As a prodrome to a series of systematic revisions of
this poorly-understood superfamily, it has been our
objective to compile a list of all names proposed or
currently used for cancellarioideans. We have attempted
to make this work as complete as possible, but recog-
nize that some taxa have undoubtedly been omitted.
We request that colleagues bring to our attention any
errors and omissions, which will be published in a
future addendum.
This catalogue of superfamily Cancellarioidea is com-
posed of three sections: an alphabetical listing of
genus-group taxa, an alphabetical listing of species-
group taxa, and a bibliography of systematic works on
cancellarioideans. The genus-group list contains all taxa
at the genus-group level, as defined by Article 42 of the
International Code of Zoological Nomenclature (Third
Edition), that have been referred to the Cancellarioidea
or any of its families or subfamilies, either originally
or subsequently. Genus-group taxa originally proposed
in Cancellarioidea but later referred to other superfamil-
ies are included in our list.
The species-group list contains all taxa proposed at
the species-group level, as defined by Article 45 of the
International Code of Zoological Nomenclature (Third
Edition). Names originally proposed as subspecies,
varieties, forma or other infrasubspecific categories are
included, with the category used by the author identi-
fied. We have listed all names originally proposed in,
or now considered as being referable to, genera pres-
ently included in Cancellarioidea. In addition, we have
included several taxa that were transferred to cancel-
larioidean genera at one time, but which we now know
to have been so placed incorrectly. Only such of these
cases as might be a source of confusion are listed.
Nomina niida, excluded from some lists of taxa, are
included here, as they have appeared in print and may
cause confusion if their status is unclear.
We have also endeavored to attribute authorship
correctly, but this task was not always easy or precise.
Many names were first published by authors having
access to other workers' manuscripts or labeled speci-
mens. There are numerous instances where an author
attempted to credit a taxon to a colleague, but did so
in a manner requiring that authorship be attributed to
the author of the work and not the colleague. A series
of examples illustrate these points. In 1817 Faujas de
Saint Fond described and illustrated several species,
citing them as though they had already been published
by Lamarck. However, there is nothing in the work to
indicate that Lamarck had provided Faujas de Saint
Fond with the names, descriptions or illustrations, and
the names must be attributed to Faujas de Saint Fond.
In this case, these taxa have been attributed to Faujas
de Saint Fond by all subsequent authors, including
Lamarck. On the other hand, Brander (1766) clearly
stated that the descriptions of the species in his work
were supplied by Solander, and the names can be
attributed to Solander. All examples are not as clear.
The taxon Triton antiquatus first appeared in print in
Reeve (June, 1844), attributed by Reeve to Hinds'
"Zoology of the Sulphur vol. ii. pl.4. fig. 78", a work
not published until July, 1844. It is obvious that Reeve
had access to a "preprint" of Hinds' work, or that
some copies were distributed prior to the stated publi-
cation date. Thus authorship is attributed to Hinds as
"Hinds in Reeve".
The bibliography lists, alphabetically by senior author,
all works referred to in either of the indices or in the
introduction. We have attempted to provide the correct
dates of publication for all of the references cited, but
some of the dales may eventually be shown to be
incorrect. A number of scientific papers [primarily 19th
century] were published in two forms: as a paper in a
journal or other scientific publication, and as a separate
or part of a series of works on a given topic or by a
single author. In most cases the journal publication
appeared first, but on several occasions the separates or
collected works were distributed prior to the publica-
tion of the journal articles. Dating of such works is
difficult, as publication dates printed within these works
may be in error by as much as several years. In some
cases, only a range of dates is given. An Appendix
discusses two works that may potentially create nomen-
clatural problems.
Conventions used in the genus-group list
Genus-group taxa are listed alphabetically, followed
by the author(s) and an abbreviated bibliographic cita-
tion. Full citations appear in the Literature Cited section
that comprises the third part of this catalogue. The type
species is then listed, followed by the geological
horizon and the geographical locality for the type
species. In some cases, our annotations follow the
geographical range.
This is a non-critical list with subjective synonyms
being shown only for a few taxa. A critical review of
these taxa is currently in preparation.
Conventions used in the species-group list
The entries are listed alphabetically by species-group
name, followed by the genus in which the taxon was
first proposed (in parentheses), then by the author(s),
date and an abbreviated citation. As in the genus list,
full citations are provided in the Literature Cited
section. If the taxon was proposed as a subspecies,
variety or other infra-subspecific form, that infonnation
follows the reference. If the taxon is proposed as a
replacement name, the abbreviation n.n. (new name or
nomen novum) is followed by the citation on which the
replacement name (either of a homonym or an incor-
rectly identified figure) is based. The geological hori-
zon and geographic locality are then listed. Objective
synonyms are indicated by "=". Subjective synonyms.
R. E. Petit and M. G. Harasewych, 1990
Page 3
indicated by "?=", are given only where the listed
taxon is a junior homonym, where the name has a
confused history, or where the subjective synonymy is
thought to be helpful to users of this compilation.
Taxa incorrectly attributed are shown with the sup-
posed author's name in single quotes followed by the
citation for the reference containing the incorrect usage.
Incorrect usages are generally not attributed, except
when this information is thought to be particularly
useful. Sources for misspellings are omitted in many
cases, as are many misspellings. Only those that could
cause confusion have been included.
If the genus-group taxon cited is not currently
included in Cancellarioidea, the current placement of the
species is given in square brackets [ ]. Conversely, if
the species was originally proposed in a genus referable
to Cancellarioidea, but is not now considered a cancel-
larioidean, current placement, if known, is also given
in square brackets [ ].
ACKNOWLEDGMENTS
Publication of these lists of taxa in the Cancellarioidea
is the culmination of a bibliographic search begun in
1964 when Mr. Druid Wilson, U.S. Geological Survey,
gave one of us (REP) a notebook containing notes he
had made on cancellariid genera over a period of many
years. We are indebted to him not only for being
responsible for starting this project, but also for his
continued assistance and encouragement.
Mr. Andre Verhecken, Mortsel, Belgium, reviewed a
draft of the list of species-group taxa and compared it
with a list he had compiled. He graciously furnished us
with references to a number of taxa that we would
otherwise have overlooked.
Mr. Jacques Le Renard, Plaisir, France, reviewed a
draft of the manuscript and provided stratigraphic data
for a number of European taxa.
Compilation of these lists would not have been
possible without assistance from many people who
brought to our attention obscure references to cancel-
larioideans, and also who supplied copies of pertinent
literature. At the risk of omitting some who rendered
assistance, and to whom we apologize for such omis-
sion, we thank;
Mr. Warren Blow, and Ms. Barbara Bedette, De-
partment of Paleobiology, National Museum of
Natural History; Dr. Thomas Gibson, United States
Geological Survey, Reston; Dr. Kenneth Boss, Dr.
Silvard Kool and Mr. Alan Kabat, Museum of
Comparative Zoology, Harvard University; Dr. Robert
Robertson, Dr. Arthur Bogan and Dr. Gary Rosen-
berg, The Academy of Natural Sciences of Philadel-
phia; Dr. William K. Emerson and Mr. Walter E.
Sage, III, American Museum of Natural History,
New York; Dr. Emily H. Yokes, Tulane University,
New Orleans; Ms. Kathie Way, Mr. John Cooper
and Mr. Patrick Nuttall, British Museum (Natural
History); Dr. Philippe Bouchet and M. Jean-Paul
Rocroi, Museum national d'Histoire naturelle, Paris;
Dr. Peter Jung, Naturhistorisches Museum Basel,
Basel; Dr. Franco Davoli, Instituto di Paleontologia,
Universita de Modena; Dr. Jordi Martinell, Univer-
sidad de Barcelona; Dr. Oleg V. Amitrov, Paleon-
tological Institute of the U.S.S.R., Moscow; Dr.
Thomas Garrard, The Australian Museum; Dr. Alan
Beu and Mr. Phillip Maxwell, New Zealand Geo-
logical Survey.
We have received advice on problems of nomencla-
ture from Drs. Raymond Manning and Frederick M.
Bayer, both of the Department of Invertebrate Zoology,
National Museum of Natural History, Washington, DC,
and Dr. L. B. Holthuis, Rijksmuseum Natuurlijke His-
toire, Leiden.
Finally, we would like to acknowledge the immeas-
urable assistance provided over the years by the Li-
brarians of the National Museum of Natural History, the
U.S. Geological Survey, Reston, and of the Library of
ConOTess.
Page 4
THE NAUTILUS, Supplement I
Genus-group Taxa
A
Admete Kroyer in Moller, 1842:88 (ex Kroyer MS.)
Type species, by monotypy, Admete crispa Moller,
1842 (?= Tritonium viriduliim Fabricius, 1780).
Recent, North Atlantic.
Admetopsis Meek, 1873:501. Type species, by subse-
quent designation of Cossmann (1895:753), Admete
gregaria Meek, 1873. Cretaceous, Utah, U.S.A. A
careful reading of Meek's text indicates that he
considered his Admete rhomhoides to be the type
of the new genus. This is particularly evident in the
discussion following the description of Admete
subfusiformis where he states that A. subfusiformis
and A. gregaria "may have to take the generic
name Titrricula.'' Although Cossmann (1899a:7)
removed Admetopsis from the Cancellariidae, the
genus appears to be cancellariid.
Admetula Cossmann, 1889:228. Type species, by origi-
nal designation, Cancellaria evulsa (Solander, 1766)
(= Buccinum evulsa Solander, 1766). Eocene, Eng-
land.
Africostoma Eames, 1957:49. Type species, by original
designation, Trigonostoma decorata Newton, 1922.
Eocene, Nigeria.
Africosveltia Eames, 1957:48. Type species, by original
designation, Cancellaria mitltiplicis Newton, 1922.
Eocene, Nigeria.
Africotriton Beu and Maxwell, 1987:29. Type species,
by original designation, Epidromiis crebriliratits
G.B. Sowerby III, 1903. Recent, South Africa.
Agatrix Petit, 1967:218. Type species, by original
designation, Trigonostoma agassizii Dall, 1889.
Recent, western Atlantic.
Anapepta Finlay, 1930b:241. Type species, by original
designation, Admete anomala Marshall and Mur-
doch, 1920. Eocene, New Zealand.
Aneiirystoma Cossmann, 1899a:23. Type species, by
original designation, Cancellaria dufourii Grate-
loup, 1832. Miocene, France.
Antepepta Finlay and Marwick, 1937:82. Type species,
by original designation, Antepepta nasitta Finlay
and Marwick, 1937. Paleocene, New Zealand.
Aphera H. Adams and A. Adams, 1854:277. Type
species, by monotypy, Cancellaria tessellata Sow-
erby, 1832. Recent, Panamic-Pacific.
Arizelostoma Iredale, 1936:318. Type species, by origi-
nal designation, Arizelostoma laseroni Iredale, 1936.
Recent. Australia.
Axelella Petit, 1988:130. Type species, by original
designation of Olssonella Petit, 1970, Cancellaria
smitliii Dall, 1888. Recent, western Atlantic. Re-
placement name for Olssonella Petit, 1970, non
Gilbert and Van de Poel, 1967.
B
Babylonella Conrad, 1865a:32. Type species, by subse-
quent designation of Cossmann (1889:231), Cancel-
laria elevata Lea, 1833. Eocene, Alabama, U.S.A.
Barkeria Addicott, 1970:118. Type species, by original
designation, Cancellaria sanjosei Anderson and
Martin, 1914. Miocene, California, U.S.A.
Benthobia Dall, 1889a: 131. Type species, by original
designation, Benthobia tryonii Dall, 1889. Recent,
North Carolina, U.S.A. Originally described in
Cancellariidae, Benthobia was transferred to Olividae
by Bouchet and Waren (1985:249).
Bivetia Jousseaume, 1887a: 163. Type species, by mono-
typy, Bivetia mariei Jousseaume, 1887 (?= Cancel-
laria indentata Sowerby, 1832). Recent, Panamic-
Pacific.
Bivetia Jousseaume, 1887b: 193. Type species, by origi-
nal designation, Cancellaria similis Sowerby, 1833.
Recent, northwestern Africa. This usage and desig-
nation is invalid due to the prior (by one month)
monotypic usage of Bivetia in the binomen Bivetia
mariei Jousseaume.
Bivetiella Wenz, 1943:1356. Type species, by original
designation of Bivetia Jousseaume, 1887b, Cancel-
laria similis Sowerby, 1833. Recent, northwestern
Africa. Replacement name for Bivetia Jousseaume,
1887b:193, non 1887a:163.
Bivetiella Marks, 1949:456. Type species, by original
designation, Cancellaria similis Sowerby, 1833.
Recent, northwestern Africa. This is a junior objec-
tive synonym and a junior homonym of Bivetiella
Wenz, 1943.
Bivetopsia Jousseaume, 1887b:193. Type species, by
subsequent designation of Cossmann (1888:784),
Cancellaria chrysostoma Sowerby, 1832. Recent,
Panamic-Pacific.
Bivetopsis Jousseaume. Unnecessary emendation of Bive-
topsia by Cossmann (1899a:9).
BonellUia Jousseaume, 1887b:223. Type species, by
original designation, Cancellaria bonellii Bellardi,
1841. Miocene/Pliocene, Italy.
Brocchinia Jousseaume, 1887b:221. Type species, by
subsequent designation of Sacco (1894:68), Brocchi-
nia mitraeformis (Brocchi) (= Volitta mitraeformis
Brocchi, 1814, non Lamarck, 1811; ?= Brocchinia
parvula tauropar\'a Sacco, 1894). Pliocene, Italy.
See Petit, 1986.
Bitccinella Perry, 1811:plate 27. Type species, by
subsequent designation of Abbott (1950:203), Buc-
cinella caerulea Perry, 1811 (?= Turbinella pyntm
(Linne, \161)). Bitccinella is included here as it has
traditionally been placed in the synonymy of Cancel-
laria s.s. until recent years. Among authors who
cited Bitccinella as a synonym of Cancellaria are
H. Adams and A. Adams (1854:275), Thiele
(1929:352) and almost all other cataloguers and
monographers. This placement was due to the fact
that Perry's first figured species, Bitccinella canit-
lata, probably represents Cancellaria reticulata
(Linne). Although Bitccinella was often listed, no
type was designated until Abbott's designation of
B. caerulea, which is presumed to be a synonym
of Turbinella pyrum (Linne). Unaware of Abbott's
R. E. Petit and M. G. Harasewych, 1990
Page 5
designation, Eames (1952:115) designated Biiccinella
canulata as type, placing Buccinella in the synon-
ymy of CanceUarki had his designation been valid.
C
Calcarata Jousseaume, 18875:214. Type species, by
original designation, Calcarata calcarata (Brocchi)
(= Valuta calcarata Brocchi, 1814). Pliocene, Italy.
Cancellaphera Iredale, 1930:80. Type species, by mono-
typy, Cancellaphera amasia Iredale, 1930. Recent,
Australia.
Cancellaria Lamarck, 1799:71. Type species, by mono-
typy. Valuta reticulata Linne, 1767. Recent, Carib-
bean.
Cancellariella G. Martin, 1904:168. Type species, by
original designation, Cancellaria (Cancellariella)
neritoidea G. Martin, 1904. Miocene, Maryland,
U.S.A.
Cancellarius Montfort, 1810:562. Type species, by
original designation. Valuta reticulata Linne, 1767.
A junior objective synonym of Cancellaria.
Cancelrana Palmer, 1937:455. Type species, by original
designation, Pleurotoma (Jaranis) finexa Harris,
1895. Eocene, Texas, U.S.A.
Caveola Stephenson, 1941:363. Type species, by origi-
nal designation, Cancellaria acuta Wade, 1926.
Cretaceous, Tennessee, U.S.A.
Charcolleria Olsson, 1942:61. Type species, by original
designation, Cancellaria {Charcolleria) perdiciana
Olsson, 1942. Miocene, Colombia.
Contortia Sacco, 1894:48. Type species, by original
designation, Cancellaria contorta Basterot, 1825.
Tertiary, France.
Coptostoma Cossmann, 1899a:34. Type species, by
original designation, Cancellaria quadrata J. Sow-
erby, 1822. Eocene, England.
Coptostomella Finlay and Marwick, 1937:83. Type
species, by original designation, Coptostomella pupa
Finlay and Marwick, 1937. Paleocene, New Zeal-
and.
Crawfordia Dall, 1918:138. Type species, by original
designation, Cancellaria crawfordiana Dall, 1891.
Recent, California, U.S.A. Not Crawfordia Pierce,
1908 (Coleoptera). Renamed Crawfordina Dall,
1919.
Crawfordina Dall, 1919:306. Type species, by mono-
typy, Cancellaria crawfordiana Dall, 1891. Recent,
California. U.S.A. Although obviously a replace-
ment name for the preoccupied Crawfordia, that
fact is not mentioned. As no type designation is
made, and it is not stated that this is a replacement
name, type designation is by monotypy.
D
Daguinia Magne, 1966:127. Type species, by mono-
typy, Daguinia vigneaiixi Magne, 1966. Miocene,
France.
Dellina Beu, 1970:223. Type species, by original desig-
nation, Waipaoa munida Ponder, 1968 (?= Anti-
zafra aoteana Dell, 1956). Recent, New Zealand.
E
Egerea Gdbor, 1936:7. Type species, by monotypy,
Egerea collectiva Gabor, 1936. Oligocene, Hungary.
Emmonsella Olsson and Petit, 1964:541. Type species,
by original designation, Trigonostoma tenerum
(Philippi, 1848) (= Cancellaria tenera Philippi,
1848). Recent, Caribbean. A junior subjective syno-
nym of Ventrilia.
Esbelta Sarasua, 1975:2. Type species, by original
designation, Ranella lanceolata Menke, 1828. Re-
cent, Caribbean.
Euclia H. Adams and A. Adams, 1854:277. Type
species, by subsequent designation of Cossmann
(1899a:10), Cancellaria cassidifarmis Sowerby, 1832.
Recent, Panamic-Pacific.
Exechoptychia Cossmann, 1903b: 189. Type species, by
original designation, Cancellaria conradiana Dall,
1890. Pliocene, Florida and the Carolinas, U.S.A.
A junior subjective synonym of Cancellaria.
Extractrix Korobkov, 1955:138. Type species, by origi-
nal designation, Pseudamalaxis extractrix (Boettger,
1906) (= Discahelix extractrix Boettger, 1906).
Miocene, Romania.
F
Fusiaphera Habe, 1961a:72, Appendix p. 27. Type
species, by original designation, Fusiaphera mac-
rospira Adams and Reeve) (= Cancellaria mac-
raspira Adams and Reeve, 1850). Recent, Japan.
G
Gerdiella Olsson and Bayer, 1972:876. Type species,
by original designation, Gerdiella gerda Olsson and
Bayer, 1972. Recent, Caribbean.
Gergovia Cossmann, 1899a: 16. Type species, by origi-
nal designation, Cancellaria platypleura Tate, 1898
(= Cancellaria laticostata Tenison- Woods, 1879).
Eocene, Australia.
Gulia Jousseaume, 1887b: 194. Type species, by subse-
quent designation of Cossmann (1888:784), Cancel-
laria acutangula Faujas de Saint Fond, 1817. Mio-
cene, France.
H
Habesolatia Kuroda, 1965:8. Type species, by original
designation, Cancellaria nodulifera Sowerby, 1825.
Recent, Japan.
Hertleinia Marks, 1949:457. Type species, by original
designation, Cancellaria mitriformis Sowerby, 1832.
Recent, Panamic-Pacific.
Hetereuclia 'Roverto' - Wenz, 1943:1357. Error for
Heteroeuclia Rovereto.
Heteroeuclia Rovereto, 1899:103. Unnecessary replace-
ment name for Euclia Adams and Adams, 1854,
which Rovereto considered to be preoccupied by
Euclea Hubner, 1816, and Euclea Newman, 1842.
I
Inennia Korobkov, 1955:328. Type species, by original
designation, Cancellaria inermis Pusch. 1837. Terti-
ary, Poland. Inermia is preoccupied and a new
name must be proposed if this is found to be a
valid genus-level group.
Page 6
THE NAUTILUS. Supplement 1
IngUsella Finlay, 1924c:513. Type species, by original
designation, Ptychatractiis pukeuriensis Suter, 1917.
Miocene, New Zealand.
Iphinoella Habe, 1958:34, 40. Type species, by mono-
typy, Iphinoella choshiensis Habe, 1958. Recent,
Japan. Placed in synonymy of Iphinopsis Dall by
Habe (1962:73) and Bouchet and Waren (1985:261).
Iphinopsis Dall, 1924:88. Type species, by original
designation, Iphiiwe kelseyi (Dall, 1908) (= Trichotro-
pis kelseyi Dall, 1908). Recent, California. U.S.A.
Transferred from Trichotropidae to Cancellariidae
bv Bouchet and Waren (1985:261).
J
Junghuhnia K. Martin, 1931:11. Type species, by
monotypy, Trigonostoma {Junghuhnia) ficus K.
Martin, 1931. Eocene, Indonesia.
K
Kapuatriton Beu and Maxwell, 1987:24. Type species,
by original designation, Kapuatriton kaitarus Beu
and Maxwell, 1987. Eocene, New Zealand.
L
Loxotaphnis G. F. Harris. 1897:165. Type species, by
original designation, Phos variciferus Tate, 1888.
Miocene, Australia.
M
Maorivetia Finlay, 1924c:513. Type species, by original
designation, Turbinella hrevirostris Hutton, 1877.
Miocene, New Zealand.
Marianarona Petuch, 1988:27. Type species, by origi-
nal designation, Cancellaria alternata Conrad, 1834.
Miocene, Maryland, U.S.A.
Mariasveltia Petuch, 1988:34. Type species, by original
designation, Cancellaria htnata Conrad, 1830. Mio-
cene, Maryland, U.S.A.
Marksella Olsson, 1964:127. Type species, by original
designation, Admele {Marksella) juniala Olsson,
1964. Pliocene, Ecuador.
Massyla H. Adams and A. Adams, 1854:278. Type
species, by monotypy, Cancellaria corrugata Hinds,
1843. Recent, Panamic-Pacific.
Mataxa Wade, 1917:455. Type species, by original
designation, Mataxa elegans Wade, 1917. Creta-
ceous, Tennessee, U.S.A.
Merica H. Adams and A. Adams, 1854:277. Type
species, by subsequent designation of Cossmann
(1899a: 13), Cancellaria melanostoma Sowerby, 1849.
Recent, Indian Ocean.
Mericella Thiele, 1929:352. Type species, by monotypy,
Cancellaria jucunda Thiele, 1925. Recent, eastern
Africa.
Microcancilla Dall, 1924:87. Type species, by original
designation, Admete microscopica (Dall, 1889) (=
Cancellaria microscopica Dall, 1889). Recent, Car-
ibbean.
Microsveltia Iredale, 1925:265. Type species, by origi-
nal designation, Microsveltia recessa Iredale, 1925.
Recent, Australia.
Misteia Janssen, 1984:16. Type species, by original
designation, Cancellaria planispira Nyst, 1845. Mio-
cene, Netherlands.
Momoebora Habe and Kikuchi, 1960:51. Nomen nu-
dum.
Momoebora "Habe (MS)', Azuma, 1960:49. Nomen
nudum.
Momoebora Kuroda and Habe, 1971:310 [Japanese],
202 [English]. Type species, by original designa-
tion, Cancellaria sinensis Reeve, 1856. Recent,
Japan.
N
Narona H. Adams and A. Adams, 1854:277. Type
species, by subsequent designation of Jousseaume
(1887b:222), Cancellaria clavatula Sowerby, 1832.
Recent, Panamic-Pacific.
Neadmete Habe, 1961a:73, Appendix p. 28. Type
species, by I. C.Z.N. Opinion 1370, Neadmete okutanii
Petit, 1974. Recent, Japan.
Nevia Jousseaume, 1887b:222. Type species, by original
designation, Cancellaria spirata Lamarck, 1822.
Recent, Australia.
Nipponaphera Habe, 1961a:72, Appendix p. 27. Type
species, by I.C.Z.N. Opinion 1052, Nipponaphera
hahei Petit, 1972. Recent, Japan.
Nivitriton Iredale, 1929b:288. Type species, by original
designation, Triton antiquatus Hinds in Reeve,
1844. Recent, Indo-Pacific.
Nothoadmete Oliver, 1982:15. Type species, by mono-
typy, Nothoadmete tumida Oliver, 1982. Recent,
Antarctica.
O
Oamaruia Finlay, 1924c:514. Type species, by original
designation, Admete suteri Marshall and Murdoch,
1920. Miocene, New Zealand.
Olssonella Petit, 1970:83. Type species, by original
designation, Cancellaria smithii Dall, 1888. Recent,
Caribbean. Not Olssonella Gilbert and Van de Poel,
1967. Renamed A.xelella Petit, 1988.
Ovilia Jousseaume, 1887b: 193. Type species, by origi-
nal designation, "O. doliaris Bast." (= Cancellaria
doUolaris Basterot, 1825). Miocene, France.
P
Paladmete Gardner, 1916:412. Type species, by original
designation, Trichotropis cancellaria Conrad, 1858.
Cretaceous, Mississippi, U.S.A.
Palaeadmete 'Gardner" - Okutani, 1964:398. Error for
Paladmete Gardner.
Palaeocancellaria Kollmann, 1976:199. Type species,
by original designation. Palaeocancellaria hoelleitenen-
sis Kollmann, 1976. Cretaceous, Austria.
Pallidonia Laseron, 1955:272. Type species, by original
designation, Pallidonia simplex Laseron, 1955. Re-
cent, Australia.
Panarona Petit, 1975:387. Type species, by original
designation, Cancellaria clavatula Sowerby, 1832.
Recent, Panamic-Pacific. An unnecessary name which
is a junior objective synonym of Narona.
Paradmete Strebel, 1908:22. Type species, by indication
(I.C.Z.N. Article 68c), Paradmete typica Strebel,
1908. Recent, Antarctica. Originally described in
Cancellariidae, Paradmete is now placed in Voluto-
mitridae.
R. E. Petit and M. G. Harasewych, 1990
Page 7
Pepta Iredale, 1925:266. Type species, by monotypy,
Admete stricta Hedley, 1907. Recent, Australia.
Perplicaria Dall, 1890:90. Type species, by monotypy,
Perplicaria perple.xa Dall, 1890. Pliocene, Florida,
U.S.A.
Peruclia Pilsbry and Olsson, 1941:24. Error for Pyi-
iiiiia Olsson, 1932.
Plesiocerithium Cossmann, 1889:232. Type species, by
original designation, "Cancellaria magloirei
Melleville, 1843" (= C. maglorii Melleville, 1843).
Lower Eocene, France.
Plesiotriton Fischer, 1884:654. Type species, by original
designation, Cancellaria volutella Lamarck, 1803.
Eocene, France.
Plicaria Fabricius, 1823:74. Fabricius' work has been
placed on the Official Index of Rejected and Invalid
Works in Zoological Nomenclature by I. C.Z.N.
Opinion 521. Prior to placement of this work on
the Official Index. Plicaria was traditionally in-
cluded in the synonymy of Cancellaria. Fames
(1952:116) designated 'T. reticulata (L.) (= Cancel-
laria reticulata (Linne))" as type.
Preangeria K. Martin, 1921:450. Type species, by
subsequent designation of Wenz (1943:1356), C.
(P.) angsanana (K. Martin) (= Preangeria angsanana
K. Martin, 1921). Tertiary, Indonesia. Martin did
not designate a type species and Wenz (1943:1356)
shows type by monotypy. This is incorrect as
Martin described two species in his new genus.
However, Wenz' citation may be taken as a subse-
quent designation. Preangeria was originally de-
scribed in Cancellariidae, but was transferred to
Purpuridae by K. Martin (1928:124).
Pristimerica Finlay and Marwick, 1937:82. Type spe-
cies, by original designation, Pristimerica dolioides
Finlay and Marwick, 1937. Paleocene, New Zeal-
and.
Procancellaria Wilckens, 1922:21. Type species, by
monotypy, Procancellaria parkiana Wilckens, 1922.
Cretaceous, New Zealand. Originally described in
Cancellariidae, Procancellaria was transferred to
Littorinidae by Finlay and Marwick (1937:81).
Progabbia Dall, 1918:138.] Type species, by original
designation, Cancellaria cooperi Gabb, 1865. Re-
cent, California, U.S.A.
Pyruclia Olsson, 1932:160. Type species, by original
designation, Cancellaria solida Sowerby, 1832. Re-
cent, Panamic-Pacific.
R
Rhomboide stoma K. Martin, 1931:12. Type species, by
monotypy, Rhomboide stoma oscitans K. Martin,
1931. Eocene, Indonesia.
S
Scalptia Jousseaume, 1887b:213. Type species, by origi-
nal designation, Cancellaria obliquata Lamarck,
1822. Recent, Indo-Pacific.
Semitriton Cossmann, 1903b: 102. Type species, by
original designation, Plesiotriton dennanti Tate,
1898. Eocene, Australia.
Solatia Jousseaume, 1887b:222. Type species, by origi-
nal designation, "Solatia solat Adams (Piscatorum
Chemn.)" (= Buccinum piscatorium Gmelin, 1791).
Recent, western Africa. "Adams" as used by
Jousseaume is an error for Adanson, whose non-
binominal work referred to the type species as Le
Solat.
Solutosveltia Habe, 1961b:433. Type species, by mono-
typy, Solutosveltia abyssicola Habe, 1961. Recent,
Japan.
Sveltella Cossmann, 1889:226. Type species, by original
designation, Cancellaria qiiantula Deshayes, 1864.
Eocene, France.
Sveltia Jousseaume, 1887b:214. Type species, by origi-
nal designation, "Sveltia varicosa Brocc." (= Vo-
luta varricosa Brocchi, 1814). Pliocene, Italy.
Sydaphera Iredale, 1929a:341. Type species, by original
designation, Sydaphera renovala Iredale, 1929. Re-
cent, Australia.
T
Tatara Fleming, 1950:247. Type species, by original
designation, Cymatium pahiense Marshall and Mur-
doch, 1921. Eocene, New Zealand.
Torellia Jeffreys, 1867:244. Type species, by monotypy,
Torellia vestita Jeffreys, 1867. Recent, North Sea.
Originally described in Cancellariidae, Torellia is
now placed in Trichotropidae.
Tribia Jousseaume, 1887b:221. Type species, by origi-
nal designation, "Tribia angasi Crosse" (= Cancel-
laria angasi Crosse, 1863). Recent, western Africa.
Trigona Perry, 1811:pl. 51. Type species, by monotypy,
Trigona pellucida Perry, 1811. Not Trigona Jurine,
1807 (Hymenoptera). This preoccupied taxon is a
subjective synonym of Trigonostoma Blainville.
Trigonaphera Iredale, 1936:319. Type species, by origi-
nal designation, Trigonostoma vinnulum Iredale,
1925. Recent, Australia.
Trigonostoma Blainville, 1827:652. Type species, by
monotypy, Delphinula trigonostoma Lamarck, 1822
(?= Buccinum scalare Gmelin, 1791). Recent, Indo-
Pacific.
Tritonoharpa Dall, 1908:319. Type species, by original
designation, Tritonoharpa vexillata Dall, 1908. Re-
cent, Galapagos Islands.
Turehua Marwick, 1943:189. Type species, by original
designation, Latirus duhius Marshall, 1919. Eocene,
New Zealand.
U
Unitas Palmer, 1947:413. Type species, by original
designation of Uxia Jousseaume, Cancellaria costu-
lata Lamarck, 1803. Eocene, France. Replacement
name for Uxia Jousseaume, 1887, non Walker,
1866.
Uxia Jousseaume, 1887b:222. Type species, by original
designation, Cancellaria costulata Lamarck, 1803.
Eocene, France. Not Uxia Walker, 1866 (Lepidop-
tera). Renamed Unitas Palmer, 1947.
Page 8
THE NAUTILUS, Supplement 1
Varicohilda Eames, 1957:40. Type species, by original
designation, Hilda tunicidata Newton, 1922. Eo-
cene, Nigeria.
Ventrilia Jousscaumc, 1887a; 164. Type species, by
monotypy, Ventrilia ventrilia Jousseaume, 1887 (=
Canccllaria tcnera Philippi, 1848). Recent, Carib-
bean.
Vercomaris Garrard, 1975:42. Type species, by original
designation, Canccllaria pcrgradata Verco, 1904.
Recent, Australia.
W
Waipaoa Marwick, 1931:122. Type species, by original
designation, Admete cristata Marwick, 1926. Mio-
cene, New Zealand.
Z
Zeadmete Finlay, 1926b:429. Type species, by original
designation, Canccllaria trailli Hutton. 1873. Re-
cent, New Zealand.
R. E. Petit and M. G. Harasewych, 1990
Page 9
Species-Group Taxa
A
ahiwrmis (Cancellaria) Gray, 1847:136. Nomcii nudum.
abnormis (Aclmete) 'Morch & Poulson MS' - Harmer,
1918:408; in synonymy of A. viridula var. couthouyi
(Jay). Nomcn nudum.
abyssicola (Soluiosvcliia) Habe, 1961b:433, pi. 23, fig.
4. Recent, Japan.
acalypta {Cancellaria) Woodring, 1970:341, pi. 53, figs.
1-2. Tertiary, Panama.
aclinica (Coluhraria) Tucker & Wilson, 1933:70, pi.
11, fig. 14. Pliocene, Florida, U.S.A. [?= Tri-
lonoharpa lanceolala (Menke, 1828)]
acosticillata {Cancellaria) Sacco, 1894:10; as var. of C.
ampullaceum (Brocchi); see Appendix, Note 2.
Pliocene, Italy.
acuminata {Cancellaria) G. B. Sowerby 1, 1832a:53;
1832b:fig. 5. Recent, Panamic-Pacific.
acuminata {Cancellaria) Bellardi, 1841:38, pi. 4, figs.
15-16. Tertiary, Europe. (= C. subacuminata d'Or-
bigny, 1852)
acuta \Cancellaria) Wade, 1926:108, pi. 35, figs. 4-5.
Cretaceous, Tennessee, U.S.A.
acutangula {Cancellaria) Faujas de Saint Fond, 1817:197,
pi. 10, figs. 1, la. Miocene, France.
acutangularis {Cancellaria) Lamarck, 1822b: 11 6. Mio-
cene, France. (= C. acutangula Faujas de Saint
Fond, 1817)
acuticarinata {Cancellaria) Weisbord, 1929:51, pi. 6,
fig. 7. Miocene, Colombia.
acutiplicata {Cancellaria) Shuto, 1962:74, pi. 10, figs.
4-5; as subsp. of C. pristina Yokoyama. Pliocene,
Japan.
adami {Cancellaria) "Eichwald" - Sherbom, 1922:70.
Error for Cassis adami Eichwald, 1830.
adelae {Cancellaria) Pilsbry, 1940:54, pi. 3, fig. 1; as
subsp. of C. reticulata (Linne). Recent, Florida,
U.S.A.
adicana {Cancellaria) Pereira Da Costa, 1867:210, pi.
25, figs. la-b. Miocene, Portugal.
aegyptiaca {Cancellaria) Oppenheim, 1906:331, pi. 26,
figs. 18a-b. Eocene, Egypt.
aequaeva {Cancellaria) Ryckholt, 1862:pl. 32, figs.
32-33. Cretaceous, Belgium.
aequicosticillata {Cancellaria) Sacco, 1894:44, pi. 3,
fig. 10; as var. of C. serrata Bronn; see Appendix,
Note 2. Miocene, Italy.
aequivaricosa {Cancellaria) Sacco, 1894:4, pi. 1, fig.
2; as var. of C. oligolongum Sacco; see Appendix,
Note 2. Oligocene, Italy.
aethiopica {Admete) Thiele, 1925:201, pi. 22, fig. 23.
Recent, eastern Africa, [not a cancellariid]
afasciata {Cancellaria) Sacco, 1894:63, pi. 3, fig. 67;
as var. of C. intermedia Bellardi; see Appendix,
Note 2. Miocene, Italy.
afenestrata {Aphera) Sacco, 1894:67; n.n. for "'C.
dufourii Grateloup" Homes, 1854:pl. 44, fig. 9; as
var. of A. hronui (Bellardi). Miocene, Austria.
affectata {Waipaoa) Marvvick, 1931:123, pi. 13, fig.
241. Miocene, New Zealand.
affinis {Cancellaria) C.B. Adams, 1852a:356. Recent,
Panamic- Pacific. (?= C. ventricosa Hinds, 1843)
affinis {Cancellaria) Reeve, 1856:pl. 9, figs. 39a-b.
Recent, Panamic-Pacific. (?= C. indentata Sowerby,
1832)
africana {Cancellaria) Petit, 1970:86; n.n. for C. imbri-
cata Watson, 1882, non Homes. 1856. Recent,
South Africa.
agalma {Cancellaria) Melvill & Standen, 1901:450, pi.
24, fig. 18. Recent, Gulf of Oman.
agassizii {Trigonostoma) Dall, 1889a: 130, pi. 35, fig.
4. Recent, Carolinas to Gulf of Mexico.
agria {Cancellaria) Mansfield, 1930:48, pi. 3, fig. 1.
Miocene, Florida, U.S.A.
alabamensis {Cancellaria) Gabb, 1860a:301, pi. 48, fig.
14. Cretaceous, Alabama, U.S.A. [Turhinella]
alaskensis {Cancellaria) B. L. Clark, 1932:832, pi. 20,
figs. 10, 12, 16, 17. Oligocene, Alaska, U.S.A.
alba {Cancellaria) Webb, 1936:125; as var. of C.
reticulata (Linne). Nomen nudum.
alba {Cancellaria) 'Sowerby' - Domenech, Espinosa,
Marquina & Martinell, 1983:45; as subsp. of C.
similis Sowerby. Nomen nudum.
alba Uphinopsis) Bouchet & Waren, 1985:263, figs.
695-697. Recent, Bay of Biscay, Europe.
albida {Cancellaria) Hinds, 1843:47; 1844b:42, pi. 12,
figs. 9-10. Recent, Panamic-Pacific.
albrechtina {Cancellaria) Mayer, 186 lb: 11 8. Oligocene,
Europe.
aldrichi {Cancellaria) Gardner, 1937:372, pi. 45, fig.
2. Miocene, Florida, U.S.A.
allophyla {Inglisella) Maxwell, 1988:70, pi. II, figs.
m-o. Miocene, New Zealand.
alta {Paladmete) Stephenson, 1941:367, pi. 71, figs.
12-13. Cretaceous, Texas, U.S.A.
altavillae {Cancellaria) Aradas, 1846:173, pi. 1, figs.
lOa-b. Tertiary, Italy.
altavillae {Cancellaria) Libassi, 1859:38, fig. 15; as
altavill. [sic]\ as var. of C. labrosa Bellardi.
Tertiary, Italy.
altavillae {Cancellaria) Libassi, 1859:40, fig. 25; as
var. of C. contorta Basterot. Tertiary, Italy.
alternata {Cancellaria) Conrad, 1834:155. Miocene,
Maryland, U.S.A.
alternicostula {Cancellaria) Sacco, 1894:19, pi. 1, fig.
53; see Appendix, Note 2. Miocene, Italy.
altispira {Cancellaria) Gabb, 1869:50, pi. 14, fig. 7.
Pliocene, California, U.S.A.
alumensis {Cancellaria) Mansfield, 1930:49, pi. 4, fig.
5; as subsp. of C. perspectiva Conrad. Miocene,
Florida, U.S.A.
alvaniopsis {Cancellaria) 'Harris' - Cossmann, 1899a:34.
Nomen nudum.
alveata {Cancellaria) Conrad, 1833:45; 1835:44, pi. 16,
fig. 19. Eocene, Alabama, U.S.A.
alveolata {Cancellaria) Tate, 1889:154, pi. 10, figs.
7a-b. Miocene, Australia.
amakusana {Trigonaphera) Petit, 1974:112, text-fig. 4.
Recent, Japan.
Page 10
THE NAUTILUS, Supplement 1
amasia (Cancellapheia) Iredale, 1930:80, pi. 9, fig. 8.
Recent, Australia.
ambigua {Admete) Hutton, 1885:320, pi. 18, fig. 18.
Pliocene, New Zealand. [Acicon]
ameghinoi (Cancellaria) von Ihering, 1897:310, pi. 3,
fig. 12; pi. 4, fig. 15. Tertiary, Argentina.
amekiensis (Bonellitia) Eames, 1957:49, pi. 7, figs.
5a-c. Eocene, Nigeria.
amoena {Cancellaria) Olsson & Harbison, 1953:179,
pi. 28, fig. 7. Pliocene, Florida, U.S.A.
ampla {Inglisella) Laws, 1935:37, pi. 6, fig. 16. Mio-
cene, New Zealand.
ampleumbilicata {Cancellaria) Sacco, 1894:33, pi. 2,
fig. 42; as var. of C. calcarala (Brocchi); see
Appendix, Note 2. Miocene, Italy.
ampulla (Cancellaria) "Broc.'. Error for C. ampullacea
Brocchi.
ampullacea {Valuta) Brocchi, 1814:313, pi. 3, figs.
9a-b. Pliocene, Italy. [Trigonostoma s.l.]
ampullacera [Cancellaria) Lesson, 1841b:253. Recent,
? New Zealand. Nonien dubium.
anachoreta {Cancellaria) Mayer, 1876:45, pi. 4, fig.
18. Tertiary, Europe.
ancycla {Cancellaria) Gardner, 1937:374, pi. 45, figs.
6-7. Miocene, Florida, U.S.A.
andaluciensis {Trigonostoma) Landau, 1984:151, pi. 2,
figs. 12-17. Pliocene, Spain.
andersoni {Cancellaria) Arnold, 1909:60, pi. 9, fig. 5.
Miocene, California, U.S.A.
andersoni {Cancellaria) B. L. Clark, 1918:80, 82, 97,
pi. 23, fig. 4. Oligocene, California, U.S.A. (?= C.
oregonensis Conrad, 1865)
angasi {Cancellaria) Crosse, 1863:64, pi. 2, fig. 8.
Recent, northwestern Africa.
angasi (Tritonium) Brazier, 1877:174. Recent, Australia.
[Tritonoharpa]
angelana {Cancellaria) G. D. Hanna, 1924:159; n.n. for
C. quadrata Moody, 1916, nan Sowerby, 1822.
Pliocene, California, U.S.A.
anglica {Cancellaria) Sacco, 1894:9; n.n. for "C
umhilicaris (Brocchi)" Wood, I874:Add. pi., fig.
10; as var. of C ampullaccum (Br.); see Appendix,
Note 2. Pliocene, England.
angosturana (Cancellaria) Marks, 1949:463, pi. 78,
figs. 1-2. Miocene, Ecuador.
angulata {Cancellaria) Eichwald, 1830:222. Tertiary,
Europe.
angulata {Cancellaria) 'Watelet' - Jousseaume,
1887b:222. Error for C. angusta Watelet.
angulata {Cancellaria) Sieber, 1936:98; as var. of C.
inermis Pusch. Miocene, Austria.
atigulatina {Cancellaria) Sacco, 1894:40, pi. 2, fig. 64;
as var. of C. dertonensis Bellardi; see Appendix,
Note 2. Miocene, Italy.
angulifera {Cancellaria) Deshayes, 1864:107, pi. 73,
figs. 13-15. Middle Eocene, France.
angulifera {Cancellaria) von Kocncn, 1885:10, pi. 1,
figs. 6a-b. Paleocene, Denmark. (= Narona ravni
Gilbert, 1960)
angulopusilla {Admete) Sacco, 1894:71; n.n. for "'C.
pusilla (Phil.)" Beyrich, 1856:pl. 28, fig. 1; as var.
of A. exilis (Philippi). Tertiary, Germany.
angulosior {Admete) Sacco, 1894:71; n.n. for "C.
subangulosa Wood" Speyer, 1867:pl. 11 [sic; error
for pi. 16], fig. 11; as var. of A. exilis (Philippi).
Tertiary, Germany.
angulovaricosa {Cancellaria) Sacco, 1894:57, pi. 3,
figs. 53a-b; see Appendix, Note 2. Miocene. Italy.
angusta {Cancellaria) Watelet, 1851:124, pi. 2, figs.
3-4. Lower Eocene, France.
angusta {Cancellaria) Almera & Bofill, 1884:46. pi.
D, figs. 21-22; as var. of C. lyrata (Brocchi).
Tertiary, Spain.
annosa {Cancellaria) Aldrich, 1898:97. Eocene, Ala-
bama, U.S.A.
annulata {Cancellaria) Stoliczka, 1867:162, pi. 13, fig.
11. Cretaceous, India.
anodosomagna {Brocchinia) Sacco, 1894:69, pi. 3, fig.
84; as var. of B. milraeformis (Brocchi). Pliocene,
Italy.
anomala {Admete) Marshall & Murdoch. 1920:132, pi.
6, fig. 6. Eocene, New Zealand.
anomoia {Cancellaria) Woodring, 1970:334, pi. 52,
figs. 1-2. Miocene, Panama.
ansonae {Tritonoharpa) Beu & Maxwell, 1987:35, pis.
13, figs, e-i; pi. 14, figs. h-k. m-p. Recent, western
Australia.
antarctica {Admete) Strebel, 1908:21, pi. 4, figs. 44a-c.
Recent, Antarctic.
antiqua {Cancellaria) 'Wagner' - Bronn, 1848:208.
Nomen nudum.
antiqua {Cancellaria) Dall, 1897:11, pi. 3, fig. 3; ex
Wagner MS. Tertiary, southeastern U.S.A.
antiquata {Cancellaria) Hinds, 1843:49; 1844b:43, pi.
12, figs. 17-18. Recent, Indo-Pacific.
antiquata {Cancellaria) 'Edwards MS' - Wrigley,
1935:372; in synonymy of Bonellitia clarendonen-
sis Wrigley. Nomen nudum.
antiquatus {Triton) Hinds in Reeve, 1844:pl. 18, fig.
80. Recent, Indo-Pacific. [Tritonoharpa]
anxifer {Cancellaria) Iredale, 1925:264; as subsp. of C.
purpuriformis Kuster [sic]. Recent, Australia.
aoteana {Antizafra) Dell, 1956:111, pi. 11, fig. 110.
Recent, New Zealand. [?= Dcllina]
apater {Cancellaria) Ryckholt, 1861:pl. 30, figs. 24-25.
Cretaceous, Belgium.
apenninensis {Cancellaria) Sacco, 1894:54, pi. 3, fig.
41; as var. of C. varicosa (Br.); see Appendix, Note
2. Tertiary, Italy.
aperta {Cancellaria) Beyrich, 1856:586, pi. 28, figs.
5a-d. Miocene, Germany.
aphrogenia {Coluhraria) Pilsbry & Lowe, 1932:62, pi.
4. fig. 10. Recent, Panamic-Pacific. [Tritonoharpa]
apimela {Cancellaria) Woodring, 1970:337, pi. 52, figs.
5-6. Miocene, Panama.
aqualica {Cancellaria) Petit & Harasewych, 1986:440,
figs. 7-8. Recent. Philippines.
araeostyla {Oamaruia) Maxwell, 1988:71, pi. II, figs.
d, h, 1. Miocene, New Zealand.
R. E. Petit and M. G. Harasewych, 1990
Paae 11
aquilara (Bucciiium) Watson, 1882b:359. [a turrid;
incorrectly placed in Cancellariidae by several
authors]
arctica (Cancellaria) Middendorff, 1849:112, pi. 9, figs.
11-12, 15. Recent, Bering Strait.
arenaria (Sveltella) Wrigley, 1935:363, pi. 32, figs.
10-11. Paleocene, England.
arnoldi {Cancellaria} Dall, 1909a:29, pi. 14, fig. 7.
Pliocene, California, U.S.A.
articiilaris (Cancellaria) G. B. Sowerby 1, 1832b:fig.
32. Recent, Sri Lanka.
ashen (Marianarona) Petuch, 1988:27, pi. 5, figs. 9-10.
Miocene, Maryland, U.S.A.
aspera {Cancellaria) Millet de la Turtaudiere, 1866:10.
Tertiary, France.
aspera {Cancellaria) "Edwards MS' - Newton, 1891:169.
Nomen nudum.
aspercella {Cancellaria) "Lamarck'. Error for asperella
Lamarck.
asperella {Cancellaria) Lamarck, 1822b: 112. Recent,
Indo-Pacific.
asperula {Cancellaria) Deshayes, 1830:187. Recent,
Indo-Pacific. (?= C. ohliquaia Lamarck, 1822)
asprella {Merica) 'Lamarck' - Habe, 1961b:434; et al.
Error for C. asperella Lamarck.
assimile {Cancellaria) 'Sowerby'. Error for similis Sow-
erby.
assimilis {Cancellaria) 'Sowerby'. Error for similis
Sowerby.
astensis {Cancellaria) Bellardi, 1840:344; as var. of C.
cancel lata Lam. [sic]. Nomen nudum.
astensis {Cancellaria) Bellardi, 1841:28, pi. 3, figs.
17-18; as var. of C. cancellata Lam. [sic]. Tertiary,
Italy.
atjehense {Trigonostoma) Oostingh, 1938:109, pi. 6,
figs. 118-120. Pliocene, Indonesia.
atopodonta {Cancellaria) Petit & Harasewych, 1986:440,
figs. 5-6, 15-16. Recent, Philippines.
atraktoides {Cancellaria) Gardner, 1937:376, pi. 45,
figs. 13-14. Miocene, Florida, U.S.A.
atiirensis {Sveltia) Peyrot, 1928:228, pi. 13, figs. 46-47.
Miocene, France.
aturensis {Tritonidea) Peyrot, 1926:242, pi. 4, figs.
50-52. Miocene, France. [Loxotaphrus]
aupouria {Zeadmete) Powell, 1940:243, pi. 29, fig. 13.
Recent, New Zealand.
auriculaperta {Cancellaria) Yokes, 1938:22, figs. 19-
20. Pliocene, Trinidad.
auriculoides {Cancellaria) Millet de la Turtaudiere,
1854:160. Tertiary, France.
aurorae {Trigonostoma) Palmer, 1937:445, pi. 73, figs.
14-16. Eocene, South Carolina, U.S.A.
australis {Cancellaria) G. B. Sowerby I, 1832b:fig. 23.
Recent, Australia.
australis {Cancellaria) Philippi, 1855:208; 1856a:164;
1856b:99. Recent, Strait of Magellan. (= Admete
philippi von Ihering. 1907).
austriaca {Cancellaria) Hoemes & Auinger, 1890:275,
pi. 33, figs. 15a-c. Miocene, Austria.
avara (Columbella) Say, 1822. [incorrectly placed in
Cancellaria by several authors]
aveniformis {Cancellaria) 'Edwards MS' - Newton,
1891:170. Nomen nudum.
awakinoensis {Inglisella) Laws, 1935:38, pi. 6, fig. 18.
Miocene, New Zealand.
azorica {Admete) Bouchet & Waren, 1985:260, fig. 690.
Recent, Azores.
aziimai {Cancellaria) 'Habe MS" - Azuma, 1960:48.
Nomen nudum.
azumai {Fusiaphera) Habe, 1961a:72, Appendix p. 28,
pi. 35, fig. 20. Recent, Japan.
B
babylonica {Cancellaria) I. Lea, 1833:138, pi. 5, fig.
134. Eocene, Alabama, U.S.A.
badensis {Cancellaria) Sacco, 1894:60; n.n. for "C.
lyrata (Brocchi)" Homes, 1854:pl. 34, fig. 4; as
var. of C. lyrata (Brocchi); see Appendix, Note 2.
Miocene, Austria.
badrii {Cancellaria) Abbass, 1972:59, pi. 4, fig. 14.
Paleocene, Saudi Arabia.
bahia {Cancellaria) Pilsbry & Olsson, 1941:24, pi. 3,
fig. 3. Pliocene, Ecuador.
balboae {Cancellaria) Pilsbry, 1931:439, pi. 41, figs.
7-8. Recent, Panama Bay.
bantamense {Trigonostoma) Oostingh, 1938:110, pi. 6,
figs. 122a-c. Pliocene, Indonesia.
barjonae {Cancellaria) Pereira Da Costa, 1867:201, pi.
25, figs. 9-13. Miocene, Portugal.
barkeri {Fossarus) P. M. Anderson in G. D. Hanna,
1924:165; unnecessary n.n. for Fossarus dalli An-
derson & Martin, 1914. [Cancellariidae]
barkeri {Zeadmete) Powell, 1952:184, pi. 36, fig. 1.
Recent, New Zealand.
barnardi {Trigonostoma) Janssen, 1984:19, pi. 3, figs.
9-14; pi. 6, fig. 3. Miocene, Netherlands.
barretti {Cancellaria) Guppy, 1866:289, pi. 17, fig. 11.
Pliocene, Jamaica.
bartonensis {Bonellitia) Wrigley, 1935:368, pi. 33, fig.
16. Eocene, England.
barystoma {Cancellaria) Woodring, 1970:342, pi. 53,
figs. 5-6. Miocene, Panama.
basicosticillata {Cancellaria) Sacco, 1894:34, pi. 2, fig.
46; as var. of C. calcarata (Brocchi); see Appendix,
Note 2. Pliocene, Italy.
basilaevis {Tritonoharpa) Beu & Maxwell, 1987:37, pi.
16, figs, a-i, k-1. Recent, western Pacific.
basisulcata {Sveltia) Peyrot, 1928:237, pi. 12, figs.
39-40; as "mut." of S. calcarata (Brocchi). Mio-
cene, France.
basteroti {Cancellaria) Deshayes, 1864:104; n.n. for
"C. buccinula Lam." Basterot, 1825. Miocene,
France.
bastropensis {Cancellaria) G. D. Harris, 1895a:66, pi.
6, fig. 5. Eocene, Texas, U.S.A.
bathyalis {Palaeadmete [sic]) Okutani, 1964:398, pi. 6,
fig. 7. Recent, Japan. [? Iphinopsis]
battersbyi {Cancellaria) Bell, 1870b:345; n.n. for '"C.
hirta (Brocc.)" Grateloup, 1847:pl. 25, fig. 25.
Tertiary, France. (= C. subhirta d'Orbigny, 1852)
Page 12
THE NAUTILUS, Supplement 1
bayeri (Adnietula) Petit, 1976:38, pi. 1, fig. 4. Recent,
Gulf of Mexico.
baylei (Cancellaria) Bezan9on, 1870:316, pi. 10, fig. 3.
Tcrtiar)'. France.
bearnensis (Cancellaria) Peyrot, 1928:206, pi. 12, fig.
1; as var. of C. harjonae Pereira da Costa. Miocene,
France.
bearnensis (.Sveltia) Peyrot, 1928:219, pi. 13, fig. 8; as
■"niut." of S. varicosa (Brocchi). Miocene, France.
beata (Cancellaria) Jung, 1965:554, pi. 75, figs. 12-14.
Miocene, Venezuela.
beatrix (Admete) Olsson, 1964:128, pi. 22, fig. 9.
Pliocene, Ecuador.
beatum {Trigonosioma) Pilsbry & Harbison, 1933:108,
pi. 3, fig. 1. Miocene, New Jersey, U.S.A.
behmi (Cancellaria) Beyrich, 1856:584, pi. 28, figs.
6a-b. Oligocene, Germany.
bella (Cancellaria) Aldrich, 1898:98; as var. of C.
i^raciloides Aldrich. Eocene, Alabama, U.S.A.
bellardii (Cancellaria) Michelotti, 1846:55. Nomen nu-
dum.
bellardii (Cancellaria) Michelotti, 1847:225. Miocene,
Italy.
bellardii (Trigonostoma) de Stefani & Pantanelli,
1878:1 16. Pliocene, Italy. (= C. exumbilicaris Sacco,
q.v.)
bellardii (Trigonostoma) Venzo & Pelosio, 1963:111,
pl. 39, figs. 3, 3a-c, 4; n.n. for "C. ampullacea
(Br.)" Bellardi, 1841:pl. 4, figs. 7-8; as subsp. of
T. ampullacea (Brocchi). Tertiary, Italy.
bellsana (Caveola) Stephenson, 1953:190, pl. 42, figs.
26-28. Cretaceous, Texas, U.S.A. [? Fasciolariidae]
benedeni (Valuta) Miiller, 1851:41, pl. 6, figs. 5a-b.
Cretaceous, Germany. [Cancellariidae]
benoisti (Trigonostoma) Peyrot, 1928:245, pi. 14, fig.
31. Miocene, France.
beraudiana (Cancellaria) Millet de la Turtaudiere,
1854:160. Nomen nudum.
beraudiana (Cancellaria) Millet de la Turtaudiere,
1866:10. Tertiary, France.
bernardii (Cancellaria) Mayer, 1861a:371, pl. 15, figs.
3-4. Tertiary, France.
bernayi (Cancellaria) Cossmann, 1889:230, pl. 7, fig.
29. Middle Eocene, France.
berolinensis (Cancellaria) Beyrich, 1847:162. Nomen
nudum.
betsiae (Trigonostoma) Olsson & Petit, 1964:544, pl.
80, figs. 1, la. Pliocene, South Carolina, U.S.A.
bettina (Cuma) Semper. 1862:102. Oligocene, Germany.
1?= Turehua tuherculata (Giebel, 1861)]
beyrichi (Cancellaria) Mayer, 1859:391, pl. 11. fig. 8.
Tertiary, Europe.
bezanconi (Cancellaria) de Raincourt, 1884:345, pl. 12,
fig. 10. Upper Eocene, France.
bezanconi (Cancellaria) Modet. 1885a:49, pl. 3, figs.
3, 3a. Eocene, France. (= C. muttienensis Morlet,
1885b).
bezanconi (Cancellaria) 'de Boury" - Meyer, 1886:74.
[Error for Scalaria bezanconi de Boun,]
bicarinata (Cancellaria) Hoemes & Auinger, 1890:281,
pl. 33, figs. 16a-c. Miocene, Austria.
bicincta (Admete) 'von Koenen' - Chavan, 1947:142.
Error for Cancellaria tricincta von Koenen.
bicolor (Cancellaria) Hinds, 1843:48; 1844b:43, pl. 12,
figs. 13-14. Recent, Indo-Pacific.
bicosticillata (Cancellaria) Sacco, 1894:56, pl. 3, fig.
45; as var. of C. varicosa (Br.); see Appendix, Note
2. Pliocene, Italy.
bicosticillata (Cancellaria) Sacco, 1894:35, pl. 2, fig.
50; as var. of C uniangulata Deshayes; see Appen-
dix, Note 2. Pliocene, Italy.
bifasciata (Cancellaria) Deshayes, 1830:181. Recent,
Indo-Pacific. (?= C. oblonga Sowerby, 1825)
bifoliata (Cancellaria) Aldrich, 1903:101, pl. 4, fig. 24.
Miocene, Florida, U.S.A.
bifurcoplicata (Cancellaria) Cossmann, 1896b:213, pl.
6, fig. 30. Middle Eocene, France.
bilineata (Bonellitia) Wrigley, 1935:367, pl. 33, fig. 15.
Middle Eocene, England.
binckhorsti (Cancellaria) Nyst, 1881:8; n.n. for C.
reticulata Binkhorst, 1861, non Linne, 1767. Creta-
ceous, Belgium.
binkhorsti (Cancellaria) Cossmann, 1899c: 178; n.n. for
C. reticulata Binkhorst, 1861, non Linne, 1767.
Cretaceous, Belgium. (= C. binckhorsti Nyst, 1881)
biperspinosa (Cancellaria) Sacco, 1894:61, pl. 3, fig.
61; as var. of C. lyrata (Br.); see Appendix, Note
2. Pliocene, Italy.
biplex (Cancellaria) G. B. Sowerby I, 1822:fig. 4. Plate
legend only; = C. suturalis G. B. Sowerby I, 1822.
biplicata (Cancellaria) Brian & Comet, 1870:12, pl. 1,
fig. 7. Paleocene, Belgium.
biplicatum (Tritonium) Ravn, 1902:228, pl. 2, figs. 11,
12, 13a-b. Tertiary, Europe. [? Admete]
biplicatus (Fusus) Lamarck, 1803:388. Eocene, France.
[?= Admetida evulsa (Solander, 1766)]
biplicifera (Cancellaria) Conrad, 1841:31. Miocene,
Mar>'land, U.S.A.
birciti (Cancellaria) Addicott, 1970:118, pl. 16, figs.
12-16. Miocene, California, U.S.A.
birmanica (Cancellaria) Vredenburg, 1921:140. pl. 15,
figs. lOa-b. Tertiary, Burma.
birmanicum (Trigonostoma) Vredenburg, 1921:141. pl.
15, figs, lla-b. Tertiary, Burma.
bistriata (Cancellaria) von Koenen, 1889:116, pl. 8,
figs. 5a-d. Oligocene, Germany.
bivet (Bivetia) Jousseaume, 1887b: 193; e.\ Adanson.
Recent, western Africa. (= Cancellaria similis Sow-
erby, 1833)
blountiana (Cancellaria) Mansfield, 1935:27, pi. 2, fig.
4. Miocene, Florida, U.S.A.
bocageana (Cancellaria) Crosse, 1863:63. Nomen nu-
dum.
bocageana (Cancellaria) Crosse & Debeaux, 1863a:77;
1863b:263, pl. 9, fig. 3. Recent, China.
R. E. Petit and M. G. Harasewych, 1990
Page 13
bocagei (Cancellaria) Crosse & Debeaux - Dautzenberg
& Fischer. 1907:149. Emendation of hocageaim.
boettgeri {Pseudomalaxis) Cossmann, 1915:143, pi. 12,
figs. 22-24; n.n. for DiscohelLx calculiformis Boettger,
1906, non Dunker, 1847. Miocene, Romania.
[Trigotwstonui s.L]
bofilli (Cancellaria) Cossmann, 1899b: 102; n.n. for C.
piilchenima Almera & Bofill, 1898, noii Lea, 1841.
Pliocene, Spain.
boivini (Cancellaria) Dareste de la Chavanne, 1910:23,
pi. 4, fig. 7. Eocene, Algeria.
bonacorsi (Cancellaria) 'Doderlein MS' - Davoli,
1982:69. Nomen nudum,
bonellii (Cancellaria) Bellardi, 1840:344. Nomen nu-
dum.
bonellii (Cancellaria) Bellardi, 1841:24, pi. 3, figs. 3-4.
Miocene, Italy.
bonneti (Sveltella) Cossmann, 1902:56, pi. 4, fig. 4.
Middle Eocene, France.
bonneti (Trigonostoma) Cossmann, 1903a: 110, pi. 3,
figs. 10-11. Pliocene, India.
borealis (Admete) A. Adams, 1855:122. Recent, Arctic
Ocean. (?= A. viridula (Fabricius, 1780))
boreobsoleta (Trigonostoma) Kautsky, 1925:142, pi. 10,
figs. 12-13. Miocene, Germany.
boucheti (Cancellaria) Petit & Harasewych, 1986:436,
figs. 1-4, 9-14. Recent, Philippines.
boucheti (Tritonoharpa) Beu & Maxwell, 1987:37, pi.
6, figs.i-o; pi. 7, figs. a-h. Recent, Mozambique.
bouczeki (Turbinella) Rzehak, 1896:250; in synonymy
of Cancellaria bouczeki Oppenheim, q.v. Nomen
nudum.
bouczeki (Cancellaria) Oppenheim, 1922:81, pi. 5, figs.
14, 14a-b; ex Rzehak list. Tertiary, Czechoslovakia.
bourdoti (U.xia) Cossmann & Pissarro, 1901:20, pi. 8,
fig. 6. Middle Eocene, France.
bournei (Cancellaria) M. A. Hanna, 1927:323, pi. 55,
figs. 1, 3, 4, 6. Eocene, California, U.S.A.
boutiiiieri (Cancellaria) Cossmann, 1889:223, pi. 7, fig.
28. Middle Eocene, France.
bradleyi (Cancellaria) Nelson, 1870:192, pi. 6, figs.
8-9. Miocene, Peru.
brandenburgi (Cancellaria) Boettger, 1902:38. Mio-
cene, Romania.
brauneana - error for brauniana.
brauniana (Cancellaria) "Nyst' - Braun, 1851:1131; et
id. Nomen nudum.
brauniana (Cancellaria) Sandberger, 1859:pl. 15, figs.
7, 7a; 1862:258; e.\ Braun; ex Nyst //( lilt. Tertiary,
Germany.
breve (Coptostoma) Wrigley, 1935:359, pi. 32, fig. 4.
Middle Eocene, England.
breviplicata (Voliita) Forbes. 1846:132, pi. 12, figs.
7a-b. Cretaceous, India, [placed in Cancellaria by
Stoliczka, 1867:163; not a cancellariid]
brevirostris (Turbinella) Hutton, 1877:596, pi. 16, fig.
10. Tertiary, New Zealand. [Maorivetia]
brevis (Cancellaria) G. B. Sowerby I, 1832a:52;
1832b:fig. 33. Recent, Panamic-Pacific.
brocchii (Cancellaria) Crosse, 1861:248; n.n. for " V.
piscatoria L." Brocchi, 1814:308, pi. 3, fig. 12.
Pliocene, Italy. (= C. hirta minor Bronn, 1848)
bronnii (Cancellaria) Bellardi, 1840:344. Nomen nu-
dum.
bronnii (Cancellaria) Bellardi, 1841:31, pi. 4, figs.
11-12. Miocene, Italy.
bruecknerii (Tritonium) Boll, 1846:162, pi. 2, fig. 9;
as briicknerii. Oligocene, Germany. [?= Admetula
evulsa (Solander, 1766)]
brunnea (Tritonoharpa) Beu & Maxwell, 1987:38, pi.
15, figs, a-e, g-h. Recent, Indonesia.
bruuni (Admete) Knudsen, 1964:132, figs. 9-10. Recent,
Kermadec Trench.
buccinea (Cancellaria) 'Basterot" - Bronn, 1848:209.
Error for C. buccinula Lamarck.
buccinoides (Cancellaria) G. B. Sowerby I, 1832a:54;
1832b:fig. 11. Recent, Panamic-Pacific.
buccinoides (Cancellaria) Couthouy, 1838:105, pi. 3,
fig. 3. Recent, North Atlantic. (= C. couthouxi Jay,
1839).
buccinoides (Cancellaria) von Koenen, 1889:106, pi.
10, figs. 9a-b, lOa-c. Oligocene, Germany.
buccinoides (Triton) K. Martin, 1879:60, pi. 14, fig. 11.
Tertiary, Indonesia. [= Cancellaria neglecta K.
Martin, 1895; = Bivetia martini Cossmann, 1899]
buccinovula (Cancellaria) Sacco, 1894:45; n.n. for "C.
evulsa (Sol.)" Speyer, 1867:pl. 11 [sic; error for
16], fig. 2; as var. of C. evulsa (Sol.); see
Appendix, Note 2. Tertiary, Germany.
buccinula (Cancellaria) Lamarck, 1 822b: 117. Tertiary,
France. (?= Admetula evulsa (Solander, 1766))
bulbrooki (Cancellaria) "Mansfield' - Marks, 1949:459.
Error for C. bullbrooki Mansfield.
bulbulus (Cancellaria) G. B. Sowerby I, I832a:55.
Recent, Panamic-Pacific.
bullata (Cancellaria) G. B. Sowerby I, 1832a:51;
I832b:fig. 35. Recent, Panamic-Pacific.
bullbrooki (Cancellaria) Mansfield, 1925:31, pi. 5, fig.
3. Miocene, Trinidad.
burdigalense (Cancellaria) Peyrot, 1928:253, pi. 14,
figs. 29, 34-35. Miocene, France.
burdigalensis (Sveltia) Peyrot, 1928:221, pi. 14, figs.
1-2. Miocene, France.
C
cacellensis (Cancellaria) Pereira Da Costa, 1867:209,
pi. 26, figs. 3a-b. Miocene, Portugal.
cacellotransiens (Cancellaria) Sacco, 1894:52; n.n. for
"C. dufourii Grat." Pereira Da Costa, I867:pl. 24,
fig. 7; as var. of C. callosa Partsch; see Appendix,
Note 2. Miocene, Portugal.
cacellowestiana (Cancellaria) Sacco, 1894:26; n.n. for
"C westiana Gr." Pereira Da Costa, 1867:pl. 25,
figs. 3-5; as var. of C. mutinensis Foresti; see
Appendix, Note 2. Miocene, Portugal.
Calais (Trigonostoma) Kautsky, 1925:140, pi. 10, figs.
7-8. Miocene, Germany.
calcarata (Valuta) Brocchi, 1814:309, pi. 3, fig. 7.
Pliocene, Italy. [Calcarata]
Page 14
THE NAUTILUS, Supplement 1
calculiformis (Discohelix) Boettger, 1906:138. [= Pseu-
domala.xis boettgeri Cossmann, 1915, q.v.]
californica {Cancellaria) Dall, 1908:296, pi. 4, fig. 4.
Recent, California, U.S.A.
callosa {Cancellaria) 'Partsch' - M. Homes, 1848:20.
Nomen nudum.
callosa {Cancellaria) M. Homes, 1854:314, pi. 34, figs.
14-16; ex Pansch MS. Miocene, Austria.
caloosahatchiensis {Cancellaria) Tucker & Wilson,
1932:7. pi. 1, fig. 4. Pliocene, Florida, U.S.A.
calvatula {Cancellaria) 'Sowerby' - Grant & Gale,
1931:942. Plate caption error for C. clavaiula
Sowerby.
calvertensis {Cancellaria) G. C. Martin, 1904:167, pi.
43, fig. 12. Miocene, Maryland, U.S.A.
calvulata {Cancellaria) Tate, 1889:153, pi. 9, fig. 3.
Miocene, Australia.
calypso {Cancellaria) White, 1887:124, pi. 10, figs.
23-24. Cretaceous. Brazil.
camdeo {Valuta) Forbes, 1846:131, pi. 12, figs. 5a-b.
Cretaceous, India. [? Cancellariidae]
camdio {Valuta) Forbes, 1846:pl. 12, figs. 5a-b; plate
legend error for V. camdea.
campbelli {Trigonostoma) Shasky, 1961:20, pi. 4, fig.
5. Recent, western Mexico.
canaliculata {Cancellaria) M. Homes, 1854:324, pi. 35,
figs. 9a-c, lOa-b. Miocene, Austria.
canaliculata {Cancellaria) Deshayes, 1864:97, pi. 72,
figs. 26-28. Eocene, France. (= C. rhabdota Bayan,
1873)
canaliculata {Narona) Janssen, 1972:40, pi. 7, figs. 8-9.
Miocene, Netherlands.
cancellangulosa {Admete) Sacco, 1894:71; n.n. for "C.
subangulosa Wood" Speyer, 1867:pl. 11 [sic; error
for 16], fig. 10; as var. of A. minuta Braun.
Tertiary, Germany.
cancellaria {Trichatrapis) Conrad, 1858:333, pi. 35, fig.
8. Cretaceous, Mississippi, U.S.A. [Paladmete]
cancellata {Valuta) Linne, 1767:1191; n.n. for Murex
scabriculus Linne, 1758, nan Valuta scabriculus
Linne, 1758. Recent, northwestern Africa. [Cancel-
laria]
cancellata {Admete) Kobelt, 1887a:12; 1887b:105, pi.
24, fig. 14. Recent, Japan.
cancellata {Lara) Otuka, 1937:1020; n.n. for "Admete
viridula Fabricius" Yokoyama, 1920:45, pi. 2, fig.
5. Pliocene, Japan. [- Admete yokoyamai Oyama,
1954]
cancellatina {Cancellaria) Sacco, 1894:45; n.n. for "C.
evulsa (Sol.)" Speyer, 1867:pl. 11 [sic; error for
16], fig. 4; as var. of C. evulsa (Solander); see
Appendix, Note 2. Tertiary, Germany.
cancellatina {Cancellaria) Sacco, 1894:12, pi. 1, fig.
30; as var. of C. tauralaevigatum Sacco; see
Appendix, Note 2. Miocene, Italy.
cancellatula {Cancellaria) Sacco, 1894:43, pi. 3, fig. 4;
as var. of C. bonellii (Bellardi); see Appendix, Note
2. Miocene, Italy.
cancellatum {Pleurotoma) Eichwald, 1830:225. Tertiary,
U.S.S.R. (= Cancellaria notahilis Eichwald, 1851]
cancellosa {Oamaruia) Marwick, 1965:40, pi. 11, figs.
6, 7. Tertiary, New Zealand.
candeana (Cancellaria) d'Orbigny, 1842:pl. 21, figs.
23-25. Recent, Caribbean, (see C. candei d'Or-
bigny) [Antillophos]
candei (Cancellaria) d'Orbigny, 1853:129. Error for, or
emendation of, C candeana d'Orbigny. Recent,
Caribbean. [Antillaphas]
Candida (Cancellaria) G. B. Sowerby I, 1832b:fig. 1.
Recent,? Panamic-Pacific.
caniilata (Buccinella) Perry, 181I:pl. 27, fig. 1. Recent,
locality unknown. [?= Cancellaria reticulata (Linne,
1767)]
caperata {Cancellaria) Tate, 1889:158, pi. 9, fig. 7.
Miocene. Australia.
capillata (Cancellaria) Tate, 1889:158, pi. 10, fig. 10.
Miocene, Australia.
carinapex (Africatriton) Beu & Maxwell, 1987:30, pi.
6, figs. a-h. Recent, Australia.
carinata (Cancellaria) Brian & Comet, 1877:14, pi. 14,
figs. 5a-c. Paleocene, Belgium.
carinata (Cancellaria) Watson, 1882a:327; 1886:275,
pi. 18, fig. 9. Recent, Kerguelen Islands. (= Zead-
mete watsoni Petit, 1970)
carinata (Sveltella) Wrigley, 1935:362, pi. 32, fig. 8.
Eocene, England.
carinatum (Coptastama) Wrigley, 1935:359, pi. 32, fig.
3; as form of C. quadratum (Sowerby). Eocene,
England.
carolinensis {Cancellaria) Emmons, 1858:254, fig. 118.
Tertiary, North Carolina, U.S.A.
carolinensis (Cancellaria) Conrad, 1863:567; n.n. for
"C. reticulata Lam." Emmons, 1858:255, fig. 119.
Tertiary, North Carolina, U.S.A.
casicalva (Cancellaria) Marks, 1949:464, pi. 78, figs.
3, 10. Miocene, Ecuador.
cassidea (Valuta) Brocchi. 1814:314, pi. 3, figs. 13a-b.
Pliocene, Italy. [Triganastama .v./.]
cassidiformis (Cancellaria) G. B. Sowerby I, 1832a:53;
1832b: fig. 22. Recent, Panamic-Pacific.
cassiniana (Cancellaria) 'Edwards MS' - Newton,
1891:170. Nomen nudum.
cassiniana (Bonellitia) Wrigley, 1935:372, pi. 33, fig.
26. Lower Eocene, England.
castexi {Sveltia) Peyrot, 1928:234, pi. 13, figs. 55-56.
Miocene, France.
cathalai (Pisanella) Doncieux, 1908:73, pi. 4, figs.
7a-b. Eocene, France. [Cancellariidae]
caudatior (Admete) Sacco, 1894:73; n.n. for C. dregeri
Hoemes & Auinger, 1890, pi. 33, fig. 19 (only);
as var. of A. nassiformis (Seguenza). Miocene,
Austria.
caveola (Paladmete) Stephenson, 1947:184. pi. 33, figs.
33-36. Cretaceous, Texas, U.S.A.
ceden (Tritanium) 'Gardner' - Sohl. 1964a:269. Error
for T. cedri Gardner.
cedri (Tritanium) Gardner, 1935:262, pi. 21, figs. 9-10.
Paleocene, Texas, U.S.A. [Plesiotritan]
centrota (Cancellaria) Dall, 1896:13; 1908:295. pi. 1,
fig. 8. Recent, Panamic-Pacific.
R. E. Petit and M. G. Harasewych, 1990
Page 15
cerithea (Cancellaria) Olsson. 1964:125, pi. 22, figs.
4, 4a. Tertiary, Ecuador.
cerithiopsis {Cancellaria) Almera & Bofill, 1898:18, pi.
11, figs. 22, 22a. Pliocene, Spain.
chainei (Admete) Peyrot, 1928:262, pi. 13, figs. 19-20.
Miocene, France.
chalmasi (Triton) Quaas, 1902:272, pi. 32, figs. 44-46.
Cretaceous, Libya. [? Plesiotriton]
chalmsi (Triton) 'Quass [sicY — Beu & Maxwell,
1987:51. Error for T. chalmasi Quaas.
charapota (Cancellaria) Olsson, 1942:60, pi. 8, fig. 3.
Miocene, Ecuador.
charh'sworthii (Cancellaria) Wood, 1872:48, pi. 3, figs.
22a-b. Tertiary, England.
chaussyensis (Cancellaria) Cossinann, 1889:231, pi. 8,
fig. 36. Middle Eocene, France.
chavani (Trigonostoma) Palmer, 1937:442, pi. 70, figs.
4, 10; as var. of T. pulcherrima (Lea). Eocene,
South Carolina, U.S.A.
chinenensis (Cancellaria) MacNeil, 1960:99, pi. 14,
fig. 12. Pliocene, Okinawa.
choshiensis (Iphinoella) Habe, 1958:34, 40, fig. 7.
Recent, Japan.
choshiensis (Admete) 'Shikama MS' - Habe, 1961b:436,
pi. 24, figs. 16, 17. Nomen nudum.
choshiensis (Admete) Shikama, 1962:47, pi. 2, figs.
13a-b, 14a-b. Recent, Japan.
christiei (Trigonostoma) Finlay, 1924b:466, pi. 49, fig.
5. Oligocene, New Zealand.
chrysostoma (Cancellaria) G. B. Sowerby \, 1832a:54;
1833:fig. 39. Recent, Panamic-Pacific.
chui (Trigonostoma) Yen, 1936:246, pi. 22, fig. 58.
Recent, China.
cibarcola (Cancellaria) F. M. Anderson, 1929:116, pi.
14, figs. 1-3. Miocene, Colombia.
ciliata (Cancellaria) Michaelis & Scherk, 1847:115; ex
Kroyer label. Nomen nudum.
ciliata (Cancellaria) 'Kroyer' - Morch, 1869:22; Paetel,
1888:329; et al. Nomen nudum.
cincta (Clathurella) Hutton, 1885:327. Miocene, New
Zealand. [Inglisella]
cingens (Cancellaria) 'Sandberger' - Semper, 1861:250.
Error for C. ringens Sandberger.
cingulata (Cancellaria) Kaunhowen, 1898:105, pi. 13,
figs. 3. 3a-b. Cretaceous, Belgium.
cingulata (Gerdiella) Olsson & Bayer, 1972:879, figs.
7-9. Recent, Caribbean.
circumcarinata (Cancellaria) 'Dall' - Dall. 1905:124.
Error for C. circumcincta Dall.
circumcincta (Cancellaria) Dall, 1873:59, pi. 2, fig. 2.
Recent, off northwestern North America.
circumspinosa (Cancellaria) Addicott, 1970:110, pi. 14,
figs. 1-2, 6-7, 16. Miocene, California. U.S.A.
citharella (Cancellaria) Lamarck, 1822b: 114. [not a
cancellariid]
citharella (Valuta) Brongniart, 1823:64, pi. 6, fig. 9.
Tertiary, Poland. [= Cancellaria puschi Semper,
18611
clarendonensis (Cancellaria) 'Edwards MS' - Newton,
1891:170. Nomen nudum.
clarendonensis (Bonellitia) Wrigley, 1935:372, pi. 34,
fig. 39. Lower Eocene, England.
clarki (Perplicaria) M. Smith, 1947:55, pi. 2, fig. 9.
Recent, Panamic-Pacific.
clathrata (Cancellaria) Lamarck, 1822b: 11 6. Tertiary,
France.
clathrata (Cancellaria) A. Adams, 1855:123. Recent,
Panamic-Pacific. (?= C. jayana Keen, 1958)
clatskaniensis (Admete) Anderson & Martin, 1914:88,
pi. 8, figs. 3a-b. Tertiary, Oregon, U.S.A.
clavatula (Cancellaria) G. B. Sowerby I, 1832a:52;
1832b:fig. 12. Recent, Panamic-Pacific.
clenchi (Brocchinia) Petit, 1986:24, figs. 1-4. Recent,
eastern Atlantic.
clewistonensis (Cancellaria) Olsson & Harbison,
1953:178, pi. 28, fig. 2; as subsp. of C conradiana
Dall. Pliocene, Horida, U.S.A.
cloezi (Cancellaria) Cossmann, 1892:74, pi. 3, fig. 16.
Paleocene, France.
coaetana (Cancellaria) Ryckholt, 1862:pl. 33, fig. 1.
Cretaceous, Belgium.
coaeva (Cancellaria) Ryckholt, 1862:pl. 33, figs. 3-4.
Cretaceous, Belgium.
coctilis (Cancellaria) Reeve, 1856:pl. 17, fig. 79. Re-
cent, Indo-Pacific.
codazzii (Cancellaria) F. M. Anderson, 1929:116, pi.
14, figs. 4-7. Miocene, Colombia.
coensis (Cancellaria) Mansfield, 1930:49, pi. 3, figs.
3-4; as coensis. Miocene, Florida, U.S.A.
collectiva (Egerea) Gabor, 1936:7, pi. 1, fig. 13.
Oligocene, Hungary.
colligens (Cancellaria) Sacco, 1894:22, n.n. for "C
inermis Pusch" Homes, 1854:pl. 34, figs. 11-13;
as var. of C inermis Pusch; see Appendix, Note 2.
Miocene, Austria.
colligens (Cancellaria) Sacco, 1894:30, pi. 2, fig. 31;
as var. of C. hirta (Brocchi); see Appendix, Note
2. Pliocene, Italy.
colombiana (Cancellaria) Olsson, 1942:63, pi. 9, fig.
4. Miocene, Colombia.
colpodes (Sveltia) Cossmann, 1899a:21, 192, pi. 2, figs.
18-19. Miocene, France.
cominella (Cancellaria) Pilsbry & Olsson, 1941:23, pi.
3, fig. 7. Pliocene, Ecuador.
complicata (Cancellaria) Handmann, 1882:264. Mio-
cene, Austria.
compressa (Cancellaria) Sacco, 1894:25; n.n. for "C
westiana Grat." Homes, 1854:pl. 35, fig. 13; as
var. of C. e.xwestiana Sacco; see Appendix, Note
2. Miocene, Austria.
compressohirta (Cancellaria) Sacco, 1894:23, pi. 2, fig.
11; as var. of C. geslini Basterot; see Appendix,
Note 2. Miocene, Italy.
compressospira (Cancellaria) Sacco, 1894:40, pi. 2, fig.
63; as var. of C dertonensis (Bellardi); see Appen-
dix, Note 2. Miocene, Italy.
Page 16
THE NAUTILUS, Supplement 1
compressula (Cancellaria) Sacco, 1894:44, pi. 3, fig.
9; as var. of C. serrata Bronn; see Appendix, Note
2. Miocene, Italy.
compressula (.Cancellaria) Sacco, 1894:15, pi. 1, figs.
38a-b; as var. of C. michelinii Bellardi; see Appen-
dix, Note 2. Miocene, Italy.
concava (Oamaruia) Marvvick, 1931:120, pl. 13, fig.
237. Pliocene, New Zealand.
concinna {Cancellaria) Wood, 1842:538. Nomen nu-
dum.
condoni {Cancellaria) F. M. Anderson, 1905:200, pl.
15, figs. 49-50. Miocene, California, U.S.A.
confirmans {Cancellaphera) Ludbrook, 1958:78, pl. 6,
fig. 5. Pliocene, Australia.
conoidea {Cancellaria) von Koenen, 1885:9, pl. 1, figs.
3a-c. Paleocene, Denmark.
conradiana {Cancellaria) Dall, 1889a: 129; 1889b: 104.
Nomen nudum.
conradiana {Cancellaria) Dall, 1890:42, pl. 3, fig. 13.
Tertiary, southeastern U.S.A.
consobrina {Admete) Powell, 1951:167, pl. 10, fig. 63.
Recent, South Georgia Islands.
constantinensis {Uxia) Cossmann & Pissarro, 1901:21,
pl. 8, figs. 7-9. Middle Eocene, France.
contabidata {Cancellaria) G. B. Sowerby I, 1832b:fig.
28. Recent, Indo-Pacific.
contabulata {Admete) Friele, 1879:276. Recent, North
Atlantic. ('?= A. viridula (Fabricius, 1780))
continua {Cancellaria) 'Tate' - Tate & Dennant,
1893:221. Nomen nudum.
contorta {Cancellaria) Basterot, 1825:47, pl. 2, fig. 3.
Tertiary, France.
convexior {Cancellaria) Sacco, 1894:31, pl. 2. fig. 38;
as var. of C. doderleini Mayer; see Appendix, Note
2. Miocene, Italy.
convexior {Cancellaria) Sacco, 1894:17, pl. 1, fig. 44;
as var. of C. subacuminata d'Orbigny; see Appen-
dix, Note 2. Miocene, Italy.
convexosimplex {Cancellaria) Sacco, 1894:55; n.n. for
"C. varicosa (Br.)" Fontannes, 1880:pl. 9, fig. 6;
as var. of C. varicosa (Brocchi); see Appendix,
Note 2. Tertiary, France.
cooperii {Cancellaria) Gabb, 1865:186. Recent, Califor-
nia, U.S.A.
corbicula {Cancellaria) Dall, 1908:294, pl. 1, fig. 4.
Recent, California, U.S.A.
corbula {Cancellaria) Conrad, 1843:308; printed in
some copies as corbulu. Miocene, Maryland, U.S.A.
Species inquirendum.
corbuliformis {Paladmete) Stephenson, 1941:366, pl.
71, figs. 9-11. Cretaceous, Texas, U.S.A.
corbulu {Cancellaria) Conrad, 1843. See corbula.
cordensis {Fasciolaria) Stanton, 1920:43, pl. 8, figs.
8a-b, 9-10. Paleocene, North Dakota, U.S.A. [Cancel-
lariidaej
cornidei {Admete) Altimira, 1978:170, fig. 1. Recent,
northwestern Africa.
cornigera {Cancellaria) Braun, 1851:1131. Nomen nu-
dum.
coronadosensis {Cancellaria) Durham. 1950:102, pl.
26, figs. 2, 8. Pleistocene. Mexico.
coronata {Cancellaria) Hoeninghaus, 1831:145. Nomen
nudum.
coronata {Cancellaria) Scacchi, 1835:5, pl. 1, fig. 15.
Tertiary/Recent, Italy.
corrosa {Cancellaria) Reeve, 1856:pl. 14, fig. 64.
Recent, Panamic-Pacific.
corrugata {Cancellaria) Hinds, 1843:48; 1844b:42, pl.
12, figs. 1-2. Recent, Panamic-Pacific.
corrugata {Valuta) Binkhorst, 1861:14. pl. 5, figs. la-b.
Cretaceous, Belgium. [? CancellariidaeJ
cossnianni {Cancellaria) Morlet, 1888:209, pl. 9, figs.
10, lOa-b. Upper Eocene, France.
cossnianni {Cancellaria) Olsson, 1922:81, pl. 6, figs.
9, 11. Pliocene, Costa Rica. (= C. petiti Olsson,
1967)
cossnianni {Plesiotriton) Oppenheim, 1906:311, pl. 24,
figs. 23a-b. Eocene, Egypt.
costata {Cancellaria) G. B. Sowerby I, 1822:fig. 2.
Recent, northwestern Africa. (?= C. cancellata
(Linne, 1767)).
costata {Cancellaria) G. B. Sowerby I, 1833:fig. 42, ex
Gray MS. Recent, western Africa. (?= Scalptia
scala (Gmelin, 1791))
costata {Cancellaria) I. Lea, 1833:141, pl. 5, fig. 140.
Eocene, Alabama, U.S.A. (?= C. gemmata Conrad.
1833)
costata {Cancellaria) Calcara, 1845:281. Not newly
described; transfer of Buccinuni costulatus Calcara,
1840 to Cancellaria.
costata {Brocchinia) Harmer, 1918:397, pl. 39, fig. 41;
as var. of B. mitraeformis (Brocchi). Pliocene,
England.
costatissima {Cancellaria) Sacco, 1894:15, pl. 1, fig.
41; as var. of C. michelinii Bellardi; see Appendix,
Note 2. Miocene, Italy.
costatonodosa {Cancellaria) Sacco, 1894:47, pl. 3, fig.
19; as var. of C. evulsa (Solander); see Appendix,
Note 2. Oligocene, Italy.
costellata {Cancellaria) 'Nyst' - Gilbert, 1952a: 131.
Error for C. costulata Lamarck.
costellifer {Mure.x) J. Sowerby, 1818:225, pl. 199, fig.
3. Pliocene, England. [?= Admete viridula (Fabri-
cius, 1780)]
costicillatina (Cancellaria) Sacco, 1894:14, pl. 1, fig.
36; as var. of C. scrobiculatum Homes; see Appen-
dix, Note 2. Miocene, Italy.
costicillatissima {Cancellaria) Sacco, 1894:38, pl. 2,
fig. 56; as "anom." of C. cancellata pleuricosticil-
lata Sacco; see Appendix, Note 2. Pliocene. Italy.
costicillatissima {Cancellaria) Sacco, 1894:64, pl. 3,
fig. 73; as var. of C. tribulus (Brocchi); see
Appendix, Note 2. Pliocene, Italy.
costifera (Cancellaria) G. B. Sowerby I, 1832b:fig. 31.
Recent, Indo-Pacific.
costigera (Trigonostoma) 'Sowerby' - Oliver, 1982:16,
19. Error for T. costifera (Sowerby).
costulata (Cancellaria) Lamarck, 1803:63. Middle Eo-
cene, France.
R. E. Petit and M. G. Harasewych, 1990
Page 17
costulatior (Cancellaria) Sacco, 1894:17, pi. 1, fig. 43;
as var. of C. subaciiminatiim d'Orbigny; see Appen-
dix, Note 2. Miocene, Italy.
costulatus {Buccinitm) Calcara, 1840:50. Tertiary, Italy.
[transferred to Cancellaria by Calcara, 1845:281,
under the incorrect spelling costata]
couthonyi {Achnete) 'Jay' - Habe, 1961a:72; 1961b:436.
Error for A. couthonyi (Jay).
couthouyi (Cancellaria) Jay, 1839:77; n.n. for C. buc-
cinoides Couthouy, 1838, non Sowerby, 1832.
Recent, North Atlantic. (?= Admete viridula (Fabri-
cius. 1780))
couturieauxi (Admete) Gilbert, 1938:118, pi. 4, fig. 9,
text-fig. 33. Oligocene, Belgium.
couvana (Cancellaria) Yokes, 1938:20, fig. 21. Mio-
cene, Trinidad.
coxi (Triton) Brazier, 1872:22, pi. 4, fig. 9. Recent,
Australia. [Tritonoharpa]
crassa (Cancellaria) Nomland, 1917:237, pi. 12, figs.
7, 7a. Pliocene, California, U.S.A.
crassa (Cancellaria) Waring, 1917:66, pi. 9, fig. 5.
Cretaceous, California, U.S.A. (= C. simiana Hanna,
1924)
crassevaricosa (Cancellaria) Sacco, 1894:46, pi. 3. fig.
16; as var. of C. eviilsa (Solander); see Appendix,
Note 2. Miocene, Italy.
crassicosta (Cancellaria) Bellardi, 1841:23, pi. 2, figs.
7-8. Miocene, Italy.
crassicostata (Narona) Nordsieck, 1972:88; as form of
N. fusiformis (Cantraine). Miocene, Europe.
crassicosticillata (Admete) Sacco, 1894:72, pi. 3, fig.
96; as var. of A. costellifera [sic] (Sowerby).
Pliocene, Italy.
crassicosticillata (Cancellaria) Sacco, 1894:27, pi. 2,
fig. 21; as var. of C. piscatoria (Gmelin); see
Appendix, Note 2. Pliocene, Italy.
crassitwdosa (Brocchinia) Sacco, 1894:70, pi. 3, figs.
90a-b. Miocene, Italy.
crassispiralis (Turehiia) Beu & Maxwell, 1987:18, pi.
1, figs. k-1. Eocene. New Zealand.
crassistria (Cancellaria) von Koenen, 1889:112, pi. 8,
figs. 6a-c, 7a-d. Oligocene, Germany.
crassistriata (Cancellaria) Wood. 1879:22, pi. 3, figs.
16a-b. Tertiary, England.
crassocostata (Cancellaria) Sacco, 1894:13, pi. 1, fig.
32; as var. of C. imbricatum Homes; see Appendix,
Note 2. Miocene, Italy.
crassonana (Cancellaria) Sacco, 1894:23, pi. 2, fig. 7;
as var. of C. geslini Basterot; see Appendix, Note
2. Miocene, Italy.
crassopostica (Cancellaria) Sacco, 1894:23, pi. 2, fig.
10; as var. of C. geslini Basterot; see Appendix,
Note 2. Miocene. Italy.
crawfordiana (Cancellaria) Dall, 1891:182, pi. 6, fig.
1. Recent, California, U.S.A.
crebriliratus (Epidromus) G. B. Sowerby ID, 1903:220,
pi. 4. fig. 4. Recent, South Africa. [Africoiriton]
cremata (Cancellaria) Hinds. 1843:48; 1844b:42. Re-
cent, Panamic-Pacific.
crenata (Cancellaria) M. Homes, 1856:679, pi. 52, figs.
4a-b. Miocene, Austria.
crenifera (Cancellaria) G. B. Sowerby I, 1832b:fig. 29.
Recent, Indo-Pacific.
crenulata (Cancellaria) Deshayes, 1835:501, pi. 79,
figs. 31-33. Lower Eocene, France.
crenulata (Cancellaria) A. Adams, 1855:124. Recent, ?
China.
crenulatus (Triton) 'Pease' - Carpenter, 1865:517; ex
Pease MS, in synonymy of Triton antiquatits Hinds.
Nomen nudum.
crepini (Cancellaria) Briart & Comet, 1877:13, pi. 14,
figs. 6a-c. Paleocene, Belgium.
cretacea (Cancellaria) Nyst, 1881:8; n.n. for C. obtusa
Binkhorst, 1861, non Deshayes, 1830. Cretaceous,
Belgium.
cretacea (Cancellaria) E. A. Smith, 1899b:245. Recent,
India. (= C. quasilla Petit, 1987)
cretaceus (Plesiotriton) Sohl, 1960:128, pi. 18, figs. 35,
41-42. Cretaceous, Mississippi, U.S.A.
crispa (Cancellaria) G. B. Sowerby I. 1832b:fig. 30.
Recent, Indo-Pacific.
crispa (Admete) Moller, 1842:88. Recent, North Atlan-
tic. (?= Admete viridula (Fabricius, 1780))
crispata (Admete) 'Muller' - Nyst, 1881:12. Error for
A. crispa Moller.
crispata (Cancellaria) 'Sowerby' - G. B. Sowerby II,
1849b:452; et al. Error for C. crispa G. B. Sowerby
I.
crispata (Cancellaria) Seguenza, 1880:110; as var. of
C. italica D'Ancona. Tertiary, Italy.
cristata (Admete) Marwick. 1926:323, pi. 73, fig. 16.
Miocene, New Zealand.
crossei (Cancellaria) Semper, 1861:257; n.n. for C.
serrata Reeve, 1856, non Bronn, 1831. Recent.
Indo-Pacific.
crossletensis (Cancellaria) Covacevich & Frassinetti,
1986:45, pi. 2, figs, la-c, 2a-c, 3a-c, text-figs. 5-6.
Miocene, Chile.
cruzialis (Cancellaria) von Ihering. 1907:214; n.n. for
"C. medinae Philippi" Ortmann, 1902:236, pi. 36,
figs. 4a-b. Tertiary, Argentina.
cubapatriae (Colubraria) Sarasua. 1975:4, figs. 1-2.
Recent, Cuba. [Tritonoharpa]
culminata (Inglisella) Beu, 1970:223, pi. 1, figs. 4-5.
Miocene, New Zealand.
cumingiana (Cancellaria) Petit de la Saussaye, 1844:pl.
112. Recent, Panamic-Pacific.
cumingii (Trigonostoma) 'Sowerby" - H. Adams & A.
Adams, 1854:276; et al. Error for C. cuminigiana
Petit de la Saussaye.
carta (Cancellaria) von Koenen, 1885:12, pi. 1, figs.
8a-b. Paleocene. Denmark.
cytharella (Cancellaria) 'Brongniart' - Pusch, 1837:pl.
11, fig. 16; plate caption error for citharella.
D
dalli (Fossarus) Anderson & Martin, 1914:70, pi. 7,
figs. 13a-b. Miocene, Califomia, U.S.A. [Cancel-
lariidae; see F. barkeri F. M. Anderson, 1924]
Page 18
THE NAUTILUS, Supplement 1
dalli (Cancellaria) Bartsch, 1915:33, pi. 4, fig. 2.
Recent, "South Africa". (?= C. buHata Sowerby,
1832)
dalliana {Cancellaria) F. M. Anderson, 1905:199, pi.
15, figs. 39-42. Miocene, California, U.S.A.
dampierensis {Fiisiaphera) Garrard, 1975:17, pi. 2, fig.
8. Recent, Australia.
danieli {Cancellaria) Morlet, 1885a:51, pi. 3, figs. 2,
2a. Middle Eocene, France.
dariena {Cancellaria) Toula, 1909:703, pi. 25, fig. 13;
pi. 28, fig. 2. Miocene, Panama.
darienensis {Cancellaria) Toula - Cossmann, 1913a:51.
Emendation of dariena.
daniini {Cancellaria) Petit, 1970:85, pi. 1, figs. 4a-c.
Recent, Galapagos Islands.
daulzenbergi {Cancellaria) Cossmann, 1896b:210, pi.
4, figs. 34-35. Middle Eocene, France.
davidsoni {Triton) d'Archiac & Haime, 1854:312, pi.
30, figs. 3, 3a. Eocene, Burma, [placed in Cancel-
laria by Noetling, 1901:331; not a cancellariid]
decapensis {Admete) Barnard, 1960:439, fig. la. Recent,
South Africa.
decaptyx {Cancellaria) Brown & Pilsbry, 1911:346, pi.
24, figs. 5-6. Miocene, Panama.
decorata {Trigonostoma) Newton, 1922:43, pi. 5, figs.
18-19. Eocene, Nigeria.
decussata {Cancellaria) G. B. Sowerby I, 1832a:55;
1832b:fig. 8. Recent, Panamic-Pacific.
decussata {Cancellaria) Nyst, 1838:115, pi. 1, fig. 5.
?Recent, unknown locality. (?= C. oblonga Sow-
erby)
decussata {Cancellaria) Bellardi, 1841:26. Tertiary,
Europe.
decussata {Cancellaria) G. B. Sowerby I, 1847:421, pi.
20, fig. 27. Tertiary, Portugal.
decussata {Cancellaria 'Smith, 1847'. Error for C.
decussata Sowerby, 1847.
decussata {Cancellaria) Grateloup, 1847:pl. 25, fig. 20;
see Appendix, Note 1. Tertiary, France.
decussata {Coralliophila) de Loriol, 1882:19, pi. 2, figs.
22-25. ? Cretaceous, Europe. [? Cancellariidae]
defuniak {Cancellaria) Gardner, 1937:365, pi. 44, figs.
1-2. Miocene, Florida, U.S.A.
delecta {Cancellaria) Deshayes, 1864:99; n.n. for C.
elegans Deshayes, 1835, non Sowerby, 1822. Mid-
dle Eocene. France.
deleta {Oamanna) Finlay, 1930b:241, pi. 43, fig. 20.
Recent, New Zealand.
delicatula {Admete) E. A. Smith, 1907:4, pi. 1, figs. 5,
5a. Recent, Antarctic.
deliciosa - error for delicosa.
delicosa {Sydaphera) Laseron, 1955:269, figs. 5, 5a.
Recent, Australia.
dein {Waipaoa) Beu, 1970:224, pi. 4, fig. e. Miocene,
New Zealand.
dennanti {Plesiotriton) Tate, 1898:383, pi. 19, fig. 1.
Eocene, Australia.
densata {Paladmete) Wade, 1926:108, pi. 35, figs. 7-8.
Cretaceous, Tennessee, U.S.A.
densestriata {Sveltia) Peyrot, 1928:226, pi. 13, fig. 44;
as var. of S. colpodes Cossmann. Miocene, France.
dentifera {Cancellaria) Deshayes, 1864:98, pi. 73, figs.
8-10. Lower Eocene, France.
deperdita {Cancellaria) Michelotti, 1861:102, pi. 11,
figs. 5-6. Oligocene, Italy.
depressa {Cancellaria) Tuomey & Holmes, 1856:143,
pi. 28, fig. 17. Pliocene, South Carolina, U.S.A.
depressa {Cancellaria) Dall, 1915:48, pi. 10, fig. 4.
Oligocene, Florida, U.S.A. {- Trigonostoma tam-
paensis Petit, 1967)
depresseplicata {Brocchinia) Sacco, 1894:69, pi. 3, fig.
85; as var. of B. mitraeformis (Br.). Pliocene, Italy.
depressicosta {Cancellaria) Sacco, 1894:22, pi. 2, fig.
2ter; as var. of C. acutangula Faujas de Saint Fond;
see Appendix, Note 2. Miocene, Italy.
depressocostata {Cancellaria) Sacco, 1894:11, pi. 1, fig.
25; as var. of C. fene strata Eichwald; see Appen-
dix, Note 2. Miocene, Italy.
deroyae {Agatrix) Petit, 1970:85, pi. 1, figs. 3a-b.
Recent, Galapagos Islands.
dertinflata {Cancellaria) Sacco, 1894:25, pi. 2, figs.
17a-c; as var. of C. miitinensis Foresti; see Appen-
dix, Note 2. Miocene, Italy.
dertocacellensis {Cancellaria) Sacco, 1894:49; n.n. for
"C. contorta Basterot" Pereira da Costa, 1867:pl.
24, fig. 6; as var. of C. contorta Basterot; see
Appendix, Note 2. Miocene, Portugal.
dertocacellensis {Cancellaria) Sacco, 1894:30; n.n. for
C. harjonae Pereira da Costa. 1867:pl. 25, fig. 9
only; as var. of C. piscatoria (Gmelin); see Appen-
dix, Note 2. Miocene, Portugal.
dertocassideum {Cancellaria) Sacco, 1894:8, pi. 1, figs.
14a-b; see Appendix, Note 2. Miocene, Italy.
dertocontorta {Cancellaria) Sacco, 1894:49, pi. 3, figs.
27a-b; see Appendix, Note 2. Miocene, Italy.
dertoconvexula {Cancellaria) Sacco, 1894:60, pi. 3, fig.
59; as var. of C. lyrata (Br.); see Appendix, Note
2. Miocene, Italy.
dcrtoconvexulata {Cancellaria) Sacco, 1894:62, expl.
to pi. 3; error for dertoconvexula.
dertocostatissima {Cancellaria) Sacco, 1894:60, pi. 3,
fig. 60; as var. of C. lyrata (Br.); see Appendix,
Note 2. Miocene, Italy.
dertocosticillata {Cancellaria) Sacco, 1894:13, pi. 1,
figs. 31a-b; as var. of C. imbricata Homes; see
Appendix, Note 2. Miocene, Italy.
dertocrenata {Cancellaria) Sacco, 1894:44, pi. 3, figs.
lla-b. Miocene, Italy; see Appendix, Note 2.
[Ranellidae]
dertofusula {Cancellaria) Sacco, 1894:49; n.n. for "C.
contorta Bast." Homes, 1854:pl. 34, fig. 7; as var.
of C. contorta Basterot; see Appendix, Note 2.
Miocene, Austria.
dertogranosa {Bonellitia) 'Sacco" - Ferrero Mortara,
1984:171. Nomen nudum.
derlolyratoides {Cancellaria) Sacco, 1894:36, pi. 2, fig.
54; as var. of C. uniangulata Deshayes; see Appen-
dix, Note 2. Miocene, Italy.
R. E. Petit and M. G. Harasewych, 1990
Page 19
dertonassoides (Cancellaria) Sacco, 1894:40, pi. 2, fig.
62bis; as var. of C. dertonensis Bellardi; see
Appendix, Note 2. Miocene, Italy.
dertonensis {Cancellaria) Bellardi, 1840:344; as var. of
C. honelli Bellardi. Nomen nudum.
dertonensis {Cancellaria) Bellardi, 1840:344; as var. of
C. cancellata Lk. [sic]. Nomen nudum.
dertonensis {Cancellaria) Bellardi, 1841:24, pi. 3, figs.
11-12; as var. of C. bonellii Bellardi. Tertiary, Italy.
dertonensis {Cancellaria) Bellardi, 1841:28, pi. 3, figs.
13-14; as var. of C. cancellata (Lamarck) [sic].
Tertiary, Italy.
dertopana {Cancellaria) Sacco, 1894:7; n.n. for "C
scahra Desh." Homes, 1856:pl. 51 [sic: error for
pi. 52], fig. 7; as var. of C. scabra Deshayes; see
Appendix, Note 2. Miocene, Austria.
dertopercostata {Cancellaria) Sacco, 1894:34, pi. 2, fig.
47: as var. of C. calcarata (Brocchi); see Appendix,
Note 2. Miocene, Italy.
dertopercostulata {Cancellaria) Sacco, 1894:36, pi. 2,
fig. 53; as var. of C. uniangulata Deshayes; see
Appendix, Note 2. Miocene, Italy.
dertopostica {Cancellaria) Sacco. 1894:12, pi. 1, fig.
27; as var. of C fenestrata Eichwald; see Appen-
dix, Note 2. Miocene, Italy.
dertoscalata {Cancellaria) Sacco, 1894:17, pi. 1, figs.
45a-b, 45bis a-b; see Appendix, Note 2. Miocene,
Italy.
dertosimplex {Cancellaria) Sacco, 1894:30, pi. 2, fig.
32; as var. of C. hirta (Brocchi); see Appendix,
Note 2. Miocene, Italy.
dertosuturata {Cancellaria) Sacco, 1894:56, pi. 3, fig.
46; as var. of C. varicosa (Brocchi); see Appendix,
Note 2. Miocene, Italy.
dertoturrita {Cancellaria) Sacco, 1894:30, pi. 2, fig. 33;
as var. of C. barjonae Pereira da Costa; see
Appendix, Note 2. Miocene, Italy.
dertovaricosa {Cancellaria) Sacco, 1894:57, pi. 3, figs.
49a-b; see Appendix, Note 2. Miocene, Italy.
dertumbilicata {Cancellaria) Sacco, 1894:10, pi. 1, fig.
21; as var. of C gradata Homes; see Appendix,
Note 2. Miocene, Italy.
dertumbilicatior {Trigonostoma) "Sacco' - Ferrero Mor-
tara, 1984:160. Nomen nudum.
deshayesana {Cancellaria) Grateloup, 1832:338. Terti-
ary, France, (incorrectly attributed to Desmoulins
by later authors, including Grateloup)
deshayesi {Cancellaria) 'Desm.' - Cossniann, 1899a:16.
Error for C. deshayesana Grateloup.
deshayesiana {Cancellaria) 'de Moulins' - Crosse,
1861:248; et al. Error for deshayesana Grateloup.
deshayesianus {Plesiotriton) Beu & Maxwell, 1987:25,
pi. 23, figs, a-e, h; n.n. for Triton turriculatum
Deshayes, 1835, non 1833. Eocene, France.
deshayesii {Cassidaria) Duval, 1841:278. Recent, Sene-
gal. [Loxotaphrus]
desmotis {Cancellaria) Gardner, 1937:373, pi. 45, fig.
5. Miocene, Florida, U.S.A.
desori {Cancellaria) Mayer, 1876:44, pi. 4, fig. 10.
Tertiary, Europe.
deucalionis {Cancellaria) 'Eichwald' - Sherborn,
1922:1888. Error for Cassis deucalionis Eichwald,
1830.
dewalquei {Cancellaria) 'Nyst' - Dewalque, 1868:419.
Nomen nudum.
deydieri {Cancellaria) Fontannes, 1878:78, pi. 1, figs.
4a-b; 1879a:515, pi. 5, figs. 4a-b. Tertiary, France.
diadela {Cancellaria) Woodring, 1970:338, pi. 53, figs.
7, 9. Pliocene, Panama.
diadema {Cancellaria) Watelet, 1853:22, pi. 2, fig. 12.
Middle Eocene, France.
diamantina {Trigonostoma) Garrard, 1975:22, pi. 3, fig.
4. Recent, Australia.
dictyella {Cancellaria) Cossmann, 1899a:24. Nomen
nudum.
dingdensis {Narona) H. J. Anderson, 1964:274, pi. 30,
fig. 216. Miocene, Germany.
dingdensis {Pseudomalaxis) H. J. Anderson, 1964:203,
pi. 15, figs. 109a-c. Miocene, Germany. [Trigono-
stoma]
dinota {Cancellaria) Woodring, 1970:340, pi. 54, figs.
1-2; pi. 56, figs. 5-6. Miocene, Panama.
disparstriata {Cancellaria) Holzl, 1958:257, pi. 21, fig.
1 1 ; as var. of C. trochlearis Faujas de Saint Fond.
Miocene, Germany.
disticha {Inglisella) Marwick, 1965:41, pi. 11, figs.
9-10. Pliocene, New Zealand.
distincta {Admete) Leche, 1878:48, pi. 1, figs. 14a-b;
as var. of A. viridula (Fabricius). Recent, North
Atlantic.
distincta {Cancellaria) Mayer, 1876:44, pi. 4, fig. 11.
Tertiary, Europe.
distinguenda {Charcolleria) Petit, 1970:84, pi. 1, figs.
2a-b. Miocene, Florida, U.S.A.
doboi {Chijsodomus) Noszky, 1936:66, pi. 5, fig. 5.
Oligocene, Hungary. [Turehua]
doderleini {Cancellaria) Mayer, 1868:108, pi. 2, fig. 5.
Tertiary, Italy.
doderleini {Cancellaria) Moroni, 1958:74, pi. 1, figs.
I, la. Miocene, Italy. (?= C. barjonae Pereira da
Costa, 1867)
doliaris - error for doliolaris.
dolioides {Cancellaria) Pilsbry & Olsson, 1941:22, pi.
3, fig. 5. Pliocene, Ecuador.
dolioides {Pristimerica) Finlay & Marwick, 1937:82, pi.
II, figs. 2-3. Paleocene, New Zealand.
doliolaris {Cancellaria) Basterot, 1825:46, pi. 2, fig.
17. Miocene, France.
doncieuxi {Cancellaria) Dareste de la Chavanne, 1910:24,
pi. 4, figs. 8a-b. Eocene, Algeria.
dregeri {Cancellaria) Hoemes & Auinger. 1890:280, pi.
33, figs. 18a-c, 19a-c. Miocene, Austria.
druentica {Cancellaria) Fontannes, 1878:76, pi. 1, fig.
2; 1879a:514, pi. 5, fig. 2. Tertiary, France.
druidarum {Cancellaria) Gardner, 1937:367, pi. 44,
figs. 7-8. Miocene. Florida, U.S.A.
druidi {Trigonostoma) Olsson & Petit, 1964:544, pi. 80.
figs. 6, 6a; pi. 82, fig. 8. Pliocene, Florida, U.S.A.
dubia {Cancellaria) Deshayes, 1864:105, pl. 73, figs.
25-27. Eocene, France.
Page 20
THE NAUTILUS. Supplement 1
dubia (Cancellaria) 'Edwards MS' - Wrigley, 1935:379.
Nomen nudum.
dubius {Latirus) Marshall, 1919:229, pi. 16, fig. 6.
Eocene, New Zealand. [Turehua; see Latirus mar-
shal li Finlay]
dubusi (Uxia) Cossmann & Pissarro, 1901:19, pi. 15.
figs. 19-20. Middle Eocene, France.
dufourii (Cancellaria) Grateloup, 1832:342. Miocene,
France.
dumasi (Sveltella) Cossmann, 1899a:30, 194, pi. 2, fig.
12. Miocene, France.
dunkeri {Cancellaria) Holzapfel, 1888:93, pi. 9, figs.
2a-c. Cretaceous, Germany.
duponti (Cancellaria) Briart & Comet, 1877:12, pi. 14,
figs. 4a-c. Paleocene, Europe.
E
eburnaeformis (Cancellaria) Reeve, 1856:pl. 5, fig. 21.
Recent, locality unknown. (?= C. obesa Sowerby,
1832)
echinata (Cancellaria) 'Doderlein' - Sacco, 1894:7.
Nomen nudum.
ecuadoriana (Trigonostoma) Pilsbry & Olsson, 1941:25,
pi. 3, figs. 8-9: pi. 5, fig. 2. Pliocene, Ecuador.
effosa (Cancellaria) Handmann, 1882:263. Miocene,
Austria.
egregia (Cancellaria) von Koenen, 1889:142, pi. 12,
figs. 13a-c. Oligocene, Germany.
elata (Cancellaria) Hinds, 1843:48; 1844b:42, pi. 12,
figs. 3-4. Recent, Panamic-Pacific.
elatior (Cancellaria) von Koenen, 1889:135, pi. 12,
figs. 5a-c; as var. of C. niiens Beyrich. Oligocene,
Germany.
elatocostata (Cancellaria) Sacco, 1894:37; as subvar.
of C. cancellata pluricusticillata Sacco; see Appen-
dix, Note 2. Tertiary, Italy.
elatocosticillata (Cancellaria) Sacco, 1894:38; as subvar.
of C. cancellata pluricosticillata Sacco; see Appen-
dix, Note 2. Tertiary, Italy.
elegans (Cancellaria) G. B. Sowerby I, 1822:fig. 3.
Recent, Indo-Pacific.
elegans (Cancellaria) "Bonelli' - Sacco, 1894:35; et
al.\ in synonymy of C. uniangulata Deshayes.
Nomen nudum.
elegans (Cancellaria) 'Gene' - Michelotti, 1838:396; et
al.; in synonymy of C. uniangulata Deshayes.
Nomen nudum.
elegans (Cancellaria) Deshayes, 1835:502, pi. 79, figs.
24-26. Middle Eocene, France. (= C. delecta De-
shayes, 1864)
elegans (Cancellaria) Karsten, 1849:25. Oligocene, Ger-
many. (?= Babylonella pusilla (Philippi, 1843))
elegans (Mataxa) Wade, 1917:456, pi. 23, figs. 1-3.
Cretaceous, Tennessee, U.S.A.
elegans (Paladmete) Stephenson, 1941:368, pi. 71, figs.
16-17. Cretaceous. Texas, U.S.A.
elegantula (Oamaruia) Beu, 1970:222, pi. 3, fig. f.
Miocene, New Zealand.
elegantula (Trigonostoma) M. Smith, 1947:54, pi. 2,
fig. 3. Recent, Panamic-Pacific.
elevata (Cancellaria) I. Lea, 1833:141, pi. 5, fig. 139.
Eocene, Alabama, U.S.A.
elevata (Admetopsis) D. W. Johnson, 1903:203, pi. 1,
fig. 14. Cretaceous, New Mexico, U.S.A.
elizabethae (Trigonostoma) Olsson & Petit, 1964:543.
pi. 80, figs. 2, 2a. Pliocene, South Carolina, U.S.A.
ellapsa (Cancellaria) Conrad, 1865d:212, pi. 20 [sic;
error for pi. 21], fig. 8. ? Eocene, Texas, U.S.A. (?=
Paladmete cancellaria (Conrad, 1858))
ellipsis (Cancellaria) Pilsbry, 1922:333. pi. 22, figs.
8-9. Tertiary, Dominican Republic.
elodiae (Cancellaria) Carson, 1926:49, pi. 1, fig. 1.
Pliocene, California, U.S.A.
elongata (Admete) Leche, 1878:48, pi. 1. figs. 13a-b;
as var. of A. viridula (Fabricius). ? Tertiary, Novaya
Zemlya.
elongata (Cancellaria) Nyst, 1845:476, pi. 12, figs.
21a-b. Tertiary, Europe.
elongata (Cancellaria) Wood, 1847:354. Nomen nudum,
elongata (Cancellaria) Grateloup, 1847:index p. 2, pi.
25, fig. 31; as var. of C. geslini Basterot. Tertiary,
France.
elongata (Cancellaria) 'Karsten' - Semper. 1861:255;
in synonymy of C. pusilla (Philippi). Error for C.
elegans Karsten.
elongata (Cancellaria) 'Sowerby' - Cossmann, 1903a:108.
Error for C. oblonga Sowerby.
elongata (Cancellaria) Kobelt, 1904:210, pi. 79, figs.
3-4; as var. of C cancellata (L.). Recent, northern
Africa.
elongata (U.xia) Cuvillier. 1935:66, pi. 5, figs. 22-23.
Eocene, Egypt.
elongata (Unitas) Traub, 1979:114, pi. 16, figs. 4a-b.
Paleocene, Austria. (= U. oichingensis Traub, 1984)
elongata (Olssonella) Dockery in MacNeil & Dockery,
1984:163, pi. 21, figs. 10. 12. Oligocene, Missis-
sippi, U.S.A.
elsmerensis (Cancellaria) English, 1914:216, pi. 23, fig.
8. Pliocene, California, U.S.A.
emmae (Admete) Albrecht & Valk, 1943:75, pi. 19, figs.
725-728. Oligocene, Netherlands.
emydis (Cancellaria) Dall & Ochsner, 1928:105, pi. 2,
fig. 7. Pleistocene, Galapagos Islands.
enderbyensis (Admete) Powell, 1958:201, pi. 1. fig. 10.
Recent, Antarctic.
engonata (Cancellaria) Conrad, 1841:32. Miocene, Mary-
land, U.S.A.
eocaenica (U.xia) 'Cossmann' - Cossmann & Pissarro,
1913:pl. 47, fig. 212-11; see infraeocaenica Coss-
mann.
eogassinense (Cancellaria) Sacco, 1904:119. pi. 25, fig.
1; see Appendix. Note 2. Eocene, Italy.
eoreticosa (Cancellaria) Cossmann, 1899c: 178. Unnec-
essary n.n. for C. reticulata 'Newton', a nomen
nudum.
epidromiformis (Cancellaria) Tate, 1889:154, pi. 8, fig.
9. Miocene, Australia.
R. E. Petit and M. G. Harasewych, 1990
Page 21
epistomifera (Cancellaria) Guppy, 1876:520, pi. 28, fig.
9. Miocene, Dominican Republic.
epomis (Trihia) Woodring, 1928:223, pi. 12, fig. 10.
Pliocene, Jamaica.
errata (Cancellaria) Sacco, 1894:15; n.n. for "C.
mkhelinii Bell." Pereira da Costa, 1867:pl. 25, fig.
8; as var. of C. michelinii Bellardi; see Appendix,
Note 2. Miocene, Portugal.
esi (Cancellaria) Koperberg, 1931:66, pi. 2, fig. 19.
Tertiary, Indonesia.
esmeralda (Cancellaria) Olsson, 1964:119, pi. 21, figs.
6, 6a-b. Pliocene, Ecuador.
etheridgei (Cancellaria) Johnston, 1880:32. Miocene,
New Zealand.
eucheea (Cancellaria) Gardner, 1947:636, pi. 52, fig.
43. Miocene, Florida, U.S.A.
euclethra (Cancellaria) Maury, 1925b: 188, pi. 9, fig.
13. Miocene, Brazil.
eudeli (Cancellaria) G. B. Sowerby III, 1893:27, text.
fig. Recent, ? Malaysia. (?= C. angasi Crosse)
euetrios (Cancellaria) Barnard, 1959:14, text-fig. 3.
Recent, South Africa.
eiifaulensis (Cancellaria) Gabb, 1860b:390, pi. 68, fig.
8. Cretaceous, Alabama, U.S.A.
eutaeniata (Cancellaria) Cossmann, 1896b:212, pi. 6,
figs. 23-24. Middle Eocene, France.
euthymei (Cancellaria) Barnard, 1960:438, text-fig. lb.
Recent, South Africa.
eva (Fusiaphera) Petit, 1980:215, figs. 5-6. Recent,
Mozambique.
evulsum (Buccinum) Solander, 1766:13, pi. 1, fig. 14.
Eocene, England. [Admetula]
exaltata (Cancellaria) Tate. 1889:154, pi. 8, fig. 10.
Miocene, Australia.
exampullacea (Cancellaria) Sacco, 1894:10; n.n. for
"C. ampullacea (Br.)"' Homes, 1854:pl. 35, fig. 4;
as var. of C. gradata Homes; see Appendix, Note
2. Miocene, Austria.
exbellardi (Cancellaria) Sacco, 1894:46; n.n. for "C
bellardi [sic] Mich." Wood, 1872:pl. 3, fig. 25; as
var. of C. evulsa (Solander); see Appendix, Note
2. Tertiary, England.
excassidea (Cancellaria) Sacco, 1894:8; n.n. for C
cassidea (Br.)" Homes, 1856:pl. 52, fig. 8; as var.
of C cassidea (Brocchi); see Appendix, Note 2.
Miocene, Austria.
excavata (Cancellaria) G. B. Sowerby U, 1849a:137;
1849b:449, pi. 93, fig. 18. Recent, Australia. (?= C.
spirata Lamarck, 1822)
excellens (Cancellaria) Beyrich, 1856:566, pi. 25, figs.
5a-b. Oligocene, Germany.
exdecussata (Aphera) Sacco, 1894:66; n.n. for "C.
decussata Smith [sic]" Pereira da Costa, 1867:pl.
26, fig. 5. Miocene, Portugal.
exgeslini (Cancellaria) Sacco, 1894:22; n.n. for "C.
geslini Bast." Homes, 1854:pl. 25 [sic; error for
35], fig. 3; as var of C. geslini Basterot; see
Appendix, Note 2. Miocene, Austria.
exigua (Cancellaria) E. A. Smith, 1891:439, pi. 34, fig.
11. Recent, Australia.
exiliplex (Sveliella) Hickman, 1980:73, pi. 9, figs. 7-9.
Oligocene, Oregon, U.S.A.
exilis (Fusus) Philippi, 1843:25, 60, pi. 4, fig. 12.
Oligocene, Germany. [?= Bahylonella piisilla
(Philippi, 1843)]
eximbricata (Cancellaria) Sacco, 1894:50, pi. 3, fig.
28; as var. of C. dertocontorta Sacco; see Appen-
dix, Note 2. Miocene, Italy.
eximia (Narona) Stoliczka, 1867:166, pi. 13, figs.
15-16. Cretaceous, India.
eximia {Cancellaria) 'Edwards MS.' - Newton, 1891:171.
Nomen nudum.
exopleura (Cancellaria) Dall, 1908:294. Recent, Panamic-
Pacific.
expidea (Cancellaria) - Tate, 1889:153. ? Error for C.
cassidea (Brocchi). Nomen nudum.
explicata (Anapepta) Laws, 1935:38, pi. 6, fig. 20.
Miocene, New Zealand.
expusilla (Admete) Sacco, 1894:71; n.n. for "C. pusilla
(Phil.)" Beyrich, 1856:pl. 28, fig. 2; as var. of A.
minuta Braun. Tertiary, Germany.
exquisita (Cancellaria) Preston, 1905:3, pi. 1, fig. 9.
Recent, Sri Lanka.
exscrobiculata {Cancellaria) Sacco, 1894:17; n.n. for
"C. scrobiculata Homes" Pereira da Costa, 1867:pl.
26, fig. 2; as var. of C dertoscalata Sacco; see
Appendix, Note 2. Miocene, Portugal.
extractrix (Discoheli.x) Boettger, 1906:138. Miocene,
Romania. [Extractrix]
exumbilicaris (Cancellaria) Sacco, 1894:6; ? n.n. for C.
[sic] bellardii de Stefani & Pantanelli, 1878, non
Michelotti, 1 847; as var. of C. umbilicaris (Br.); see
Appendix, Note 2. Pliocene, Italy.
exvaricosa (Cancellaria) Sacco, 1894:58; n.n. for "C.
varicosa (Br.)" Beyrich, 1856:pl. 27, fig. 6; as var.
of C. taurinia Bellardi; see Appendix, Note 2.
Tertiary, Germany.
exwestiana (Cancellaria) Sacco. 1894:24; n.n. for "C
westiana Grat." Homes, 1854:pl. 35, figs. 11-12;
see Appendix, Note 2. Miocene, Austria.
F
farafrense (Triton) Quaas, 1902:273, pi. 32, fig. 47.
Cretaceous, Libya. [?= Plesiotriton]
fenestrata (Cancellaria) Eichwald, 1830:222. Tertiary,
Europe.
fenollerae (Ovilia) Landau, 1984:149, pi. 2, figs. 1-5.
Pliocene, Spain.
fergusoni (Cancellaria) Carson, 1926:53, pi. 1, figs.
7-8. Pleistocene, California, U.S.A.
fernandoensis (Cancellaria) Amold, 1907a:535, pi. 50,
fig. 4. Pliocene, California, U.S.A.
fetzaraensis (Cancellaria) Dareste de la Chavanne,
1910:24, pi. 4, figs. 9a-d. Eocene, Algeria.
fictilis (Triton) Hinds, 1844a:21; 1844b: 12, pi. 4. figs.
11-12. Recent, South Africa. [Africotriton]
ficus (Trigonostoma) K. Martin, 1931:11, pi. 2, figs. 5,
5a. Eocene, Indonesia.
finexa (Pleurotoma) G. D. Harris, 1895a:64, pi. 5, fig.
13. Eocene, Texas, U.S.A. [Cancelrana]
Page 22
THE NAUTILUS, Supplement 1
finlayi {Anapepta) Marvvick, 1931:122, pi. 13, fig. 238.
Miocene, New Zealand.
finlayi (Zeadmete) Powell, 1940:242, pi. 29, fig. 12.
Recent, New Zealand.
fischeri (Cancellaria) A. Adams, 1860:411. Recent,
Indo-Pacific.
flemingi (Taiara) Beu & Maxwell, 1987:53, pi. 27, figs.
a-e. Eocene/Oligocene, New Zealand.
floridana {Cancellaria) Olsson & Petit, 1964:540, pi.
82, fig. 6; as subsp. of C. rotunda Dall. Pliocene,
Florida, U.S.A.
floridensis (Colubraria) Tucker & Wilson, 1932:11, pi.
4, figs. 3-4. Pliocene, Florida, U.S.A. [Tritonoliarpa]
fontinalis {Cancellaria) 'Tate' - Tate & Dennant,
1893:221. Nomen nudum.
forestieri {Cancellaria) Montrouzier in Souverbie &
Montrouzier, 1863:161, pi. 5, fig. 7. Recent, New
Caledonia. (?= C. contabulata Sowerby, 1832)
fornicis {Uxia) Wrigley, 1935:376, pi. 34, fig. 31.
Lower Eocene, England.
foveata {Cancellaria) Almera & Bofill, 1884:32, pi. C,
figs. 10-12. Tertiary, Spain.
foveolata {Cancellaria) G. B. Sowerby U, 1849a: 137;
1849b:455, pi. 93, figs. 30-31. Recent, South
Africa.
fragilis {Sveltella) Wrigley, 1935:363, pi. 35, fig. 41.
Lower Eocene, England.
fragosa {Cancellaria) Olsson, 1964:123, pi. 21, fig. 8.
Miocene, Ecuador.
fi-esvillensis {Uxia) Cossmann & Pissarro, 1901:21, pi.
8, fig. 2. Midle Eocene, France.
frigida {Admete) Rochebrune & Mabille, 1885:104.
Recent, Cape Horn.
frizzelli {Cancellaria) Marks, 1949:462, pi. 78, fig. 5.
Miocene, Ecuador.
fugleri {Cancellaria) Arnold, 1907b:433, pi. 54, fig. 9;
as var. of C. crawfordiana Dall. Pliocene, Califor-
nia. U.S.A.
fumiculata — error for funiculata.
fundata (Bonellitia) Marwick, 1931:120, pi. 13, fig.
235. Pliocene, New Zealand.
funerata {Cancellaria) Conrad, 1848a:287: 1848b:118,
pi. 11, fig. 39. Oligocene, Mississippi, U.S.A.
fiiniculata {Cancellaria) Hinds, 1843:48; 1844b:43, pi.
12, figs. 5-6. Recent, Panamic-Pacific.
funiculifera {Cancellaria) Vincent in Vincent & Lef evre,
1872:59, pi. 2, fig. 1. Oligocene, Belgium.
funigera {Admete) Staadt in Cossmann, 1913b: 188, pi.
3, fig. 212-ter-9. Paleocene, France.
fusca {Cancellaria) G. B. Sowerby HL 1889:568, pi.
28, fig. 12. Recent, Hong Kong.
fuscoapicata {Iphinopsis) Bouchet & Waren, 1985:262,
figs. 700-703. Recent, off British Isles.
fusiformis {Cancellaria) Cantraine, 1835:391. Tertiary,
Italy.
fusiformis {Cancellaria) 'Philippi, 1845' - Sherbom,
1926:2610. Error for Fasciolaria fusiformis Philippi,
1845.
fusiformis {Cancellaria) Deshayes, 1864:102, pi. 72,
figs. 31-32. Middle Eocene, France.
fusoascalaris {Cancellaria) Sacco, 1894:25. pi. 2, fig.
16; as var. of C. exwestiana Sacco; see Appendix.
Note 2. Miocene, Italy.
fusoconvexa {Admete) Sacco, 1894:71, pi. 3, fig. 95;
as var. of A. costellifera [sic] (Sowerby). Pliocene,
Italy.
fusosimplex {Cancellaria) Sacco, 1894:49; n.n. for "C.
contorta Bast." Homes, 1854:pl. 34, fig. 8; as var.
of C. contorta Basterot; see Appendix, Note 2.
Miocene, Austria.
fusospinosa {Cancellaria) Sacco, 1894:65, pi. 3, figs.
74a-b; see Appendix, Note 2. Miocene, Italy.
fusulus {Cancellaria) Bronn, 1831:43. Tertiary, Italy.
(?= C uniangulata Deshayes, 1830)
fusus {Cancellaria) 'Bronn" - Deshayes, 1843:423.
Error for C. fusulus Bronn.
G
gabbiana {Cancellaria) Pilsbry & Johnson, 1917:163.
Miocene, Dominican Republic.
gailleti {Uxia) Cossmann, 1913b:201, pi. 7, fig. 212-24.
Eocene, France.
gainfarnensis {Cancellaria) Handmann, 1882:262. Mio-
cene, Austria.
galei {Cancellaria) Addicott, 1970:119, pi. 16, figs.
6-10. Miocene, California, U.S.A.
gallica {Sveltia) Peyrot, 1928:229, pi. 14, figs. 11-14;
as mut. of S. inermis (Pusch). Miocene, France.
galvestonensis {Cancellaria) G. D. Harris, 1895b: 100,
pi. 3, fig. 11. Miocene, Texas, U.S.A.
gardnerae {Paladmete) Wade, 1926:108, pi. 35, figs. 3,
6. Cretaceous, Tennessee, U.S.A.
garrardi {Admetula) Petit, 1974:109; n.n. for C. nassoi-
des Schepman, 1911, non von Kocnen, 1889.
Recent, Indo-Pacific.
garvani {Bonellitia) Palmer, 1937:452, pi. 69, figs. 1-3.
Eocene, southeastern U.S.A.
gaudryi {Cancellaria) Fontannes, 1878:77, pi. 1, figs.
3a-b; 1879a:514, pi. 5, figs. 3a-b. Tertiary, France.
gelriana {Cancellaria) Janssen, 1972:39, pi. 8, figs.
5a-b, 6; as subsp. of C. contorta Basterot. Miocene,
Netherlands.
gemmata {Cancellaria) Conrad, 1833:35; 1835:44, pi.
16, fig. 10. Eocene, Alabama, U.S.A.
gemmata {Oamaruia) Maxwell, 1969:182. Oligocene,
New Zealand.
gemmulata {Cancellaria) G. B. Sowerby 1, 1832a:55;
1832b:fig. 7. Recent, Panamic-Pacific.
gerda {GerdJella) Olsson & Bayer, 1972:877, figs. 1-3.
Recent, Caribbean.
gerthi {Merica) K. Martin, 1931:10, pi. 2, figs. 4, 4a.
Eocene, Indonesia.
geslini {Cancellaria) Basterot, 1825:46, pi. 2, fig. 5.
Tertiary, France.
gibbera {Oamaruia) Marwick, 1931:120, pi. 13, fig.
236. Miocene, New Zealand.
gilberti {Cancellaria) Aldrich, 1921:13. pi. 1. fig. 29.
Eocene, Alabama, U.S.A.
girauxi {Admete) Cossmann, 1913b:203, pi. 7, fig.
212-ter-lO. Middle Eocene, France.
R. E. Petit and M. G. Harasewych, 1990
Page 23
glabra (Canccllaria) Tesch, 1915:40. pi. 7, figs. 84a-b.
Tertiary, Indonesia.
glahrata (Cancellaria) 'Bosquet' - Vincent, 1886:6.
Nonicii nudum.
glabricula (Cancellaria) von Koenen, 1894:1397, pi.
100, figs. 4a-c. Oligocene, Gemiany.
gladiator (Cancellaria) Petit, 1976:35, pi. 1, fig. 2.
Recent, Galapagos Islands.
gliberti (Unitas) Dolin, Dolin & Le Renard, 1980:43,
pi. 2, fig. 19. Middle Eocene, France.
gliberti (Sveltia) Janssen, 1984:15, pi. 2, figs. 14-15; pi.
5, fig. 6. Miocene, Belgium.
globularis (Admete) E. A. Smith, 1875:426. Recent,
Japan. [? Microglyphis]
globularis (Cancellaria) 'Edwards MS' - Jones, 1878:236.
Nomen nudum.
globulosa (Cancellaria) Holzapfel, 1888:93, pi. 9, figs.
I, la-b. Cretaceous, Germany.
goniostoma (Cancellaria) G. B. Sowerby I, 1832a:51;
1833:fig. 43. Recent, Panamic-Pacific.
govenderi (Cancellaria) King. 1953:67, 83, fig. 7.
Miocene, Zululand.
gracilenta (Cancellaria) Wood, 1872:46, pi. 3. fig. 23.
Tertiary. England.
gracilenta (Cancellaria) 'Edwards MS' - Newton,
1891:171. Nomen nudum.
gracilina (Cancellaria) Sacco, 1894:13, pi. 1, fig. 34;
as var. of C. crassicosta Bellardi; see Appendix,
Note 2. Miocene, Italy.
gracilior (Cancellaria) Carpenter in Gabb, 1869:50.
Tertiary, California, U.S.A.
gracilis (Cancellaria) Philippi, 1845:450. Nomen nu-
dum.
gracilis (Cancellaria) von Ihering, 1897:310, pi. 3, fig.
I I . Tertiary, Argentina.
gracilis (Merica) Friedberg. 1914:243, pi. 15, fig. 6; as
var. of M. fenestrata (Eichwald). Miocene, Poland.
graciloides (Cancellaria) Aldrich, 1898:98. Eocene, Ala-
bama. U.S.A.
gradala (Cancellaria) M. Homes, 1854:319, pi. 35, figs.
2a-b. Miocene, Austria.
gradata (Cancellaria) Tate, 1889:155, pi. 10, fig. 12.
Miocene, Australia. (= Aneurystoma tatei Coss-
mann. 1899)
grandis (Admete) Morch, 1869:22; as var. of A. viridula
(Fabricius). Recent, Arctic Ocean.
granifera (Cancellaria) Deshayes. 1830:183. Eocene,
France.
granosa (Cancellaria) G. B. Sowerby I. 1832b:figs.
16-17. Recent, Australia.
granulata (Cancellaria) Wood, 1842:538; 1872:48. No-
men nudum.
granulata (Cancellaria) Nyst, 1845:479; 1845-46:pl.
39, fig. 14. Tertiary, Belgium.
granulosa (Cancellaria) 'Sow.' - Lesson, 1842:205.
Error for C. granosa Sowerby.
grata (Waipaoa) Marwick, 1931:122, pi. 13, fig. 240.
Miocene, New Zealand.
grateloupi (Cancellaria) d'Orbigny - Crosse, 1861:247.
First emendation of C. gratteloupi d'Orbigny, here
accepted as correct emendation.
gratteloupi (Cancellaria) d'Orbigny, 1852:10; n.n. for
"C. acutangula Faujas" Grateloup, 1847, pi. 1,
figs. 2 & 4 only; see grateloupi. Tertiary, France.
gravecostata (Cancellaria) Holzl, 1958:257, pi. 21, figs.
12, 12a; as var. of C. trochlearis Faujas de Saint
Fond. Miocene, Germany.
grayi (Cancellaria) Tryon, 1885:70, pi. 3, fig. 33.
Recent, Philippines.
gregaria (Admete) Meek, 1873:501. Cretaceous, Utah,
U.S.A.
greggi (Cancellaria) G. D. Harris, 1899:26, pi. 3, fig.
6; as var. of C. quercollis Harris. Lower Eocene,
Alabama and Texas, U.S.A.
grossauensis (Cancellaria) Handmann, 1882:262. Mio-
cene, Austria.
guppyi (Cancellaria) Gabb, 1873:236. Miocene, Do-
minican Republic.
gurabis (Cancellaria) Maury, 1917:65, pi. 10, fig. 11.
Miocene, Dominican Republic.
guttoides (Sveltella) Staadt in Cossmann, 1913b: 187, pi.
3, fig. 212-bis-7. Paleocene, France.
H
habei (Nipponaphera) Petit, 1972:103; n.n. for "N.
lyrata (Adams & Reeve)" Habe, 1961a:72, pi. 35,
fig. 18. Recent, Japan.
haemastoma (Cancellaria) G. B. Sowerby I, 1832a:54;
1833:fig. 40. Recent, Galapagos Islands.
haemostoma (Cancellaria) 'Sowerby'. Error for haema-
stoma Sowerby.
hamlini (Cancellaria) Carson, 1926:51, pi. 1, figs. 4,
6. Pliocene, California, U.S.A.
hampdenensis (Bonellitia) Marshall & Murdoch,
1923:124, pi. 12, fig. 4. Eocene, New Zealand.
harmeri (Cancellaria) Gilbert, 1958:16. Nomen nudum.
harmeri (Cancellaria) Gilbert, 1960a:2, pi. 4, fig. 1.
Pliocene, Belgium.
harpa (Cancellaria) von Koenen, 1889:145, pi. 9, figs.
3a-d. Oligocene, Germany.
harpiformis (Cancellaria) Pilsbry & Olsson, 1941:23,
pi. 3, figs. 1-2. Pliocene, Ecuador.
harpovoluta (Admete) Powell. 1957:143, pi. 1, fig. 3.
Recent. Antarctic.
harrisi (Cancellaria) Maury, 1917:64, pi. 10, figs. 9-10.
Miocene. Dominican Republic.
harrisi (Trigonostoma) Palmer. 1937:444. pi. 70, figs.
3-6. Eocene, Texas, U.S.A.
hartti (Cancellaria) Maury, 1925b: 184, pi. 9, fig. 19.
Miocene, Brazil.
haswelli (Gergovia) Garrard, 1975:36, pi. 4, fig. 7.
Recent. Australia.
haweraensis (Merica) Laws, 1940:54, pi. 7, fig. 26.
Pliocene, New Zealand.
hebertiana (Cancellaria) M. Homes, 1856:680, pi. 52,
figs. 5a-b. Miocene, Austria.
helenae (Trigonostoma) Olsson & Petit, 1964:543. pi.
80, figs. 3, 3a. Pliocene, Florida, U.S.A.
Page 24
THE NAUTILUS, Supplement 1
hemphilli (Cancellaria) Dall, 1909a:30, pi. 14, fig. 5.
Pliocene, California, U.S.A.
hettneri {Cancellaria) F. M. Anderson, 1929:114, pi.
10, figs. 5-6. Miocene, Colombia.
hidalgoi {Narona) Jousseaume, 1887a: 164, fig. 3. Re-
cent, western Mexico. (?= Cancellaria clavarula
Sowerby, 1832)
hidasensis {Cancellaria) Hoemes & Auinger, 1890:276,
pi. 33, figs. 13a-b, 14a-b. Miocene, Austria.
hillegondae {Eutritonium) K. Martin, 1914:150, pi. 4,
figs. Ill, Illa-c. Eocene/01 igocene, Indonesia. [Ple-
sioiriton]
hirta {Inglisella) Laws, 1936:116, pi. 16, fig. 69.
Pliocene, New Zealand.
hirta {Valuta) Brocchi, 1814:315, pi. 4, figs. la-b.
Pliocene, Italy. [Solalia]
hirtissima {Cancellaria) Sacco, 1894:24; n.n. for "C
spinifera Grateloup" Homes, 1854:pl. 35, fig. 8;
see Appendix, Note 2. Miocene, Austria.
hirtocostata {Cancellaria) Sacco, 1894:pl. 1, fig. 46;
plate caption error for C dertoscalata van Iri-
costata Sacco.
histrio {Scalptia) "Reeve' - Jousseaume, 1887b:213.
Error for C. hystrix Reeve.
hoelleitenensis {Palaeocancellaria) Kollmann, 1976:199,
pi. 1, figs. 7-9. Cretacous, Austria.
hoerlei {Trigonostoma) Olsson, 1967:24, pi. 8, figs. 6,
6a. Pliocene, Florida, U.S.A.
hoernesi {Cancellaria) Kittl. 1887:246, pi. 8, fig. 9.
Tertiary, Europe.
Iwffmanni {Neptiinea) Gabb, 1864:90, pi. 18, fig. 41.
Cretaceous, California, U.S.A. [?= Paladmete perfo-
rata (Gabb, 1864)]
hordeola {Cancellaria) von Koenen, 1889:99, pi. 12,
figs. 9a-c, lOa-c. Oligocene. Germany.
horii {Sydaphera) Masuda, 1967:10, pi. 2, figs. 16a-b,
17a-b, 18a-b, 19. Tertiary, Japan.
hukuiensis {Admete) Nomura & Niino, 1940:74, pi. 1,
fig. 5. Recent, Japan.
hukusimana {Cancellaria) Nomura & Hatai, 1936:134,
pi. 17, figs. 6a-b. Pliocene, Japan.
humerosa {Admetopsis) Stanton, 1893:160, pi. 33, figs.
4-5. Cretaceous, Utah, U.S.A.
hypermerces {Cancellaria) Cossmann, 1896a:42, pi. 3,
figs. 19-20. Middle Eocene, France.
hystrix {Cancellaria) Reeve, 1856:pl. 14, fig. 67. Re-
cent, Mauritius.
I
igarassuensis {Cancellaria) Penna, 1965:273, pi. 1, figs.
1-2, 4. Miocene, Brazil.
imbricata {Cancellaria) M. Homes, 1854:327, pi. 35,
figs. 16a-b. Miocene, Austria.
imbricata {Cancellaria) Watson, 1882a:325; 1886:274,
pi. 18, figs. lOa-c. Recent, South Africa. (= C.
africana Petit, 1970)
imbricatum {Tritoniwn) Dareste de la Chavanne, 1910:23,
pi. 4, fig. 6. Eocene, Algeria. [Plesiotriton]
imevborei (Bonellitia) Adegoke, 1977:206, pi. 31, figs.
16-19. Paleocene, Nigeria.
imperialis {Cancellaria) Michelin, 1832:[unnumbered
page], pi. 16. Recent, locality unknown. ('?= C.
cassidiformis Sowerby, 1832)
impressa {Cancellaria) Conrad, 1865a:32. Nomen nu-
dum.
impressa {Cancellaria) Conrad, 1865b: 145, pi. 11, fig.
16. Eocene, Alabama, U.S.A. (?= C gemmata
Conrad, 1833)
inaequalis {Paladmete) Stephenson, 1941:368, pi. 71,
figs. 14-15. Cretaceous, Texas, U.S.A.
incerta {Bonellitia) Harmer, 1918:405, pi. 40, fig. 9.
Pliocene, England.
incompta {Cancellaria) Briart & Comet, 1877:16, pi.
14, figs. 7a-c. Paleocene, Belgium.
indentata {Cancellaria) G. B. Sowerby I, 1832a:54;
1832b:figs. 9-10. Recent, Panamic-Pacific.
indicum {Tritonium) Spengler, 1923:37, pi. 3, figs.
19a-b. Cretaceous, India. [? Cancellariidae]
indicum {Trigonostoma) Vredenburg, 1921:139. Nomen
nudum.
indicum {Trigonostoma) Vredenburg, 1925:96, pi. 12,
figs. 4a-c. Tertiary, India.
indoceana {Tritonoharpa) Beu & Maxwell, 1987:40, pi.
19, figs, m-o; pi. 24, figs, m-p, s. Recent, Mozam-
bique.
inermis {Cancellaria) Pusch, 1837:129, pi. 11. fig. 22;
n.n. for Buccinum mitraeforme Pusch in Andrzejow-
ski, 1830. Tertiary, Poland.
inflata {Trichotropis) Friele, 1879:275. Recent, North
Atlantic. [Iphinopsis]
inflata {Admetula) Dockery in MacNeil & Dockery,
1984:164, pi. 21, figs. 8-9. Oligocene, Mississippi,
U.S.A.
infracosticillata {Cancellaria) Sacco, 1894:38, pi. 2,
fig. 59; as var. of C. cancellata (L.); see Appendix,
Note 2. Pliocene, Italy.
infracosticillata {Cancellaria) Sacco, 1894:47, pi. 3,
fig. 23; as var. of C. tauroconvexula Sacco; see
Appendix, Note 2. Miocene, Italy.
infraeocaenica (Cancellaria) Cossmann, 1889:224, pi.
7, fig. 27.' Paleocene, France..
inopinatus {Semitriton) Cossmann & Pissarro, 1905:92,
pi. 16, figs. 19-20. Eocene, France.
inornata (Cancellaria) Noetling, 1901:329, pi. 22, figs.
9a-c. Miocene, Burma.
insularis {Cancellaria) Pilsbr>' & Johnson, 1917:163.
Tertiary, Dominican Republic.
intercedens (Cancellaria) Stoliczka, 1867:164, pi. 13,
fig. 13. Cretaceous, India.
interlaevis (Trigonaphera) Laseron, 1955:270, figs. 9,
9a. Recent, Australia.
intermedia (Cancellaria) Bellardi, 1840:344. Nomen
nudum.
intermedia (Cancellaria) Bellardi, 1840:344; as var. of
C nodulosa Lamarck. Nomen nudum.
'Cossmann & Pissarro (1913:pl. 47. fig. 212-11) listed •■(Jxia
infraeocaenica Cossmann" wiili ihe notation "err. typ. eocaenica."
No usage of "eocaenica" has been located, and we are unsure as to
Ihc meaning of this remark.
R. E. Petit and M. G. Harasewych, 1990
Page 25
intermedia (Cancellaiia) Bellardi, 1841:13, pi. 1, figs.
13-14. Tertiary, Italy.
intermedia (Cancellaiia) Holzl, 1958:258, pi. 21, fig.
13; as var. of C. irochlearis Faujas de Saint Fond.
Miocene, Germany.
interrupta (Cancellaria) Deshayes, 1864:100, pi. 73,
figs. 5-7. Lower Eocene, France.
interstrialis {Cancellaria) von Koenen,-889:130, pi. 11,
figs. 3a-d. Oligocene, Germany.
io {Cancellaria) Dall, 1896:14; 1908:295, pi. 1, fig. 2.
Recent, Panamic-Pacific.
iota {Trigonostoma) Garrard, 1975:23, pi. 3, fig. 3.
Recent, Australia.
irelaniana {Cancellaria) Cooper, 1894:42, pi. 1, fig. 5.
Cretaceous (? Eocene), California, U.S.A.
irregularisuturata {Cancellaria) Sacco, 1894:38; as
"anom." of C, cancellata pleuricosticillata Sacco;
see Appendix, Note 2. Pliocene, Italy.
islacolonis {Cancellaria) Maury, 1917:65, pi. 10, figs.
12, 12a-b. Miocene/Pliocene, Dominican Republic.
Ualica {Cancellaria) D'Ancona, 1872:112, pi. 12, figs.
5a-b, 6a-b. Pliocene, Italy.
iunior {Cancellaria) Bellardi, 1841:pl. 1, fig. 15; as
var. of C. intermedia Bellardi; plate caption only;
see C. junior Bellardi. Miocene, Italy.
iwaotakii {Nipponaphera) Habe, 1961b:431, pi. 24, fig.
22. Recent, Japan.
J
jacksonica {Cancellaria) Cooke, 1926:134, fig. 4. Eo-
cene, Mississippi, U.S.A.
jadisi {Cancellaria) Olsson, 1964:123, pi. 21, fig. 7.
Miocene, Ecuador.
japonica {Cancellaria) E. A. Smith, 1879b:216, pi. 20,
fig. 54. Recent, Japan.
japonica {Cancellaria) 'Lischke". Error for C. japonica
Smith.
jasnini {Cancellaria) Gilbert, 1952b:368, pi. 12, fig. 10.
Miocene, Belgium.
jayana {Cancellaria) Keen, 1958:249, pi. 30, fig. 5.
Recent, Panamic-Pacific.
jipijapana {Cancellaria) Pilsbry & Olsson, 1941:22, pi.
4, figs. 1, 4. Pliocene, Ecuador.
joachimi {Cancellaria) Handmann, 1882:261. Miocene,
Austria.
jocosa {Inglisella) Maxwell, 1988:70, pi. 11, figs. i-k.
Miocene, New Zealand.
joaqiiinensis {Cancellaria) F. M. Anderson, 1905:199,
pi. 15, figs. 46-48. Miocene, California. U.S.A.
jogjacartensis {Cancellaria) K. Martin, 1914:128, pi. 2,
fig. 57. Eocene, Indonesia.
jonkairiana {Cancellaria) Nyst, 1835:29, pi. 5, fig. 28.
Tertiary, Europe, (see C. lajonkairii Nyst, 1853)
jonkeiriana {Cancellaria) 'Nyst' - Nyst, 1845:475; et
al. Error for C. jonkairiana Nyst.
jonkeri {Cancellaria) Koperberg, 1931:65, pi. 2, fig. 18.
Tertiary, Indonesia.
josephinae {Trigonostoma) Janssen, 1984:23, pi. 4, figs.
10-13; pi. 6, fig. 5; as subsp. of T. geslini
(Basterot). Miocene, Netherlands.
jucunda {Cancellaria) Thiele, 1925:201, pi. 22, fig. 21.
Recent, eastern Africa.
jiimala {Admete) Olsson, 1964:127, pi. 22, figs. 3, 3a-c.
Pliocene, Ecuador.
junior {Sveltia) 'Bellardi' - Ferrero Mortara, 1984:176;
variant spelling of iunior, q.v.
junipera {Cancellaria) G. D. Harris, 1895a:65, pi. 6,
fig. 3; as var. of C. panones Harris. Eocene, Texas,
U.S.A.
K
kaiparaensis {Merica) Laws, 1939:496, pi. 65, fig. 50.
Miocene, New Zealand.
kaitarus (Kapuatriton) Beu & Maxwell, 1987:24, pi. 2,
figs, a-b, d-e. Upper Eocene, New Zealand.
karsteni {Cancellaria) F. M. Anderson, 1929:114, pi.
10, figs. 7-9, Miocene, Colombia.
kaunhoweni {Uxia) Cossmann, 1899a:38; n.n. for C.
minima Kaunhowen, 1898, non Reeve, 1856. Creta-
ceous, Germany.
keaseyensis {Sveltella) Hickman, 1980:75, pi. 9, figs.
10-11. Oligocene, Oregon, U.S.A.
keenae {Cancellaria) Addicott, 1970:119, pi. 16, figs.
1-4, 18-19. Miocene, California, U.S.A.
keepingi {Bonellitia) Wrigley, 1935:368, pi. 33, fig. 17.
Middle Eocene, England.
kelseyi {Trichotropis) Dall, 1908:254. Recent, Califor-
nia, U.S.A. [Iphinopsis]
kernensis {Cancellaria) Addicott, 1970:117, pi. 15, figs.
18-21. Miocene, California, U.S.A.
kilburni {Africotriton) Beu & Maxwell, 1987:31, pi. 8,
figs, i-p; pi. 11, figs. m-q. Recent, South Africa.
kimikoae {Cancellaria) Hatai, 1940:115, figs. 1-2. Mio-
cene, Japan.
kobayashii {Mitra) Yokoyama, 1927:173, pi. 47, fig. 5.
Neogene, Japan. [Merica]
kobayasii {Cancellaria) Otuka, 1937:1020. Nomen nu-
dum.
kochiensis {Cancellaria) Katto. 1960:110, pi. 1, fig. 5.
Oligocene, Japan.
koreanica {Cancellaria) Hatai & Kotaka. 1952:83, pi.
7, figs. 3, 5; as subsp. of C. spengleriana Deshayes.
Miocene, Japan.
kroyeri {Trichotropis) Philippi, 1849:175. Recent, North
Pacific. [? Iphinopsis]
kugleri {Cancellaria) Rutsch, 1934:90, pi. 8, figs. 3-4.
Neogene, Venezuela.
kulanda {Zeadmete) Garrard, 1975:44, pi. 3, fig. 15.
Recent, Australia.
kumeroa {Zeadmete) Fleming, 1943:206, pi. 31, fig. 34.
Pliocene/Pleistocene, New Zealand.
kunraedensis {Cancellaria) Kaunhowen, 1898:105, pi.
13, fig. 6. Cretaceous, Belgium.
kurodai {Trigonostoma) Makiyama, 1927:85, pi. 4, fig.
8. Pliocene, Japan.
L
labratula {Cancellaria) von Koenen, 1889:147, pi. 11,
figs. 5a-d, 6a-b. Oligocene, Germany.
labrosa {Cancellaria) Bellardi, 1840:343. Nomen nu-
dum.
Page 26
THE NAUTILUS, Supplement 1
labrosa (Cancellaria) Bellardi, 1841:10. pi. 1, figs. 3-4.
Tertiary, Italy.
lacondamini (Cancellaria) Olsson, 1964:121, pi. 21.
figs. 1, la-c. Miocene, Ecuador.
lactea (Cancellaria) Deshayes, 1830:180. Recent, Aus-
tralia.
lacunosa (Cancellaria) Mutton, 1885:320. Pliocene/
Pleistocene, New Zealand.
laddi (Cancellaria) Petit, 1987:154; n.n. for C. petiti
Ladd, 1982, non Olsson, 1967. Pliocene, Fiji.
laekeniana (Cancellaria) Vincent in Vincent & Lefevre,
1872:58, pi. 2, figs. 2-3. Eocene, Belgium.
laekeniensis - error for laekeniana.
laevescens (Cancellaria) Guppy, 1866:289, pi. 17, fig.
12. Pliocene, Jamaica.
laevicolligens (Cancellaria) Sacco, 1894:43, pi. 3, fig.
3; as var. of C bonellii Bellardi; see Appendix,
Note 2. Miocene, Italy.
laevicolumella (Admete) Sacco, 1894:72, pi. 3, fig. 97;
as var. of A. nassiformis (Seguenza). Miocene, Italy.
laevicosta (Cancellaria) Wood, 1842:538. Nomen nu-
dum.
laevifasciata (Brocchinia) Sacco, 1894:69, pi. 3, fig. 88;
as var. of B. mitraeformis (Brocchi). Pliocene, Italy.
laevigata (Cancellaria) G. B. Sowerby I, 1832b:fig. 24.
Recent, Australia. (?= C lactea Deshayes, 1830)
laevigata (Cancellaria) von Koenen, 1865:472. Oligo-
cene, Europe.
laevilabris (Cancellaria) 'Bon.' - Bellardi, 1840:344;
as var. of C nodulosa Lamarck. Nomen nudum.
laevilabris (Cancellaria) Bellardi, 1841:20; ex Bonelli
MS, as var. of C nodulosa Lamarck. Tertiary, Italy.
laevior (Admete) Leche, 1878:48; n.n. for "C viridula
(Fabr.)" Middendorff, 1849:pl. 9, figs. 1-2; as var.
of A. viridula (Fabric! us). Recent, Arctic Ocean.
laevior (Cancellaria) Sacco, 1894:30; n.n. for C. bar-
jonae Pereira da Costa, 1867:pl. 25, fig. 13 (only);
as var. of C. barjonae Pereira da Costa; see
Appendix, Note 2. Miocene, Portugal.
laevior (Cancellaria) Sacco, 1894:12, pi. 1, fig. 29; as
var. of C. taurolaevigatum Sacco; see Appendix,
Note 2. Miocene, Italy.
laevis (Paladmete) Sohl, 1964a:273, pi. 45, figs. 35, 41,
42, 46-48. Cretaceous, Mississippi, U.S.A.
laeviuscula (Cancellaria) J. Sowerby, 1822:84, pi. 361,
fig. 1. Lower Eocene, England.
lajonkairei (Cancellaria) 'Nyst' - Cossmann, 1899a:21;
el al. Error for C. lajonkairii Nyst.
lajonkairii (Cancellaria) Nyst, 1853:592. Emendation
of C. jonkairiana Nyst, 1835.
lamberti (Cancellaria) Souverbie in Souverbie & Mon-
trouzier, 1870:428, pi. 14, fig. 2. Recent, New
Caledonia.
lamellosa (Cancellaria) Hinds, 1843:49; 1844b:43, pi.
12, figs. 15-16. Recent, Indo-Pacific. (?= Scalptia
nassa (Gmelin, 1791))
lamyi (Cancellaria) Koperberg, 1931:69, pi. 2, fig. 20.
Tertiary, Indonesia.
lanceolata (Cancellaria) Aldrich, 1897b:27, text-fig.
Eocene, Alabama, U.S.A.
lanceolata (Ranella) Menke, 1828:87. Recent, Carib-
bean. [Tritonoharpa]
landesi (Cancellaria) Van Winkle, 1918:91, pi. 7, fig.
17. Oligocene, Washington, U.S.A.
laqua (Cancellaria) Mansfield, 1935:26, pi. 2. fig. 5.
Miocene, Florida, U.S.A.
larkinii (Cancellaria) Nelson, 1870:192, pi. 6, fig. 7.
Miocene, Peru.
laseroni (Arizelostoma) Iredale, 1936:318, pi. 24, fig.
9. Recent, Australia.
lateapertitm (Trigonostoma) Peyrot, 1928:240, pi. 13,
fig. 31; as var. of T. geslini (Basterot). Miocene,
France.
latecostata (Admete) Traub, 1938:94, pi. 8, figs. lOa-b.
Tertiary, Europe.
latecostata (Cancellaria) 'Lobbecke' - Lobbecke,
1887b:40. Error for C. laticosta Lobbecke, 1881.
latefasciata (Cancellaria) Sacco, 1894:64, pi. 3, fig. 70;
as var. of C. tribulus (Brocchi). Pliocene, Italy.
laterinensis (Inglisella) Maxwell, 1988:70, pi. 11, figs.
e-g. Miocene, New Zealand.
latesulcata (Cancellaria) von Koenen, 1885:8, pi. 1,
figs. 2a-f. Paleocene, Denmark.
laticosta (Cancellaria) Lobbecke, 1881:12, pi. 2, figs.
7-9; as var. of C. reeveana Crosse. Recent, Indo-
Pacific.
laticostata (Cancellaria) Tenison- Woods, 1880:17, pi.
2, fig. 8. Miocene, Australia, (see C. platypleura
Tate)
laticostata (Cancellaria) 'Lobbecke' - Lobbecke, 1881:13.
Error for C. laticosta Lobbecke, 1881.
laticostata (Cancellaria) Sacco, 1894:31, pi. 2, fig. 37;
as var. of C. doderleini Mayer; see Appendix, Note
2. Miocene, Italy.
latilabris (Cancellaria) 'Bon.' - Michelotti, 1838:396.
Nomen nudum.
lattorfensis (Bonellitia) Wrigley, 1935:369, pi. 33, fig.
19; n.n. for C. ovata von Koenen, 1889, non
Sowerby, 1832. Oligocene, Germany.
latum (Tritonium) Dareste de la Chavanne, 1910:22, pi.
4, fig. 4; as var. of T. turriculatum Deshayes.
Eocene, Algeria. [Plesiotriton]
laurensii (Cancellaria) Grateloup, 1832:341. Tertiary,
France.
lauta (Cancellaria) 'Tate' - Tate & Dennant, 1893:221.
Nomen nudum.
lavelana (Cancellaria) H. K. Hodson in Hodson &
Hodson, 1931:44, pi. 24, fig. 12. Miocene, Vene-
zuela.
lavescens (Cancellaria) 'Guppy' - Marks, 1949:460.
Error for C. laevescens Guppy.
laviae (Cancellaria) 'Hoffmann' - Brugnone, 1880:103;
as La Viae. Nomen nudum.
leal (Cancellaria) Crosse, 1861:255; n.n. for C. tessel-
lata Lea, 1833, non Sowerby, 1832. Eocene, south-
eastern U.S.A. (?= C. alveata Conrad, 1833)
lebrosus (Fusus) Bellardi, 1839:31. Nomen nudum. [=
C. labrosa Bellardi, 1841]
R. E. Petit and M. G. Harasewych, 1990
Page 27
leioderma (Mataxa) Sohl, 1964b:382, pi. 56, figs. 2-3.
Cretaceous, Mississippi, U.S.A.
leoiia (Coliihraria) Mansfield, 1937:610, pi. 85, fig. 3.
Pliocene, Florida, U.S.A. [?= Plesiolriton lanceolata
(Menke, 1828)]
leonensis (Cancellaria) Mansfield, 1930:46, pi. 3, fig.
12; as subsp. of C. reticulata (Linne). Miocene,
Florida, U.S.A.
leopoldinae (Cancellaria) Toumouer in Bouille, 1876:90,
pi. 1, fig. 7. Miocene, France.
lesbarritzensis (Sveltia) Vergneau-Saubade, 1968:206,
text-fig. Oligocene, France.
leuzingeri (Cancellaria) Rutsch, 1934:89, pi. 7, figs.
10-11; pi. 8, figs. 1-2, 5; as subsp. of C. reticulata
(Linne). Neogene, Venezuela.
levis (Merica) Peyrot, 1928:208, pi. 12, fig. 34; as var.
of M. contorta (Basterot). Miocene, France.
lichana - error for lickana.
lickana (Cancellaria) Anderson & Martin, 1914:84, pi.
8, figs. 6a-d. Miocene, California, U.S.A.
Ugeriana (Cancellaria) Gilbert, 1952b:367, pi. 11, fig.
lib; as forma of C. contorta Basterot. Miocene,
France.
ligustica (Cancellaria) Sacco, 1894:6, pi. 1, fig. 5; as
var. of C. umbilicaris (Brocchi); see Appendix,
Note 2. Pliocene, Italy.
lima (Cancellaria) Hoeninghaus, 1831:145. Nomen nu-
dum.
lima (Cancellaria) von Koenen, 1889:126, pi. 11, figs.
la-c. Oligocene, Germany.
limata (Cancellaria) Yokoyama, 1928:342, pi. 66, fig.
12. Pliocene, Japan.
limnaeaeformis (Admete) E. A. Smith, 1877:6, pi. 9,
fig. 4; 1879a: 172, pi. 9, fig. 4. Recent, Kerguelen
Island, [not a cancellariid; ? Toledonia]
limnaeiformis (Cancellaria) 'Smith' - Tryon, 1885:85.
Error for C. limnaeaeformis (Smith).
lindae (Aphera) Petuch, 1987:109, pi. 13, fig. 11.
Recent, Barbados.
lindeni (Trigonostoma) Janssen, 1984:21, pi. 4, figs.
1-4; pi. 6, fig. 6. Miocene, Netherlands.
lipara (Cancellaria) Woodring, 1951:76, pi. 16, figs.
13-14. Tertiary, California, U.S.A.
lipara (Cancellaria) Woodring, 1970:337, pi. 52, figs.
7-8; as subsp. of C. epistomifera Guppy. Pliocene,
Panama. (= C. sathra Woodring, 1973)
lirata (Cancellaria) Conrad, 1865a:32. Nomen nudum.
lirata (Cancellaria) Conrad, 1865b: 145, pi. 11. fig. 3.
Eocene, Texas, U.S.A. (?= C. gemmata Conrad,
1833)
lirata (Cancellaria) 'Brocchi'- de Stefani & Pantanelli,
1878:118. Error for C. lyrata (Brocchi).
lirisculpta (Admete) Cossmann, 1902:57, pi. 6, fig. 25;
ex deBoury MS. Middle Eocene, France.
lisberi (Cancellaria) 'Risso' - Sacco, 1894:27. Error for
C. listeri Risso.
lischkei (Cancellaria) Yokoyama, 1926a:264, pi. 32,
figs. 16-17. Pliocene, Japan.
listeri (Cancellaria) Risso, 1826:188. Tertiary, Europe.
littorinaeformis (Cancellaria) 'Sowerby' - G. B. Sow-
erby II, 1849:450; et al. Error for C. littoriniformi.s
G. B. Sowerby I.
littoriniformis (Cancellaria) G. B. Sowerby I, 1832b:fig.
18. Recent, Sri Lanka.
lividorupis (Turefuia) Beu & Maxwell, 1987:19, text-
fig. 2K; pi. 2, figs, c, f-j, m. Miocene, New
Zealand.
lloydi (Fasciolaria) Stanton, 1920:42, pi. 8, figs, lla-b.
Paleocene, North Dakota, U.S.A. [Cancellariidae]
lobata (Cancellaria) Swainson, 1840:305, fig. 72f. ?Re-
cent, locality unknown. (?= C. tuberculosa Sow-
erby, 1832)
loebbeckei (Cancellaria) - Kuroda & Habe, 1971:202
(English). Error for C. laticosta Lobbecke.
longispirata (Cancellaria) Yokoyama, 1926a:265, pi.
32, fig. 10. Pliocene, Japan.
longojuvenis (Cancellaria) Sacco, 1894:22, pi. 2, fig.
3; as var. of C. acutangula Faujas de Saint Fond;
see Appendix, Note 2. Miocene, Italy.
longonassoides (Cancellaria) Sacco, 1894:52, pi. 3, fig.
34; as var. of C. deshayesiana Desm. [sic]; see
Appendix, Note 2. Miocene, Italy.
losquemadica (Cancellaria) Maury, 1917:66, pi. 10, fig.
13. Miocene, Dominican Republic.
luffa (Admete) Olsson, 1929:27, pi. 8, figs. 3-4. Eocene,
Peru.
lunata (Cancellaria) Conrad, 1830:222, pi. 9, fig. 4.
Miocene, Maryland, U.S.A.
luscinia (Cancellaria) Melvill & Standen, 1903:319, pi.
23, figs. 14-15. Recent, Arabian Sea.
lyra (Cancellaria) - Deshayes, 1833:30. Error for C
lyrata (Brocchi).
lyrata (Voluia) Brocchi, 1814:311. pi. 3, fig. 6. Plio-
cene, Italy. [Sveltia]
lyrata (Cancellaria) Adams & Reeve, 1850:42, pi. 10,
fig. 4. Recent, ? Panamic-Pacific. (?= C. funiculata
Hinds, 1843)
M
macconkeyi (Scalptia) Jousseaume. 1894:201. Recent,
Aden. (?= Scalptia hystrix (Reeve, 1856))
maccoyi (Cancellaria) Pritchard & Gatliff, 1899:182, pi.
20, fig. 6. Recent, Australia. (?= C. purpuriformis
Kiener, 1841)
macnairyensis (Cancellaria) Sohl, 1964a:268, pi. 44,
figs. 1-2. Cretaceous, Tennessee, U.S.A.
macneili (Cancellaria) Mansfield, 1937:609. pi. 85, figs.
1. 4. Miocene. Rorida. U.S.A.
macrospira (Cancellaria) Adams & Reeve. 1850:41. pi.
10, fig. 2. Recent, Indo-Pacific.
macrospiratoides (Fusiaphera) Habe, 1961b:433, pi. 23,
fig. 10; pi. 24, fig. 10. Recent, Japan.
macrostoma (Cancellaria) du Bois de Montpereux,
1831:32. pi. 3. figs. 36-37. [not a cancellariid]
macrostoma (Cancellaria) Doderlein. 1863:21. Nomen
nudum.
macrostoma (Cancellaria) 'Eichwald' - Sacco. 1894:25.
? Error for C. macrostoma du Bois de Montpereux.
magellanica (Admete) Strebel, 1905:594, pi. 22, figs.
29a-d. Recent, Magellanic Province.
Page 28
THE NAUTILUS, Supplement 1
magloirei (Plesiocerithium) 'Melleville' - Cossmann,
1889:232; et al. Error for P. maglorii (Melleville).
maglorii (Cancellaria) Melleville, 1843:112, pi. 9, figs.
1-3. Lower Eocene, France.
magnoturrita (Cancellaria) Sacco, 1894:57, pi. 3, fig.
50; as var. of C. dertovaricosa Sacco; see Appen-
dix, Note 2. Miocene, Italy.
maior (Cancellaria) Bellardi, 1841:19, pi. 2, figs. 1-2:
as var. of C. nodiilosa Lamarck. Pliocene, Italy.
major (Cancellaria) "Bellardi" - Sacco, 1894:28; as var.
of C. hirta (Br.). Variant spelling of C. maior
Bellardi.
major (Cancellaria) Grateloup, 1847:2, pi. 1, fig. 29;
as var. of C. ditfourii Grateloup; see Appendix,
Note 1. Tertiary, France.
major (Cancellaria) von Ihering, 1899:35, pi. 2, fig.
10; as var. of C. gracilis von Ihering. Tertiary,
Argentina.
major {Cancellaria) Pallary, 1900:259; as var. of C.
scahriiiscula (Linne). Nomen nudum.
major (Oamaruia) Marwick, 1965:40, pi. 11, figs. 12,
15. Pliocene, New Zealand.
malachitensis (Cancellaria) Stanton, 1893:158, pi. 33,
figs. 6-7. Cretaceous, Colorado, U.S.A.
malaisei (Cancellaria) Briart & Comet, 1877:15, pi. 14,
figs. 2a-c; as malaisi on p. 15, but as malaisei on
p. 16 and in later publications by the authors, et al.
Paleocene, Belgium.
malaisi (Cancellaria) 'Briart & Cornet' - Brian &
Comet, 1877:15. Error for C. malaisei Briart &
Comet.
maldonadoi (Cancellaria) Olsson, 1964:122, pi. 21,
figs. 5, 5a. Miocene, Ecuador.
mangelioides (Cancellaria) Reeve, 1856:pl. 15, figs.
69a-b. Recent, Indo-Pacific.
mangyschlakica (Cancellaria) Bajarunas, 1912:46, pi.
3, figs. 5-7. Upper Oligocene, U.S.S.R.
maorium (Admete) Marshall & Murdoch, 1921:82, pi.
18, figs. 7-8. Miocene, New Zealand.
margaritata (Cancellaria) Vinassa de Regny, 1896:270,
pi. 18, figs. 16-17. Tertiary, Italy.
mariae (Bivetia) 'Jousseaume' - Marks, 1949:456. Error
for B. mariei Jousseaume.
marieana (Cancellaria) Aldrich, 1897a: 179, pi. 1, fig.
6. Lower Eocene, Alabama, U.S.A.
mariei (Bivelia) Jousseaume, 1887a:163, fig. 1. Recent,
locality unknown. (?= C. indentata Sowerby, 1832)
marksi (Cancellaria) Olsson, 1964:125, pi. 37, fig. 6.
Pliocene, Ecuador.
marshalli (Cerittudea [sic\) Vignal in Cossmann,
1921:181; n.n. for Cerithidea minuta Marshall,
1919, non Gabb, 1873. Eocene, New Zealand.
[Brocchinia]
marshalli (Latirus) Finlay, 1924a: 102; unnecessary n.n.
for Latirus duhius Marshall, 1919, q.v.
marshalli (U.xia) Allan, 1926:342, pi. 77, figs. 3a-b.
Eocene, New Zealand.
marthae (Trigonostoma) Olsson, 1967:25, pi. 7, fig. 3.
Pliocene, Florida, U.S.A.
martini (Bivetia) Cossmann, 1899a: 10; n.n. for C.
neglecta Martin, 1895, non Michelotti, 1861. Terti-
ary, Indonesia.
martiniana (Cancellaria) Noetling, 1901:332, pi. 22,
figs, lla-d, 12, 12a, 13a-c. Miocene, Burma, [not
a cancellariid]
marwicki (Waipaoa) Dell, 1956:112, fig. 113. Recent,
New Zealand.
marylandica (Cancellaria) G. C. Martin, 1904:165, pi.
43, fig. 8. Miocene, Maryland, U.S.A.
marylandica (Marianarona) Petuch, 1988:27. pi. 5, figs.
13-14. Miocene, Maryland, U.S.A.
marysvillensis (Cancellaria) Dickerson, 1916:442. No-
men nudum.
masferreri (Cancellaria) Almera & Bofill, 1884:30; as
var. of C. gradata Homes. Tertiary, Spain.
matsoni (Cancellaria) Stephenson, 1941:362, pi. 69,
figs. 4-5. Cretaceous, Texas, U.S.A.
mauryae (Cancellaria) Olsson, 1922:82, pi. 6, fig. 5.
Miocene, Dominican Republic.
mediangulata (Cancellaria) Sacco, 1894:36; n.n. for
"C uniangulata Deshayes" H5mes, 1853:pl. 32,
fig. 2; see Appendix, Note 2. Miocene, Austria.
medicinensis (Petersia) Cragin, 1894:11. Cretaceous,
Kansas, U.S.A. [? Cancellariidae]
medinae (Cancellaria) Philippi, 1887:63, pi. 7, fig. 4.
Miocene, Chile.
megapex (Bonellitia) Yokes, 1939:129, pi. 18. figs. 4-5.
Eocene, California, U.S.A.
megastoma (Cancellaria) Peyrot, 1928:206, pi. 12, fig.
7; as var. of C. barjonae Pereira da Costa. Miocene,
France.
melanostoma (Cancellaria) G. B. Sowerby H, 1849a: 137;
1849b:447, pi. 95, fig. 78. Recent, Indian Ocean.
menadensis (Cancellaria) Schepman, 1907:164, pi. 10,
figs. 5, 5a. Post-Tertiary, Indonesia.
mercadoi (Scatptia) Old, 1968:286, pi. 43, figs. 1-3,
text-figs. 1-2. Recent, Philippines.
mericana (Cancellaria) - Cossmann, 1899a:30. ? Error
for C. marieana Aldrich.
mericoides (Cancellaria) Sacco, 1894:52, pi. 3, fig. 35;
as var. of C. deshayesiana Desm. [sic\: see Appen-
dix, Note 2. Miocene, Italy.
meridionalis (Cancellaria) Woods, 1906:326, pi. 40,
figs. 10-11. Cretaceous, South Africa.
metuloides (Cancellaria) Olsson, 1964:119, pi. 37, figs.
7. 7a. Miocene, Ecuador.
michelinii (Cancellaria) Bellardi, 1841:37, pi. 4, figs.
5-6. Miocene, Italy.
micra (Cancellaria) Tate, 1889:158, pi. 10, fig. 8.
Eocene, Australia.
microscopica (Cancellaria) Dall, 1889a:131. Recent,
Caribbean.
microsoma (Cancellaria) Dall, 1908:296, pi. II, fig.
10. Recent, westem Mexico.
microstoma (Cancellaria) 'Dubois' - Bellardi. 1841:3.
Error for C. macrostoma du Bois de Montpereux.
microstoma (Cancellaria) 'Chariesworth MS' - Morris,
1854:239; Newton, 1891:171. Nomen nudum.
R. E. Petit and M. G. Harasewych, 1990
Page 29
microstoma (Cancellarla) Newton, 1895:327, pi. 22,
tigs. 3-4; ex Charlesworth MS. Eocene, England.
middendorffiana (Admete) Dull, 1885:524; n.n. for
"Admeie viridula Fabricius" Middendorff, 1849:pl.
9, figs. 13-14. Recent, Alaska, U.S.A. (= Admete
laevior Leche, 1878)
mihelici (Cancelaria [sic]) Suklje, 1929:30, pi. 4, figs.
3a-b. Miocene, Yugoslavia.
milleri {Thgonostoma) Burch, 1949:3, text-figs. Recent,
Panamic-Pacific.
milletii (Cancellaria) Deshayes, 1830:181. Tertiary,
France.
minhuensis (Cassidarid) Noetling, 1895:28, pi. 6, figs.
4, 4a-b. Oligocene/Miocene, India. [Loxotaphrus]
minima (Cancellaria) Reeve, 1856:pl. 17, figs. 77a-b.
Recent, Canary Islands.
minima (Cancellaria) Geinitz, 1874a:265, pi. 59, figs.
2a-b. Cretaceous, Germany.
minima (Cancellaria) Kaunhowen, 1898:104, pi. 7, figs.
9, 9a. Cretaceous, Belgium. (= Uxia kaunhoweni
Cossmann, 1899)
minor (Cancellaria) Grateloup, 1847:2, pi. 1, fig. 26;
as var. of C. diifourii Grateloup; see Appendix,
Note 1. Tertiary, France.
minor (Cancellaria) Bronn, 1848:210; n. n. for "K
piscatoria (Gm.)" Brocchi, 1814:pl. 3, fig. 12; as
var. of C. hirta (Br.). Pliocene, Italy. (= C. brocchii
Crosse, 1861)
minor (Cancellaria) Beyrich, 1856:557; as var. of C.
evnlsa (Sol.). Tertiary, Europe.
minor (Cancellaria) Almera & Bofill, 1884:56, pi. E,
figs. 31-32; as var. of C. cancellata (L.). Tertiary,
Spain.
minor (Cancellaria) G. B. Sowerby III, 1889:568; as
var. of C fiisca Sowerby. Recent, Hong Kong.
minor (Cancellaria) Almera & Bofill, 1892:12; as var.
of C. hirta (Br.). Nomen nudum.
minor (Cancellaria) Pallary, 1900:259; as var. of C.
scabriuscula (L.). Nomen nudum.
minor (Cancellaria) Pallary, 1920:27, fig. 15; as var.
of C. piscatoria (Gmelin). Recent, Morocco.
minuta (Cancellaria) Nyst, 1845:482, pi. 12, figs.
23a-c. Tertiary, Belgium.
minuta (Cancellaria) Brown, 1849:253, pi. 33*, fig. 73.
? Tertiary, ? British Isles. Nomen dubium.
minuta (Cancellaria) Braun, 1851:1131. Nomen nudum.
minuta (Cancellaria) Sandberger, 1859:pl. 15, figs. 9,
9a; 1862:259; ex Braun. Tertiary, Germany. (= C.
moguntina Crosse, 1861)
minuta (Cancellaria) 'Harris' - Kennedy, 1895:130.
Nomen nudum.
minuta (Cerithidea) Marshall, 1919:226, pi. 15, fig. 11.
Eocene, New Zealand. [= Cerittudea marshalH
Vignal, q.v.]
miocaenica (Cancellaria) 'Doderl.' - Cossmann,
1899a:21. Error for C miocenica Doderlein.
miocaenica (Cancellaria) Cossmann, 1913a:54, pi. 4,
figs. 11-12. Miocene, Panama.
miocenica (Cancellaria) Doderlein, 1863:21; as var. of
C lyrata (Br.). Nomen nudum.
miocenica (Cancellaria) Doderlein, 1863:21; as var. of
C. varicosa (Br.). Nomen nudum.
miocenica (Cancellaria) Doderlein, 1863:21; as var. of
C cassidea (Br.). Nomen nudum.
miocenica (Cancellaria) Sacco, 1894:61, pi. 3, fig. 62;
as var. of C. lyrata (Brocchi); ex Doderlein; see
Appendix, Note 2. Miocene, Italy.
miocenica (Cancellaria) Sacco, 1894:56, pi. 3, figs.
47a-b; as var. of C. varicosa (Brocchi); ex Doder-
lein; see Appendix, Note 2. Miocene, Italy.
miocenica (Cancellaria) Sacco, 1894:8, pi. 1, figs.
15a-b; as miocenicum; ex Doderlein; see Appendix,
Note 2. Miocene, Italy.
miocenica (Zeadmete) Finlay, I930a:78. Miocene, New
Zealand.
mioconnectens (Cancellaria) Sacco, 1894:58, pi. 3, fig.
55; as var. of C. taurinia Bellardi; see Appendix,
Note 2. Miocene, Italy.
miocrassa (Cancellaria) Sacco, 1894:7; n.n. for "C.
ampullacea (Br.)" Grateloup, 1847:pl. 25, figs. 28,
32; see Appendix, Note 2. Tertiary, France.
miohirta (Cancellaria) Sacco, 1894:27; n.n. for "C.
hirta (Br.)" Almera & Bofill, 1884:pl. B, figs. 6-7;
see Appendix, Note 2. Tertiary, Spain.
miolonga (Cancellaria) Sacco, 1894:46, pi. 3, fig. 17;
as var. of C. evulsa (Solander); see Appendix, Note
2. Miocene, Italy.
miolyrata (Cancellaria) Sacco, 1894:62; n.n. for "C.
lyrata (Br.)" Beyrich, 1857:pl. 27, figs. 7-8; see
Appendix, Note 2. Tertiary, Germany.
mioparva (Cancellaria) Sacco, 1894:33; n.n. for "C.
calcarata (Br.)" Beyrich, 1857:pl. 28, fig. 3; see
Appendix, Note 2. Tertiary, Germany.
mioquadrata (Cancellaria) Sacco, 1894:66; n.n for "C.
laurensii Grat." Hoemes & Auinger, 1890:pl. 33,
figs. 1-2; see Appendix, Note 2. Miocene, Austria.
mioscalaroides (Cancellaria) Sacco, 1894:54; n.n. for
"C. scalaroides Wood" Beyrich, 1857:pl. 27, fig.
5; see Appendix, Note 2. Tertiary, Germany.
mirabilis (Plesiotriton) Beu & Maxwell, 1987:27, pi.
24, figs, a-1, q-r; text-figs. 2E-F. Recent, New
Britain.
mississippiensis (Cancellaria) Conrad, 1848a:287;
1848b:118, pi. 11, fig. 38. Oligocene, Mississippi,
U.S.A.
mistense (Trigonostoma) Janssen, 1984:16, pi. 3, figs.
4a-b. Miocene, Netherlands.
mitraeforme (Buccinum) Pusch in Andrzejowski, 1830:94,
pi. 4, figs. la-b. Tertiary, Poland. [= Cancellaria
inermis Pusch, 1837, q.v.]
mitraeformis (Voluta) Brocchi, 1814:645, pi. 15, fig.
13. Pliocene, Italy, [non Voluta mitraeformis La-
marck, 1811; ?= Brocchinia pan-ula tauroparva
Sacco, 1894]
mitraeformis (Cancellaria) "Eichwald' - Crosse,
1861:251. Error for C. mitraeformis (Pusch).
mitraeformis (Cancellaria) 'Sowerby' - Bellardi, 1841:9;
et al. Error for C mitriformis Sowerby.
mitriformis (Cancellaria) G. B. Sowerby I, 1832a:51;
1832b:fig. 14. Recent, Panamic-Pacific.
Page 30
THE NAUTILUS. Supplement 1
mitrodita (Cancellaria) Gardner, 1937:369, pi. 44, fig.
11. Miocene, Florida, U.S.A.
mUroides {Cancellaria) Fischer von Waldheim, 1807:173.
Fossil, locality unknown. Nonieti inquirendum.
mudesta (Cancellaria) Carpenter, 1864:628, 660. Re-
cent, northwestern coast of U.S.A.
modeslina {Cancellaria) Tate, 1889:157, pi. 9, fig. 4.
Miocene, Australia.
moguntina {Cancellaria) Crosse, 1861:252; n.n. for C.
minuta Braun, non Nyst, with ref. to Sandberger,
[1859]:pl. 15, fig. 9. Tertiary, Germany.
moharrami {Cancellaria) Abbass, 1972:58, pi. 4, fig.
15. Paleocene, Saudi Arabia.
montrouzieri {Cancellaria) Souverbie in Souverbie &
Montrouzier, 1863:163, pi. 5, fig. 8. Recent, New
Caledonia. (?= C. contabulata Sowerby, 1832)
montserratensis {Cancellaria) Maury, 1925a: 194, pi
35, figs. 6, 8. Miocene, Trinidad.
moorei (Cancellaria) Guppy, 1866:289, pi. 17, fig. 7
Pliocene, Jamaica.
moravica (Cancellaria) Oppenheim, 1922:80, pi. 5, figs
15, 15a-b. Tertiary, Czechoslovakia.
morgani (Sveltia) Cossmann, 1903a: 107, pi. 3, figs
3-4. Pliocene, India.
mourloni (Cancellaria) Briart & Cornet, 1877:12, pi
14, figs. 3a-c. Paleocene, Belgium.
mucronatum (Trigonostoma) Peyrot, 1928:256, pi. 14
figs. 9, 33. Miocene, France.
miilticostata (Cancellaria) Bellardi & Michelotti,
1840:147; as var. of C. uniangulata Deshayes.
Tertiary, Italy.
miilticostata (Cancellaria) Bellardi, 1840:344; as var.
of C. uniangulata Deshayes. Nomen nudum.
miilticostata (Cancellaria) Bellardi. 1841:17, pi. 2, figs.
5-6; as var. of C. uniangulata Deshayes. Tertiary,
Italy.
miilticostata (Cancellaria) Bellardi, 1841:8; as var. of
C. uinpullacca (Br.). Nomcn nudum.
miilticostata (Cancellaria) Bellardi, 1841:20, pi. 2, figs.
13-14; as var. of C. nodulosa Lamarck. Tertiary,
Italy.
miilticostata (Cancellaria) Bellardi, 1841:31, pi. 4, figs.
19-20; as var. of C. huccinula Basterot [sic].
Tertiary, Italy.
miilticostatum (Triton) Favre, 1869:89, pi. 10, fig. 15.
Cretaceous, France. [Plesiolriton]
miiltienensis (Cancellaria) Morlet, 1885b: 196; n.n. for
C. hezanconi Morlet, 1885, non de Raincourt, 1884.
Middle Eocene, France.
miiltiensis (Cancellaria) 'Morlet' - Cossmann, 1899:38.
I'jTor for C. multienensis Morlet.
multilinea (Cancellaria) 'Edwards MS' - Newton,
1891:171. Nomen nudum.
miiltilineata (Bonellitia) Wrigley, 1935:370, pi. 33, figs.
21-22. Middle Eocene, England.
miiltilira (Mata.xa) Stephenson, 1941:366, pi. 70, figs.
6-7; as subsp. of M. valida Stephenson. Cretaceous,
Texas, U.S.A.
miiltinodiilatus (Africotriton) Beu & Maxwell, 1987:32,
pi. 10. figs. a-s. Recent, South Africa.
multiplicata (Cancellaria) I. Lea, 1833:139, pi. 5, fig.
135. Eocene, Alabama, U.S.A.
multiplicata (Cancellaria) Lesson, 1841a:37. Recent, ?
Panamic-Pacific. (?= C. chrysostoma Sowerby, 1832)
multiplicis (Cancellaria) Newton, 1922:42, pi. 3, fig.
26. Eocene, Nigeria.
multistriata (Cancellaria) Bellardi, 1840:344; as var. of
C. taurinia Bellardi. Nomen nudum.
multistriata (Cancellaria) Beyrich, 1856:567, pi. 26.
figs. 6a-d. Oligocene, Europe.
multistriata (Sveltia) Ravn, 1939:87, pi. 3, figs. 23a-b,
24a-b. Paleocene, Denmark.
munida (Waipaoa) Ponder, 1968:46, pi. 4, fig. 57.
Recent, New Zealand. (?= Dellina aoteana (Dell,
1956))
muratana (Cancellaria) Nomura & Onisi, 1940:185, pi.
17, fig. 14. Neogene, Japan.
murayamai (Cancellaria) Yokoyama, 1926b:384, pi.
44, fig. 3. Pliocene, Japan.
muricata (Cancellaria) Risso, 1826:188. Tertiary, Europe.
miiricata (Cancellaria) Wood, 1847:354. Nomen nu-
dum.
mutica (Cancellaria) 'Basterot' - Grateloup, 1832:338.
Nomen nudum.
mutinensis (Cancellaria) Foresti, 1884:302, pi. 1, figs.
la-c. Tertiary, Italy.
mutsuana (Cancellaria) Hatai, Masuda & Suzuki.
1961:29, pi. 4, figs, la-b, 2a-b. Pliocene, Japan.
myrayamai (Cancellaria) 'Vok.' - Zoological Record
for 1930, 67(8):72. Error for C. murayamai
Yokoyama.
N
nakayamai (Neadmete) Habe, 1961a:73, Appendix p.
29, pi. 36, fig. 3. Recent, Japan.
namnetica (Cancellaria) Vasseur, 1881:249. Nomen
nudum.
nana (Cancellaria) Deshayes, 1864:106, pi. 73, figs.
11-12. Eocene, France.
nana (Cancellaria) 'Doderlein' - Davoli. 1980:228.
Nomen nudum.
nana (Trigonostoma) Ozaki, 1956:2, pi. 1, fig. 5.
Miocene, Japan.
nancellaria (Cancellaria) Woodring, 1970:341, pi. 53,
figs. 3-4. Miocene/Pliocene, Panama.
nanggulanensis (Cancellaria) K. Martin, 1914:126, pi.
2, fig. 55. Tertiary, Indonesia.
narica (U.xia) Vredenburg, 1925:99, pi. 7, fig. 7.
Tertiary, India.
naroniformis (Uxia) Finlay, 1930a:80. Miocene, New
Zealand.
nassa (Voluta) Gmelin, 1791:3464. Recent, Indo-Pacific.
[Scalptia]
nassaeformis (Cancellaria) 'Wood MS" - Newton,
1891:171. Nomen nudum.
nassaeformis (Uxia) Wrigley, 1925:246, fig. 12. Eo-
cene, Enaland.
R. E. Petit and M. G. Harasewych, 1990
Page 31
nassaeformis (Egerea) Noszky, 1936:73, pi. 5, fig. 12;
as van of E. collectiva Gabor. Oligocene, Hungary.
nassariformis {Cancellaria) Covacevich & Frassinetti,
1986:49, pi. 2, figs. 4a-c, 5a-c, text-fig. 9. Miocene,
Chile.
nassiformis {Cancellaria) Lesson, 1842:204. Recent,
western Mexico. [?= Nassariiis corpiilentus (C. B.
Adams, 1852)]
nassiformis (Cancellaria) Seguenza, 1880:110, pi. 11,
fig. 28. Tertiary, Italy. (?= C. dregeri Hoemes &
Auinger, 1890)
nassoides {Cancellaria) von Koenen, 1889:149, pi. 12,
figs. 13a-c. Oligocene, Germany.
nassoides {Cancellaria) Schepman, 1911:263, pi. 18,
fig. 9. Recent, Indonesia. (= Admetula garrardi
Petit, 1974)
nasuta {Antepepta) Finlay & Marwick, 1937:83, pi. 11,
fig. 1. Paleocene, New Zealand.
nausorensis {Neadmete) Ladd, 1982:58, pi. 14, figs.
19-22. Pliocene, Fiji.
neavolutella {Cancellaria) Noetling, 1901:328, pi. 22,
figs. 7a-b, 8, 8a. Miocene, Burma. [?= Rimella
javana K. Martin, 1879]
neglecta {Cancellaria) Michelotti, 1861:104, pi. 11,
figs. 9-10. Tertiary, Italy.
neglecta {Cancellaria) K. Martin, 1895:47, pi. 7, fig.
112; n.n. for Triton buccinoides K. Martin, 1880).
Tertiary, Indonesia. (= Bivetia martini Cossmann,
1899)
neritoidea {Cancellaria) G. C. Martin, 1904:168, pi. 43,
figs. 13a-b. Miocene, Maryland, U.S.A.
neudorfensis {Cancellaria) Oppenheim, 1922:82, pi. 6
[sic; error for pi. 5], figs. 13, 13a-b. Tertiary,
Czechoslovakia.
neugeboreni {Cancellaria) M. Homes, 1856:680, pi. 52,
figs. 6a-b. Miocene, Austria.
neuvillei {Bivetia) Peyrot, 1928:203, pi. 12, figs. 35-36.
Miocene, France.
nevadensis {Cancellaria) Anderson & Martin, 1914:85,
pi. 8, figs. 5a-d. Miocene, California, U.S.A.
newhallensis {Cancellaria) Carson, 1926:56, pi. 3, fig.
3. Pliocene, California, U.S.A.
nitens {Cancellaria) Beyrich, 1856:561, pi. 27, figs.
la-c. Oligocene, Germany.
nitida {Cancellaria) A. Adams, 1855:123. Recent, Indo-
Pacific.
nitida {Cancellaria) Reeve, 1856:pl. 17, figs. 78a-b; ex
Adams MS. Recent, ? Indo-Pacific.
nitida {Cancellaria) von Koenen, 1889:100, pi. 12, figs.
lla-c, 12a-c. Oligocene, Germany.
nitida {Scalptia) 'Hinds' - Jousseaume, 1887b:214.
Error for S. nitida (Adams).
nUidula {Valuta) Muller, 1851:41, pi. 5, figs. 25a-b.
Cretacous, Germany. [Cancellariidae]
nodigera {Cancellaria) 'Edwards MS' - Newton,
1891:172. Nomen nudum.
nodosa (Admete) Verrill & Smith in Verrill, 1885:419,
pi. 44, fig. 9. Recent, Delaware Bay, U.S.A.
nodosissima {Cancellaria) Sacco, 1894:47, pi. 3, fig.
20; as var. of C. evul.sa (Solander); see Appendix,
Note 2. Oligocene, Italy.
nodosivaricosa {Agatrix) Petuch, 1979:11, figs. 26-27.
Recent, Philippines.
nodulatus {Epidromus) Tate, 1888:128, pi. 6, fig. 11.
Eocene, Australia. [Triionoharpa]
nodulifera {Cancellaria) G. B. Sowerby I, 1825:Appen-
dix p. 15. Recent, Japan.
nodulifera {Cancellaria) Beyrich, 1856:569, pi. 27, figs.
3-4, 4a. Miocene, Austria. (= C. rothi Semper,
1861)
noduliformis {Cancellaria) 'Sowerby' - Otuka,
1937:1004. Error for C. nodulifera Sowerby.
nodulosa {Cancellaria) Lamarck, 1822b: 113. Recent,
western Africa. (= C. piscatoria (Gmelin, 1791))
nodulosissima {Cancellaria) Sacco, 1894:30, pi. 2, fig.
34; as var. of C. barjonae Pereira da Costa; see
Appendix, Note 2. Miocene, Italy.
nodus {Oamaruia) Finlay, 1930a:78. Miocene, New
Zealand.
notabilis {Cancellaria) Eichwald, 1851:103, pi. 8, figs.
14a-c. Tertiaiy, U.S.S.R.
nucleosa {Anapepta) Marwick, 1931:122, pi. 13, fig.
239. Miocene, New Zealand.
nuda {Trichotropis) Dall, 1927:104. Recent, Florida,
U.S.A. [Iphinopsis]
nympha {Inglisella) Garrard, 1975:40, pi. 4, fig. 14.
Recent, Australia.
nysti {Cancellaria) M. Homes, 1854:305, pi. 34, figs.
la-c. Miocene, Austria.
nysti {Pentadactylus) Briart & Comet, 1870:43, pi. 3,
figs. lOa-c. Paleocene, Europe. [Unitas]
O
obesa {Cancellaria) G. B. Sowerby I, 1832a:52;
1832b:figs. 3-4. Recent, Panamic-Pacific.
obliqua {Cancellaria) 'Lamarck' - Crosse, 1861:233.
Error for C. obliquata Lamarck.
obliquata {Cancellaria) Lamarck, 1822b: 115. Recent,
Indo-Pacific.
obliquata {Cancellaria) 'Kiener' - Sherbom, 1929:4461.
Error for C. obliquata Lamarck.
oblita {Cancellaria) Michelotu, 1861:103. pi. 11, figs.
7-8. Tertiary, Italy.
oblonga (Cancellaria) G. B. Sowerby I, 1825: Appendix
p. 15. Recent, Indo-Pacific.
obnixa (Sydaphera) Iredale, 1936:318, pi. 23, fig. 6.
Recent, Australia.
observatoria (Cancellaria) von Ihering, 1907:214; n.n.
for "C gracilis Ihering" Ortmann, 1902:235, pi.
36, figs. 3a-b; as var. of C. gracilis von Ihering.
Tertiary, Argentina.
obsoleta (Cancellaria) M. Homes, 1856:678, pi. 52,
figs. 3a-b. Miocene, Austria.
obsoleta (Cancellaria) Brugnone, 1880:103, pi. 1, fig.
2; as var. of C. hirta (Br.). Tertiary, Italy.
obtusa (Cancellaria) Deshayes, 1830:187. Recent, Panamic-
Pacific.
obtusa (Cancellaria) Binkhorst, 1861:5, pi. 2, figs. 2a-b.
Cretaceous, Belgium. {- C. cretacea Nyst, 1881)
Page 32
THE NAUTILUS, Supplement 1
occulta (Cancellaria) Beyrich, 1856:576, pi. 28, figs.
7a-b. Oligocene, Germany.
ocoyana (Cancellaria) Addicott, 1970:110, pi. 14, figs.
4-5, 8-9; pi. 16, fig. 20. Miocene, California, U.S.A.
oichingensis (Unitas) Traub, 1984:6; n.n. for U. elon-
gata Traub, 1979, non U. elongata (Cuvillier,
1935). Paleocene, Austria.
okinawana (Merica) Noda, 1980:37, pi. 5, figs. 5a-b;
as subsp. of Merica asprella [sic] Lamarck. Plio-
cene, Okinawa.
okiitanii {Neadmete) Petit. 1974:110, n.n. for "Nead-
inete japonica (Smith)'" Habe, 1961a:73, Appendix
p. 28, pi. 36, fig. 2. Recent, Japan.
oldroydia (Cancellaria) Carson, 1926:51, pi. 1, fig. 5.
Pliocene, California, U.S.A.
oligocancellata (Cancellaria) Sacco, 1894:46, pi. 3, fig.
18; as var. of C. evulsa (Solander); see Appendix,
Note 2. Oligocene, Italy.
oligoevulsa (Cancellaria) Sacco, 1894:45; n.n. for "C.
evulsa Sow." von Koenen, 1889:pl. 10, figs. 1-3;
see Appendix, Note 2. Tertiary, Germany. (= C.
evulsa var. parisiensis Cossmann, 1889)
oligolongum (Cancellaria) Sacco, 1894:4, pi. 1, figs.
la-b; see Appendix, Note 2. Oligocene, Italy.
oligopercostata (Admete) Sacco, 1894:71; n.n. for "C.
subangulosa Wood" Speyer, 1867:pl. 11 [sic; error
for 16], fig. 10. Tertiary, Germany.
olssoni (Admete) Rivera, 1957:186, pi. 4, fig. 27.
Eocene, Peru.
onubensis (Ovilia) Landau, 1984:149, pi. 1, fig. 14; pi.
2, figs. 6-11. Pliocene, Spain.
oppenheimi (Sveltella) Cossmann & Pissarro, 1901:22,
pi. 8, figs. 4-5. Middle Eocene, France.
orbignyana (Cancellaria) Briart & Comet, 1868:21, pi.
2, figs. 9-10. Cretaceous, Belgium.
orbignyi (Cancellaria) 'Blainville,1829' - Sherbom,
1929:4602. Refers to transfer of Buccinum d' or-
bignyi Payraudeau to Cancellaria by Blainville.
[Cantharus]
orcitans (Trigonostoma) 'Martin' - Marks, 1949:458.
Error for Rhomboidestoma oscitans K. Martin.
oregonensis (Cancellaria) Conrad, 1865c:151. Miocene,
Oregon, U.S.A.
oregonensis (Cancellaria) Dall, 1909a:28, pi. 2, fig. 7.
Tertiary, Oregon, U.S.A. (?= C. crawfordiana Dall,
1891)
oregonsis (Cancellaria) "Dair - Dall, 1909a:pl. 2, fig.
7. Plate caption error for C. oregonensis Dall.
ornata (Turbinella) Watelet, 1853:21, pi. 2, fig. 17.
Eocene. France. [?= Cancellaria maglorii Melleville,
1843]
ornata (Cancellaria) Deshayes, 1864:101, pi. 73, figs.
19-20. Lower Eocene, France.
ornata (Cancellaria) Ovechkin, 1954:82, pi. 12, fig. 14;
pi. 13, figs. 4-5; ex Alekseyev MS. Paleogene,
U.S.S.R.
ornata (Admete) Il'ina, 1955:78, pi. 30, fig. 16; ex
Alekseyev MS. Paleogene, U.S.S.R.
ornata (Cancellaria) Alekseyev, 1963:124. pi. 22, figs.
11-12, 15-20. Paleogene. U.S.S.R.
ornatissima (Cancellaria) Boettger, 1906:50; as var. of
C. geslini Basterot. Miocene, Romania.
oscitans (Rhomboidestoma) K. Martin, 1931:12, pi. 2,
figs. 6, 6a. Tertiary, Indonesia.
otagoensis (Zeadmete) Dell, 1956:113, fig. 104. Recent,
New Zealand.
ovalis (Cancellaria) 'Friele' - Paetel, 1888:331. ? Error
for Pleurotoma ovalis Friele.
ovalis (Borsonia) Marshall. 1918:269, pi. 18, figs. 10.
10a. Miocene, New Zealand. [Bonellitia]
ovalis (Zeadmete) Dell, 1956:114, fig. 103. Recent, New
Zealand.
ovata (Admete) E. A. Smith, 1875:426. Recent, ? Japan.
ovata (Cancellaria) G. B. Sowerby I, 1832a:53; 1832b:fig.
2. Recent, Panamic-Pacific.
ovata (Cancellaria) von Koenen, 1889:104, pi. 12, figs.
6a-c, 7a-b. Oligocene, Germany. (= Bonellitia lat-
torfensis Wrigley, 1935)
ovatocrassa (Aphera) Sacco, 1894:67. pi. 3, figs. 79a-b.
Miocene, Italy.
ovatoventricosa (Cancellaria) Sacco, 1894:15; as var.
of C. michelinii Bellardi; ex Grateloup, 1847:pl. 1,
fig. 13; see Appendix, Note 2. Tertiary, France.
ovilia (Ovilia) 'Bast.' - Jousseaume, 1887b:194. Nomen
nudum.
ovoidolaevis (Cancellaria) Sacco, 1894:18, pi. 1. fig.
47; see Appendix, Note 2. Tertiary, Italy.
ovulum (Cancellaria) Geinitz, 1874a:265, pi. 59, figs.
3a-b. Cretaceous, Germany.
ozawai (Cancellaria) Otuka, 1937:1004. Nomen nudum.
ozawai (Cancellaria) 'Otuka MS' - Hatai & Nisiyama,
1940:119, 131. Nomen nudum.
P
pabloensis (Cancellaria) B. L. Clark, 1915:503, pi. 68.
figs. 9, 11. Miocene, California, U.S.A.
pachia (Cancellaria) M. Smith, 1940:45, pi. 2, fig. 2;
as subsp. of C. moorei Guppy. Pleistocene, Florida,
U.S.A.
pacifica (Cancellaria) F. M. Anderson, 1905:199, pi.
15, figs. 43-45. Miocene, Califomia, U.S.A.
pacifica (Cancellaria) Pilsbry & Olsson, 1941:23, pi.
3, fig. 4. Pliocene, Ecuador. (= C. surpacifica
Olsson, 1967)
pahiense (Cymatium) Marshall & Murdoch. 1921:81,
pi. 18, fig. 5. Eocene, New Zealand. [Tatara]
pajana (Cancellaria) Pilsbry & Olsson. 1941:25. pi. 3,
fig. 6. Pliocene, Ecuador.
pakistanicum (Trigonostoma) Fames, 1952:117, pi. 4,
fig. 99. Eocene, Pakistan.
pallida (Cancellaria) E. A. Smith, 1899a:313, text-fig.
Recent, Australia.
palmeri (Cancellaria) Carson, 1926:55, pi. 2, fig. 4.
Pliocene, Califomia, U.S.A.
panamensis (Coluhraria) M. Smith, 1947:55, pi. 2, fig.
6. Recent. Panamic-Pacific. [Tritonoharpa]
panamica (Olssonella) Petit, 1976:35, pi. 2, fig. 1.
Pliocene, Panama.
panamuna (Cancellaria) Garrard, 1975:14, pi. 2, fig.
7. Recent. Australia.
R. E. Petit and M. G. Harasewych, 1990
Page 33
panones (Cancellaria) G. D. Harris, 1895a:65, pi. 6,
fig. 1. Eocene, Texas, U.S.A.
papulosa (Cancellaria) Doderlein, 1863:21. Noincn nu-
dum.
papulosa {Cancellaria) Coppi, 1872:17, pi. 1, figs.
29a-b; ex Doderlein. Tertiary, Italy. (?= C. doder-
leini Mayer, 1868)
papyracea {Cancellaria) Grateloup, 1832:344. Tertiary,
France.
paraguanensis {Cancellaria) H. K. Hodson in Hodson
& Hodson, 1931:44, pi. 24, fig. 10. Miocene,
Venezuela.
paramoorei {Cancellaria) 'Gardner MS' - Mansfield,
1925:31. Nomen nudum.
paramoorei {Cancellaria) Gardner, 1937:372, pi. 45,
figs. 3-4. Miocene, Florida, U.S.A.
parcestriata {Cancellaria) Bronn in Reiss & Bronn,
1862:27, pi. 1, fig. 7; as parcestriata. Tertiary,
Azores.
parilis {Bonellitia) Palmer, 1937:453, pi. 70, figs. 21-25.
Eocene, Louisiana and Texas, U.S.A.
parisiensis {Cancellaria) Cossmann, 1889:228; n.n. for
"C. evulsa (Sol.)" von Koenen, 1889:117, pi. 10,
figs. 1-3; as var. of C. evulsa (Sol.). Tertiary,
Europe, (see C. oligoevulsa Sacco, 1894)
parnensis {Cancellaria) Cossmann, 1896a:42, pi. 3,
figs. 21-22. Middle Eocene, France.
partschi {Cancellaria) M. Homes, 1854:307, pi. 34,
figs. 3a-b. Miocene, Austria.
parva {Cancellaria) I. Lea, 1833:142, pi. 5, fig. 141.
Eocene, southeastern U.S.A.
parva {Cancellaria) Philippi, 1860:187, pi. 7, fig. 18.
Recent, Chile. (= Sveltella philippii Cossmann,
1899) [? Engina]
parva {Cancellaria) Brugnone, 1880:103; as var. of C.
hirta (Br.). Tertiary, Italy.
parva {Inglisella) Laws, 1935:37, pi. 6, fig. 17. Mio-
cene, New Zealand.
parvicarinata {Sveltia) Kautsky, 1925:139, pi. 10, fig.
4; as var. of S. lyrata (Br.). Miocene, Germany.
parvillima {Admete) Sacco, 1894:71; n.n. for "C. pu-
silla Phil." Beyrich, 1857[=1856]:pl. 27, fig. 9; as
var. of Admete minuta (Braun). Tertiary, Germany.
par\'ocrassa {Cancellaria) Sacco, 1894:9; n.n. for "C
ampullacea (Br.)" Almera & Bofill, 1884:pl. C,
figs. 13-14; as var. of C. ampullacea (Br.); see
Appendix, Note 2. Miocene, Spain.
parvotriangula {Cancellaria) Sacco, 1894:6, pi. 1, fig.
6; as var. of C. umbilicaris (Br.); see Appendix,
Note 2. Pliocene, Italy.
parvoturrita {Cancellaria) Sacco, 1894:58, pi. 3, fig.
56; as var. of C taurinia Bellardi; see Appendix,
Note 2. Miocene, Italy.
panula (Cancellaria) Beyrich, 1856:576, pi. 28, figs.
8a-b. Tertiary, Germany.
paschalis {Cancellaria) Thiele, 1925:201, pi. 22, fig.
22. Recent, eastern Africa.
patricia {Cancellaria) Thiele, 1925:200, pi. 22, fig. 20.
Recent, eastern Africa.
patula {Cancellaria) Mayer, 1858:80, pi. 3, fig. 8.
Tertiary, France.
patuxentia {Cancellaria) G. Martin, 1904:167, pi. 43,
figs, lla-b. Miocene, Maryland, U.S.A.
paucicostata {Cancellaria) Sacco, 1894:8, pi. 1, fig. 13;
as var. of C. cassidea (Brocchi); see Appendix,
Note 2. Pliocene, Italy.
paucicostata {Cancellaria) G. B. Sowerby III, 1894:160,
pi. 12, fig. 26. Recent, Persian Gulf.
paucicostata {Merica) Friedberg, 1914:243, pi. 15, fig.
5; as var. of M. fenestrata (Eichwald). Miocene,
Poland.
paucicostata {Sveltia) Peyrot, 1928:218, pi. 14, fig. 7;
as "mut." of S. varicosa (Brocchi). Miocene,
France.
paucicostulata {Brocchinia) Sacco, 1894:69, pi. 3, fig.
86; as var. of B. milraeformis (Brocchi). Pliocene,
Italy.
pauciplicata {Cancelrana) Shuto, 1962:77, pi. 9, figs.
2, 6; pi. 11, figs. 5, 9; text-fig. 14; as subsp. of C.
lischkei (Yokoyama). Tertiary, Japan.
pauciserrata {Cancellaria) Sacco, 1894:44, pi. 3, fig.
7; as var. of C. serrata Bronn; see Appendix, Note
2. Miocene, Italy.
paucivaricatum {Tritonium) Gabb, 1864:95, pi. 28, figs.
209, 209a. Eocene, California, U.S.A. [Admetula]
paytensis {Admete) Olsson, 1930:29, pi. 11, fig. 5.
Eocene, Peru.
paytensis {Plesiotriton) Olsson, 1930:60, pi. 10, figs. 8,
10-11. Eocene, Peru.
pearlensis {Cancellaria) Meyer & Aldrich in Meyer,
1887:7, pi. 1, fig. 4. Eocene, Mississippi, U.S.A.
pellucida {Trigona) Perry, 1811:pl. 51, figs. 1, 2.
Recent, Indo-Pacific. [?= Trigonostoma scalare (Gme-
lin, 1791)]
pemrosei {Cancellaria) 'Harris' - Cossmann, 1899:34.
Error for C. penrosei Harris.
peninsularis {Cancellaria) Olsson, 1942:62, pi. 11, fig.
9. Pliocene, Costa Rica.
penita {Cancellaria) Olsson, 1942:59, pi. 8, figs. 4, 8.
Pliocene, Costa Rica.
penrosei {Cancellaria) G. D. Harris, 1895a:66, pi. 6,
fig. 4. Eocene, Texas, U.S.A.
peracuminata {Cancellaria) Sacco, 1894:29, pi. 2, fig.
26; as var. of C. hirta (Brocchi); see Appendix,
Note 2. Pliocene, Italy.
percostata {Cancellaria) de Gregorio, 1890:48, pi. 3,
figs. 21-22. Eocene, Alabama, U.S.A.
percostatoacuta {Cancellaria) Sacco, 1894:15, pi. 1,
figs. 40a-b; as var. of C. michelinii Bellardi; see
Appendix, Note 2. Miocene, Italy.
percostatula {Cancellaria) Sacco, 1894:14, pi. 1, figs.
35a-b; as var. of C. crassicosta Bellardi; see
Appendix, Note 2. Miocene, Italy.
percrassulata {Cancellaria) Sacco, 1894:15, pi. 1, figs.
39a-b; as var. of C. michelinii Bellardi; see Appen-
dix, Note 2. Miocene, Italy.
perdiciana {Cancellaria) Olsson, 1942:61, pi. 8, fig. 5.
Miocene, Colombia.
Page 34
THE NAUTILUS, Supplement 1
perforata (Neptunea) Gabb, 1864:89, pi. 18, fig. 39.
Cretaceous, California, U.S.A. [Paladmele]
pergradata (Cancellaria) Verco, 1904:142, pi. 26, fig.
19. Recent, Australia.
perhirta (Cancellaria) Sacco, 1894:30; n.n. for "C.
harjanae Pereira da Costa" Almera & Bofill,
1884:pl. B, figs. 4-5; as var. of C. harjonae Pereira
da Costa; see Appendix, Note 2. Tertiary, Spain.
perla (Coliihraria) M. Smith, 1947:55, pi. 2, fig. 2.
Recent. Panamic-Pacific. [Tritonoharpa]
perplexa (Perplicaria) Dall. 1890:90, pi. 3, fig. 1.
Pliocene, Florida, U.S.A.
perproducta {Cancellaria) Sacco, 1894:25, pi. 2, fig.
15; as var. of C. exv,x'stiana Sacco; see Appendix,
Note 2. Miocene, Italy.
perrini (Cancellaria) Carson, 1926:56, pi. 3, fig. 4.
Pliocene, California, U.S.A.
perscalarata (Cancellaria) Sacco, 1894:38, pi. 2, fig.
58; as var. of C. cancellata (L.); see Appendix,
Note 2. Pliocene, Italy.
perscalaris (Cancellaria) Sacco, 1894:64, pi. 3, fig. 71;
as var. of C. trihuliis (Br.); see Appendix, Note 2.
Pliocene, Italy.
perspectiva (Cancellaria) Conrad, 1834:136. Miocene,
Virginia, U.S.A.
persuturata (Cancellaria) Sacco, 1894:56, pi. 3, fig. 44;
as var. of C. varicosa (Br.); see Appendix, Note 2.
Pliocene, Italy.
pertypica (Cancellaria) Sacco, 1894:54, pi. 3, fig. 42;
as var. of C. varicosa (Br.); see Appendix, Note 2.
Pliocene, Italy.
peruana (Aphera) Nelson, 1870:190, pi. 6, fig. 3.
Miocene, Peru.
peruviana (Cancellaria) 'Strong & Hertlein' - Burch,
1945:5. Nomen nudum.
peruviana (Cancellaria) Strong, 1954:17; n.n. for C.
granosa Sowerby, 1832b:fig. 16 (only). Recent,
locality unknown. (?= C. undulata Sowerby, 1849)
peterana (Trigonostoma) Olsson & Harbison, 1953:181,
pi. 28, figs. 6, 6a-b; as subsp. of T. sericea (Dall).
Pliocene, Florida, U.S.A.
petiti (Cancellaria) Olsson, 1967:44; n.n. for C. coss-
manni Olsson, 1922, non Morlet, 1888. Pliocene,
Costa Rica.
petiti (Cancellaria) Ladd, 1982:57, pi. 14, figs. 16-18.
Pliocene, Fiji. (= C. laddi Petit, 1987)
petiti (Africotriton) Beu & Maxwell, 1987:33, pi. 7,
figs, i-m; pi. 8, figs. a-h. Recent, South Africa.
peyreirensis (Sveltia) Peyrot, 1928:235, pi. 13, figs.
9-10. Miocene, France.
pharaonica (Adrnete) Cuvillier, 1933:61, pi. 8, figs. 27,
33. Eocene, Egypt.
pliilippii (Adrnete) von Ihering, 1907:212; n.n. for C.
australis Philippi, 1855, non Sowerby, 1832. Re-
cent, Strait of Magellan.
philippii (Adnwle) Carcelles. 1950:65; n.n. for C. aus-
tralis Philippi, 1855, non Sowerby, 1832. Recent,
Strait of Magellan. (= Adrnete philippii von Ihering,
1907)
philippii (Sveltella) Cossmann, 1899a:30; n.n. for Cancel-
laria parva Philippi, 1860, non Lea, 1833. Recent,
Chile. [? Engina]
pinguis (Cancellaria) Gardner, 1937:370, pi. 44, figs.
12-13. Miocene, Florida, U.S.A.
pinguis (Caveola) Stephenson, 1953:190, pi. 42, figs.
6-7. Cretaceous, Texas, U.S.A. [? Fasciolariidae]
pinolensis (Cancellaria) B. L. Clark, 1918:80. Nomen
nudum.
pirabensis (Cancellaria) Maury, 1925b: 186, pi. 9, fig.
7. Miocene, Brazil.
pisanelloides (Tritonidea) Cossmann, 1923:101, pi. 6,
figs. 45-46. Eocene, France. [?= Turehua]
piscatorium (Buccinum) Gmelin, 1791:3496. Recent,
western Africa. [Solatia]
plagiostoma (Cancellaria) Conrad, 1834:136. Miocene,
Virginia, U.S.A.
planasuturata (Trigonostoma) Richards & Harbison,
1942:217, pi. 21, figs. 4-5. Miocene, New Jersey,
U.S.A.
planispira (Cancellaria) Nyst. 1845:481, pi. 12, figs.
22a-b. Tertiary, Belgium.
planistria (Cancellaria) von Koenen, 1885:11. pi. 1,
figs. 7a-c. Paleocene, Denmark.
planistria (Cancellaria) von Koenen, 1889:110, pi. 8,
figs. 8a-c, 9a-d; as var. of C. quadrata Sowerby.
Oligocene, Europe.
planospira (Cancellaria) 'Nyst' - Dewalque, 1868:419.
Error for C. planispira Nyst.
planospira (Cancellaria) Grant & Gale, 1931:613, pi.
27, fig. 4; as var. of C ohesa Sowerby. Pliocene,
California, U.S.A.
plateaui (Sveltella) Cossmann, 1902:56, pi. 4. fig. 5.
Paleocene, France.
platypleura (Cancellaria) 'Tate' - Tate & Dennant,
1893:221. Nomen nudum.
platypleura (Cancellaria) Tate, 1898:389; unnecessary
n.n. for C. laticoslata Tenison-Woods.
plebeja (Cancellaria) Thiele, 1925:200, pi. 22, fig. 19.
Recent, South Africa.
plicata (Cancellaria) I. Lea, 1833:139, pi. 5, fig. 136.
Eocene, Alabama, U.S.A. (?= C. gemmatum Conrad)
pliocenica (Cancellaria) Sacco, 1894:24; n.n. for "C
spinifera Grateloup" D'Ancona. 1872:92, pi. 11,
fig. 11; see Appendix, Note 2. Pliocene, Italy.
pliocenica (Zeadmete) Finlay, 1930a:77. Pleistocene,
New Zealand.
plioligustica (Cancellaria) Sacco, 1894:63, pi. 3, fig.
68; as var. of C. intermedia Bellardi; see Appendix,
Note 2. Pliocene, Italy.
pliopostica (Cancellaria) Sacco, 1894:19; n.n. for "C.
trochlearis Faujas" Cocconi, 1873:170, pi. 4, figs.
9-11; see Appendix, Note 2. Pliocene, Italy.
plummeri (Cancellaria) Olsson, 1922:85, pi. 6, figs.
2-3. Miocene, Panama.
pluricarinata (Brocchinia) Sacco, 1894:70, pi. 3, fig.
92; as var. of B. parvula (Beyrich). Miocene, Italy.
pluricostata (Trigonostoma) Kautsky, 1925:140, pi. 10,
fig. 6; as var. of T. spinifera (Grateloup). Miocene,
Germany.
R. E. Petit and M. G. Harasewych, 1990
Page 35
pluricosticillata (Cancellaria) Sacco, 1894:37, pi. 2, fig.
55; as var. of C. canccllaia (L.); see Appendix,
Note 2. Pliocene, Italy.
pluricostulata (Cancellaria) Sacco, 1894:36, pi. 2, fig.
52; as var. of C. itniangulata Deshayes; see Appen-
dix. Note 2. Pliocene, Italy.
plurimixta (Cancellaria) de Gregorio, 1882:218. Terti-
ary, Italy.
poecilma (Paladmete) Harbison, 1945:87, pi. 3, figs.
17-18. Cretaceous, Mississippi, U.S.A.
poetzleinsdorfensis (Cancellaria) Sieber, 1936:91, pi.
3, figs. 5a-b; as potileinsdorfensis; as var. of C.
exwestiana Sacco. Miocene, Austria.
polonica (Cancellaria) Pusch, 1837:128, pi. 11, figs.
17a-b; as var. of C. acutangula Faujas de Saint
Fond. Tertiary, Poland.
polygona (Cancellaria) Bellardi, 1840:344. Nomen nu-
dum.
ponderi (Tritonoharpa) Beu & Maxwell, 1987:42, pi.
18, figs, a-j, 1. Recent, Australia.
ponscuspidis (Turehua) Beu & Maxwell, 1987:20, PI.
2, figs, k-1, n-r ; text-fig. 2G. Eocene, New Zealand.
porrecta (Cancellaria) 'Edwards MS' - Newton,
1891:172. Nomen nudum.
porrectum (Coptostoma) Wrigley, 1935:358, pi. 32, fig.
2; as form of Coptostoma quadrata (Sowerby).
Eocene, England.
portoricana (Cancellaria) Maury, 1920:69, pi. 7, fig.
10; as var. of C. laevescens Guppy. Miocene,
Puerto Rico.
postera (Cancellaria) Beyrich, 1856:557, pi. 17, figs.
3a-b, 4a-c, 5a-b; as var. of C. evulsa (Solander).
Oligocene, Europe.
posteror (Cancellaria) "Beyrich' - Kuster-Wendenburg,
1973:126. Error for C. postera Beyrich.
postypica (Cancellaria) Sacco, 1894:58; n.n. for "C.
suessi Hoemes" Hoemes & Auinger, 1890:pl. 25
[sic\ error for pi. 35], fig. 9; see Appendix, Note
2. Miocene, Austria.
posunculensis (Cancellaria) Anderson & Martin, 1914:86,
pi. 8, figs. 7a-c. Miocene, California, U.S.A.
potomacensis (Cancellaria) Clark & Martin, 1901:129,
pi. 21, figs. 3, 3a. Eocene, Maryland, U.S.A.
potzJeinsdorfensis (Cancellaria) Sieber - see poetzleinsdor-
fensis.
pouwi (Trigonostoma) Janssen, 1984:24, pi. 4, figs. 5-9;
pi. 6, fig. 7. Miocene, Netherlands.
praecedens (Cancellaria) Beyrich, 1856:571, pi. 27,
figs. 2a-b; as var. of C. cancellata (L.). Tertiary,
Germany.
praecursoria (Admete) Suter, 1915:3. Nomen nudum,
praeevulsa (Cancellaria) Cossmann, 1898:155; n.n.
for C similis Kaunhowen, 1898, non Sowerby,
1833. Cretaceous, Belgium.
praeevulsa (Bonellitia) Cossmann, 1899a:33: unneces-
sary n.n. for C. similis Kaunhowen, 1898, non
Sowerby, 1833. (= Cancellaria praeevulsa Coss-
mann, 1898)
praeindentata (Cancellaria) Maury, 1925b:188, pi. 9,
fig. 14. Miocene, Brazil.
praetextum (Triton) Bellardi, 1873:227, pi. 14, fig. 13.
Miocene, Italy. [?= Tritonoharpa]
praetiosa (Cancellaria) Oppenheim, 1922:80, pi. 5, figs.
10, lOa-b; ex Rzchak, in litt. Tertiary, Czechoslova-
kia.
priama (Cancellaria) G. D. Harris, 1895a:49, pi. 1, fig.
2. Eocene, Alabama, U.S.A.
prior (Perplicaria) Maury, 1910:17, pi. 4, fig. 8.
Miocene, Florida, U.S.A.
pristina (Mitra) Yokoyama, 1923:8, pi. 1, figs. 8-12.
Pliocene, Japan. [Cancellariidae]
pristina (Paladmete) Stephenson, 1954:39, pi. 8, figs.
24-26. Cretaceous, New Jersey, U.S.A.
producta (Admete) Sars, 1878:217, pi. 13, fig. 2; as
var. of A. viridula (Fabricius). Recent, Arctic Ocean.
producta (Cancellaria) 'Edwards MS' - Jones, 1878:236.
Nomen nudum.
producta (Cancellaria) G. B. Sowerby III, 1903:220,
pi. 4, fig. 5. Recent, South Africa.
producta (Caveola) Stephenson, 1941:364, pi. 70, figs.
11-12. Cretaceous, Texas, U.S.A.
profunda (Babylonella) Tabanelli, 1985:21, figs. 1-5;
as subsp. of B. nassiformis (Seguenza). Pliocene,
Italy.
profundicola (Neadmete) Okutani, 1964:419, pi. 3, fig.
12; pi. 7, fig. 5. Recent, Japan.
profundior (Oamaruia) Cotton & Godfrey, 1932:55; as
subsp. of O. pergradata (Verco). Recent, Australia.
promensis (Merica) Vredenberg, 1921:140, pi. 15, figs.
12a-b. Tertiary, Burma.
propegemmata (Cancellaria) de Gregorio, 1890:46, pi.
3, figs. 14-15. Eocene, Alabama, U.S.A.
propevenusta (Cancellaria) Mansfield, 1929:pl. 16, fig.
2; 1930:47, pi. 17. fig. 2. Miocene/Pliocene, Flor-
ida, U.S.A.
prophylactica (Cancellaria) Icke & Martin, 1907:230,
pi. 14, figs. 11, Ua. Tertiary, Indonesia.
propinqua (Cancellaria) Kaunhowen, 1898:105, pi. 13,
figs. 4-5. Cretaceous, Belgium.
protrigonostoma (Cancellaria) Sacco, 1894:4, pi. 1,
figs. 3a-b; see Appendix, Note 2. Miocene, Italy.
proxima (Cancellaria) Risso, 1826:187, pi. 6, fig. 84.
Tertiary, Europe. Nomen dubium.
prunicola (Cancellaria) G. Martin, 1904:164, pi. 43,
figs. 6a-b. Miocene, Maryland, U.S.A.
pseudangasi (Tritonoharpa) Beu & Maxwell, 1987:44,
pi. 16, figs, j, m-n; pi. 19, figs, a-1; pi. 22, figs. 1,
o. Recent, Indo-Pacific.
pseudaustralis (Cancellaria) 'Tate' - Dennant, 1889:44.
Nomen nudum.
pseudocancellata (Cancellaria) Noetling, 1901:330, pi.
22, figs. lOa-c. Miocene, Burma.
pseudocoronata (Cancellaria) Sacco, 1894:35, pi. 2,
fig. 51; as var. of C. uniangulata Deshayes; see
Appendix, Note 2. Pliocene, Italy.
pseudoevulsa (Cancellaria) d'Orbigny, 1852:11; n.n. for
"C. evulsa Sowerby [sic]" Nyst, 1845:pl. 39, fig.
13; as pseudo-evulsa. Tertiary, Europe.
Page 36
THE NAUTILUS. Supplement 1
pseudonassoides (Cancellaria) Sacco, 1894:38, pi. 2,
fig. 60; as var. of C. cancellata (L.); see Appendix,
Note 2. Pliocene. Italy.
pseudotribulus (Cancellaria) Sacco, 1894:18, pi. 1. fig.
49; as var. of C. dertoscalata Sacco; see Appendix,
Note 2. Miocene, Italy.
pseudotypica (Cancellaria) Sacco, 1894:27, pi. 2, fig.
20; as var. of C. piscatoria (Gmelin); see Appendix,
Note 2. Pliocene, Italy.
pseudovilia (Cancellaria) Sacco, 1894:20, pi. 1, figs.
55a-b, 55bis; as var. of C trochlearis Faujas de
Saint Fond; see Appendix, Note 2. Miocene, Italy.
pseudumbilicare (Trigonostoma) Peyrot, 1928:240, pi.
14, figs. 27-28, 32. Tertiary, Europe.
psinifera (Trigonostoma) 'Grat' - Jousseaume, 1888:23.
Error for T. spinifera (Grateloup).
ptychotropis (Cancellaria) Tate, 1889:156. pi. 9, fig. 5.
Eocene, Australia.
pukeuriensis (Ptychatractus) Suter, 1917:26, pi. 12, fig.
24. Miocene, New Zealand. [Inglisella]
pukeuriensis (Merica) Finlay, 1930a:79, pi. 3, fig. 33.
Miocene, New Zealand.
pulcherrima (Cancellaria) H. C. Lea, 1841:99, pi. 1,
fig. 15. Eocene. Alabama, U.S.A.
pulcherrima (Cancellaria) Almera & Bofill, 1892:12;
Almera, 1894:58. Nomen nudum.
pulcherrima (Cancellaria) Almera & Bofill, 1898:17,
pi. 1, fig. 4. Pliocene, Spain. (= C. bofilli Coss-
mann, 1899)
pulchra (Cancellaria) G. B. Sowerby I, 1832a:50;
I833:fig. 37. Recent, Panamic-Pacific.
pulchra (Cancellaria) 'Briart & Comet' - Dewalque.
1868:388. Nomen nudum.
punctulata (Cancellaria) 'Hinds' - Paetel, 1888:330.
Error for C. funiculara Hinds.
punjabense (Coptostoma) Fames, 1952:118, pi. 5. figs.
106, I07a-b. Eocene, Pakistan.
pupa (Cancellaria) "Edwards MS" - Newton, 1891:172.
Nomen nudum.
pupa (Coptostomella) Finlay & Marwick, 1937:83, pi.
1 1 , fig. 4. Paleocene, New Zealand.
purpuriformis (Cancellaria) Kiener, 1841:37, pi. 7, fig.
4: ex Valenciennes. Recent, Australia.
puruensis (Cancellaria) K. Martin, 1914:127, pi. 2, fig.
56. Tertiary, Indonesia.
puschi (Cancellaria) Semper, 1861:257; n.n. for "C.
citharella (Brong.)" Pusch, 1837:pl. 11, fig. 16.
Tertiary, Poland.
puschi (Cancellaria) Hoernes & Auinger, 1890:276;
n.n. for "C. michelinii Bell." Homes, 1854:326, pi.
35, figs. 14-15. Miocene, Austria.
pusilla (Cancellaria) G. B. Sowerby I, 1832b:fig. 34.
Recent. Indo-Pacific. (?= C. contahulala Sowerby)
pusilla (Fasciolaria) Philippi, 1843:59, pi. 4. fig. 11.
Oligocene, Germany. [Bahylonella]
pusilla (Cancellaria) H. Adams, 1869:274, pi. 19, fig.
12. Recent, Canary Islands. {?= Brocchinia clenchi
Petit, 1986)
pycta (Cancellaria) Olsson, 1964:122, pi. 21, figs. 3,
3a. Miocene, Ecuador.
pygmaea (Cancellaria) C. B. Adams, 1852a:360;
1852b: 136. Recent, Panamic-Pacific.
pygmaea (Paladmete) Sohl, 1964a:273, pi. 45, figs.
36-40; as subsp. of P. gardnerae Wade. Cretaceous,
Mississippi, U.S.A.
pyramidalis (Brocchinia) Sacco, 1894:70, pi. 3, fig. 89;
as var. of B. mitraeformis (Brocchi). Pliocene, Italy.
pyramidata (Cancellaria) Sacco, 1894:25, pi. 2, fig. 14;
as var. of C e.xwestiana Sacco; see Appendix, Note
2. Miocene, Italy.
pyramidatospira (Cancellaria) Sacco, 1894:25, pi. 2,
figs. 18a-b; as var. of C. mutinensis Foresti; see
Appendix, Note 2. Tertiary, Italy.
pyramidalula (Brocchinia) Sacco, 1894:70; n.n. for "C.
mitraeformis (Br.)" Nyst, 1881 [sic; error for
1878]:pl. 38 [sic\ error for pi. 28], fig. 9. Tertiary.
Europe.
pyramidum (Uxia) Cuvillier, 1933:60, pi. 8, figs. 1-4,
26, 32. Eocene, Egypt.
pyrastriformis (Muricites) Schlotheim, 1820:142. Oligo-
cene, Germany. [?= Admetula evulsa (Solander,
1766)]
pyrenaica (Sveltia) Peyrot, 1928:223, pi. 14, figs. 36-37.
Miocene, France.
pyrgota (Cancellaria) Edwards in Lowry. 1866:pl. 3.
Upper Eocene, England.
pyrozonias (Buccinum) Gmelin, 1791:3488. Recent, lo-
cality unknown. [?= Cancellaria cancellata (Linne,
1767)]
pyruliformis (Turbinella) Nyst, 1845:486, pi. 12, figs.
24a-b. Oligocene, Europe. [Turehua]
pyruloides (Egerea) Noszky, 1936:73; as var. of E.
collectiva Gabor. Oligocene, Hungary.
pyrum (Cancellaria) Adams & Reeve, 1950:42, pi. 10,
fig. 16. Recent, Panamic-Pacific.
Q
quadrata (Buccinella) Perry, 1811:pl. 27, fig. 3. Recent,
locality unknown. Nomen inquirendum. [Cancel-
lariidae]
quadrata (Cancellaria) J. Sowerby, 1822:83, pi. 360.
Eocene, England.
quadrata (Cancellaria) Moody, 1916:56. pi. 1. fig. 6.
Pliocene, California, U.S.A. (= C angelana Hanna,
1924)
quadrulata (Cancellaria) Almera & Bofill, 1884:40, pi.
C, figs. 15-16; as var. of C. calcarata (Brocchi).
Tertiary. Spain.
quantula (Cancellaria) Deshayes, 1864:106, pi. 72,
figs. 29-30. Middle Eocene, France.
quasilla (Cancellaria) Petit, 1987:154, fig. 1; n.n. for
C. cretacea E. A. Smith, 1899, non Nyst, 1881.
Recent, India.
quatuorcostata (Admete) Sacco, 1894:73, pi. 3, fig. 98;
as var. of A. nassiformis (Seguenza). Tertiary,
Europe.
quercollis (Volutilithes) G. D. Harris, 1896:85, pi. 8,
fig. 4. Paleocene, Alabama. U.S.A. [Cancellariidae]
R
rakhiense (Coptostoma) Eames, 1952:118, pi. 4, fig.
100. Eocene, Pakistan.
R. E. Petit and M. G. Harasewych, 1990
Page 37
rameum {Coptostoma) Palmer, 1944:323, pi. 25, fig. 3.
Eocene, Texas, U.S.A.
ramonensis {Cancellaria) B. L. Clark, 1918:186, pi. 23,
fig. 7. Oligocene, California, U.S.A.
ranellifonnis (Cancellana) Chenu, 1859:274, fig. 1810.
?Fossil, locality unknown. (?= C. suturalis Sow-
erby, 1822)
rapa (Cancellaria) Nomland, 1917:240, pi. 11, figs. 1,
la. Pliocene, California, U.S.A.
rapella {Cancellaria) C. W. Johnson, 1904:143, text-fig.
Pliocene, North Carolina. U.S.A.
rara {Cancellaria) Aoki, 1954:38, pi. 2, fig. 17. Mio-
cene, Japan.
raricosta {Cancellaria) Sacco, 1894:31, pi. 2, fig. 39;
as var. of C. doderleini Mayer; see Appendix, Note
2. Miocene, Italy.
raulini {Cancellaria) Mayer, 1858:81, pi. 3, fig. 7.
Tertiary, France.
raulini {Cancellaria) 'Grateloup MS' - Peyrot, 1928:220.
Nomen nudum.
ravni {Narona) Gilbert, 1960b:73; n.n. for Cancellaria
angulifera von Koenen, 1885, non Deshayes, 1864.
Paleocene, Denmark.
recessa {Microsveltia) Iredale, 1925:265, pi. 43, fig. 16.
Recent, Australia.
reedii {Admeie) Bell, 1870a:213. Tertiary, England.
reevana {Cancellaria) "Crosse' - Grant & Gale, 1931:613.
Error for C. reeveana Crosse.
reeveana {Cancellaria) Crosse, 1861:237; unnecessary
n.n. for C. elegans Sowerby, 1822.
regina {Admete) Dall, 1911:20. Recent, Bering Sea.
regularia {Admetula) Dockery in MacNeil & Dockery,
1984:164, pi. 58, fig. 16. Oligocene, Mississippi,
U.S.A.
renovata (Sydaphera) Iredale, 1929a:341, pi. 38, fig. 3.
Recent, Australia. (?= Cancellaria undulata Sow-
erby, 1849)
reticulata {Valuta) Linne, 1767:1190. Recent, Carib-
bean. [Cancellaria]
reticulata {Cancellaria) Binkhorst, 1861:66, pi. Va2,
figs. 8a-b. Cretaceous, Belgium. (= C. binckhorsti
Nyst, 1881)
reticulata {Cancellaria) 'Edwards MS' - Newton,
1891:172. Nomen nudum.
reticulatoides {Cancellaria) G. Martin, 1904:164, pi.
43, fig. 7. Miocene, Maryland, U.S.A.
revolutum {Cymatium) Finlay, 1924b:456, pi. 51, figs.
2a-b. Eocene, New Zealand. [Tatara]
rewaensis {Hindsia) Ladd, 1982:47, pi. 11, figs. 10-11.
Pliocene, Fiji. [?= Neadmete nausorensis Ladd,
1982]
reyesi {Cancellaria) Covacevich & Frassinetti, 1986:50,
pi. 2, figs. 6a-c, 7a-c. 8a-c, text-fig. 11. Miocene,
Chile.
rhabdota {Cancellaria) Bayan, 1873:108; n.n. for C.
canaliculata Deshayes, 1864, non Homes, 1854.
Middle Eocene, France.
rhombea {Cancellaria) von Koenen, 1889:114, pi. 8,
figs, la-d, 2a-d. Oligocene, Germany.
rhomboides {Admete) Meek, 1873:501. Cretaceous, Utah,
U.S.A.
rhyssa {Admete) Dall, 1919:306. Recent, western Mex-
ico.
richardpetiti {Cancellaria) Petuch, 1987:17, pi. 2, figs.
5-6. Recent, Florida, U.S.A.
ricinus {Cancellaria) Pecchioli, 1864:505, pi. 5, figs.
6-7. Pliocene, Italy.
rigida {Cancellaria) G. B. Sowerby I, 1832a:53; 1833:fig.
41. Recent, Panamic-Pacific. (?= C. goniostoma
Sowerby, 1832)
ringens {Cancellaria) Sandberger, 1859:pl. 15, figs. 8,
8a-c; 1862:256. Tertiary, Germany.
ringiculaefonnis {Cancellaria) 'Almera & Bofill' -
Almera, 1894:110. ? Error for Marginella ringicu-
laeformis Almera & Bofill.
ripleyana {Trigonostoma) Sohl, 1964a:268, pi. 44, figs.
25, 27. Cretaceous, Mississippi, U.S.A. [not a
cancellariid]
rissoiaeformis {Brocchinia) Cossmann, 1899a:20, 193,
pi. 2, fig. 15. Pliocene, France.
rivalis {Bonellitia) Wrigley, 1935:369, pi. 33, fig. 20;
pi. 35, fig. 48. Middle Eocene, England.
robustum {Tritonium) Geinitz, 1874a:264, pi. 59, figs.
14a-c. Cretaceous, Germany. [Cancellariidae]
rosea {Cancellaria) 'Beck' - G. B. Sowerby II,
1849b:453. Nomen nudum.
rosewateri {Cancellaria) Petit, 1983:250, figs. lA-D.
Recent, Gulf of Mexico.
rothi {Cancellaria) Semper, 1861:255; n.n. for C.
nodulifera Beyrich, 1856, non Sowerby. 1825.
Miocene, Austria.
rotunda {Cancellaria) Dall, 1892:224; as var. of C.
conradiana Dall. Pliocene, Florida, U.S.A.
rotunda {Cancellaria) Anderson & Martin, 1914:87, pi.
8, figs. 4a-b. Miocene, Oregon, U.S.A. (= C.
siletzensis Anderson, 1924)
rotundata {Buccinella) Perry, 1811:pl. 27, fig. 2. Re-
cent, "South Seas". [?= Cancellaria cancellata
(Linne, 1767)]
rotundata {Cancellaria) von Koenen, 1889:103, pi. 12,
figs. 15a-b; as var. of C. subangulosa Wood.
Oligocene, Germany.
rotundata {Merica) Friedberg, 1914:242, pi. 15, fig. 4;
as var. of M. fenestrata (Eichwald). Miocene,
Poland.
rotundulatior {Cancellaria) Sacco, 1894:46, pi. 3, fig.
14; as var. of C. evulsa (Solander); see Appendix,
Note 2. Miocene, Italy.
rotundulina {Cancellaria) Sacco, 1894:50, pi. 3, fig.
29; as var. of C. dertocontorta Sacco; see Appen-
dix, Note 2. Miocene, Italy.
rougeyroni {Cancellaria) Souverbie in Souverbie &
Montrouzier, 1870:427, pi. 14, fig. 1. Recent, New
Caledonia. (?= C. contahulata Sowerby, 1832)
rowelli {Cancellaria) Dall in Guppy & Dall, 1896:307,
pi. 29, fig. 1. Miocene, Dominican Republic.
roydonensis {Cancellaria) 'Edwards MS' - Newton,
1891:172. Nomen nudum.
Page 38
THE NAUTILUS, Supplement 1
rudis (Cancellaria) 'Edwards MS' - Newton, 1891:172.
Nomen nudum.
rudis (Cancellaria) Whitfield, 1892:214, pi. 33, figs.
1-2. Eocene, New Jersey, U.S.A.
rudolphi {Cancellaria) Covacevich & Frassinetti, 1986:51,
pi. 1, figs. 6a-c, text-fig. 8. Miocene, Chile.
ruellensis (Bonellitia) Wrigley, 1935:367; as var. of B.
evulsa (Sol.); ex de Boury MS. Upper Eocene,
France.
rugosa {Cancellaria) Lamarck, 1822b: 115. Recent, Car-
ibbean.
rugosa {Cancellaria) von Koenen, 1889:120, pi. 10,
figs. 8a-c: pi. 11, figs. 7a-b, 8a-c, lOa-d, lla-c.
Oligocene, Germany.
rugosior {Cancellaria) Sacco, 1894:58; n.n. for "C
excellens Beyrich" von Koenen, 1889:pl. 11, fig.
4; as var. of C. excellens Beyrich; see Appendix,
Note 2. Oligocene, Germany.
runchaena {Cancellaria) Gardner, 1937:375, pi. 45,
figs. 8-9. Miocene, Florida, U.S.A.
S
sabinetownensis {Trigonostoma) Le Blanc, 1942:142,
pi. 18, figs. 1-2. Eocene, Texas, U.S.A.
saccoi {Cancellaria) Hoemes & Auinger, 1890:274; n.n.
for "C. bellardi Michelotti" Homes, 1854:314. pi.
34, figs. 17-18. Miocene, Austria.
sacellum {Trigonostoma) Petit, 1976:42, pi. 2, fig. 4.
Pliocene, Panama.
sadko {Admete) Gorbunov, 1946:310, pi. 1, figs. 5a-b.
Recent, North Atlantic.
sagamiensis {Neadmete) Kuroda & Habe, 1971:313
(Japanese), 204 (English), pi. 109, fig. 24. Recent,
Japan.
salbriacensis {Sveltia) Peyrot, 1928:220, pi. 13, figs.
25-26. Miocene, France.
salomacensis {Cancellaria) Peyrot, 1928:206, pi. 12,
figs. 4-5, 8; as var. of C. barjonae Pereira da Costa.
Miocene, France.
salomacensis {Merica) Peyrot, 1928:210, pi. 13, figs.
13-14. Miocene, France.
sanctaemariae {Cancellaria) Carson, 1926:57, pi. 3, fig.
5; as sanctae-mariae. Pliocene, California, U.S.A.
sandbergeri {Cancellaria) Toumouer, 1879:470, pi. 10,
figs. 2a-b. Oligocene, France.
sandbergeri {Rissoa) Muller, 1851:77, pi. 6, fig. 22.
Cretaceous, Germany. [?= Voluta nitidula Muller; ?
CancellariidaeJ
sanjoseensis {Cancellaria) 'Anderson & Martin' - Loel
& Corey, 1932:172. Emendation of, or error for, C.
sanjosei Anderson & Martin.
sanjosei {Cancellaria) Anderson & Martin, 1914:87, pi.
6, figs. 2a-b. Miocene, California, U.S.A.
santa (Gerdiella) Olsson & Bayer, 1972:877, figs. 4-6.
Recent, Caribbean.
santiagensis {Cancellaria) Marks, 1949:462, pi. 78, fig.
6. Miocene, Ecuador.
sathra {Cancellaria) Woodring, 1973:481; n.n. for C.
lipara Woodring, 1970, non 1951; as subsp. of C.
epistomifera Guppy. Pliocene, Panama.
scaberrima {Bonellitia) Chavan. 1947:140. pi. 2, figs.
14-15, text-fig. 1. Tertiary, Jordan.
scabra {Cancellaria) Deshayes, 1830:190. Tertiary,
Europe.
scabriculus {Murex) Linne, 1758:751. [= Voluta cancel-
lata Linne, 1767, q.v.]
scabriuscula {Cancellaria) Moroni, 1958:77, pi. 1, tigs.
2, 2a-b; as subsp. of C. doderleini Moroni. Mio-
cene. Italy.
scabriusculus (Murex) 'Linne" - Linne, 1767:1191.
Error for M. scabriculus L.
scabroides (Cancellaria) Sacco, 1894:10, pi. 1, fig. 18;
as var. of C. ampullacea (Br.); see Appendix, Note
2. Pliocene, Italy.
scala (Murex) Gmelin, 1791:3551. Recent, western
Africa. [Scalptia]
scalaratula (Cancellaria) Sacco, 1894:33, pi. 2, fig. 43;
as var. of C. calcarata (Br.); see Appendix, Note
2. Pliocene. Italy.
scalare (Buccinum) Gmelin, 1791:3495. Recent, Indo-
Pacific. [Trigonostoma]
scalariformis (Cancellaria) Lamarck, 1822b: 11 3. Re-
cent, Indo-Pacific.
scalariformis (Mitra) Borson, 1825:306, pi. 19. fig. 27.
Miocene, Italy. [?= Cancellaria acutangula Faujas
de Saint Fond, 1817]
scalarina (Cancellaria) Lamarck. 1822b: 113. Recent,
Indo-Pacific.
scalarina (Cancellaria) G. B. Sowerby II. 1849b:452,
pi. 96, figs. 87-88. Recent, Indo-Pacific. (?= C.
thomasiana Crosse, 1861)
scalarina (Cancellaria) Conrad. 1863:567. Nomen nu-
dum.
scalarina (Cancellaria) Conrad. 1866:68, pi. 4, fig. 17.
Miocene, Maryland, U.S.A. (?= C. lunata Conrad,
1830)
scalaris (Cancellaria) Michelotti, 1838:396. Nomen nu-
dum.
scalaris (Cancellaria) Bellardi, 1840:344. Nomen nu-
dum.
scalaroides (Cancellaria) Wood, 1857:316, pi. 31, fig.
9. Tertiar)', England.
scalata (Cancellaria) G. B. Sowerby I, 1832b:fig. 27.
Recent, Mauritius.
scalatella (Cancellaria) Guppy, 1873:78, pi. 2, fig. 4.
Pliocene. Jamaica.
scheibei (Cancellaria) F. M. Anderson, 1929:115. pi.
10, figs. 1-4. Miocene, Colombia.
schroeckingeri (Cancellaria) Hoernes & Auinger,
1890:275, pi. 33, figs. 12a-b. Miocene, Austria.
schucherti (Cancellaria) Olsson, 1932:162, pi. 17, figs.
3-4. Miocene, Peru.
schwartzi (Neadmete) Mount, 1970:3, fig. 1. Pliocene,
California, U.S.A.
schythei (Cancellaria) Philippi. 1855:208; 1856a: 164.
Recent, Strait of Magellan.
scobina (Cancellaria) Hedley & Petterd, 1906:222, pi.
38, fig. 12. Recent, Australia.
scopalveus (Aphera) Finlay, 1926a:246, pi. 56, figs.
14-15. Miocene, New Zealand.
R. E. Petit and M. G. Harasewych, 1990
Page 39
scrobiciilata (Cancellaria) M. Homes, 1854:318, pi. 35,
figs. la-b. Miocene, Austria.
sculptura {Cancellaria) I. Lea, 1833:140, pi. 5, fig.
137. Eocene, Alabama, U.S.A. (?= C. alveata
Conrad, 1833)
sculpturata {Cancellaria) "Lea' - Conrad, 1865a:31.
Error for C. sculptura Lea.
secutoriim {Coptostoma) Palmer, 1947:413, pi. 63, figs.
23-25; as var. of C. ulmulum (Harris). Eocene,
Louisiana, U.S.A.
seftoni {Admeie) Berr>', 1956:155, figs. 2, 9. Recent,
California, U.S.A.
selectiim {Trigonostoma) Palmer, 1947:411. pi. 63, figs.
20-22. Eocene. Louisiana, U.S.A.
seniiacostata {Cancellaria) Sacco, 1894:46, pi. 3, fig.
15; as var. of C. evulsa (Solander); see Appendix,
Note 2. Miocene, Italy.
semiclathrata {Cancellaria) Morlet, 1885a:50, pi. 3,
figs. 4, 4a. Middle Eocene, France.
semicostata {Cancellaria) Tate, 1889:157, pi. 10, fig.
3. Miocene. Australia.
semicostata {Bonellitia) 'Sacco' - Harmer, 1918:pl. 40,
fig. 14; et al. Error for B. sertiiacostata (Sacco).
semidisjuncta {Cancellaria) G. B. Sowerby 11, 1849a: 137;
1849b:458, pi. 95, figs. 62-63. Recent, South
Africa.
semilunaris (Murex) Gmelin, 1791:3549. Recent, Sene-
gal. [?= C. piscatoria (Gmelin, 1791)]
semipelliicida {Cancellaria) Adams & Reeve, 1850:42,
pi. 10, figs. 3, 3a. Recent, Japan.
semiplicata {Valuta) Nyst, 1845:593, pi. 15, figs. lOa-b.
Oligocene, Europe. [?= Turehua subgranulata
(Schlotheim, 1820)]
semota {Cancellaria) Jung. 1969:541, pi. 58, figs. 4-5.
Miocene, Trinidad.
semperi {Cancellaria) Speyer, 1867:181, pi. 16, figs. 9,
9a-c. Oligocene, Germany.
semperiana {Cancellaria) Crosse, 1863:65, pi. 2, fig.
7. Recent, New Caledonia.
senarium {Trigonostoma) Petit & Hoerle, 1976:44, pi.
2, fig. 5. Pliocene, Rorida, U.S.A.
sendoi {Cancellaria) Hatai, 1941:110, pi. 3, figs. 5-6;
as s-endoi. Miocene, Japan.
separata {Cancellaria) Deshayes, 1864:97, pi. 72, figs.
20-22. Middle Eocene, France.
septemcostata {Cancellaria) Odhner, 1917:55, pi. 2, fig.
57. Recent, Australia. (?= C. bicolor Hinds, 1843)
septemlirata {Cancellaria) Gabb, 1860c:94, pi. 2, fig.
10. Cretaceous, New Jersey, U.S.A. [not a cancel-
lariid]
septentrionalis {Inglisella) Finlay, 1930b:240, pi. 43,
fig. 14. Recent, New Zealand.
sericea {Cancellaria) Dall. 1892:224, pi. 14, figs. 7, 7a.
Pliocene, Florida, U.S.A.
serramata {Cancellaria) Olsson, 1967:44; n.n. for C.
serrata Olsson, 1964, nan Bronn, 1831. Neogene,
Ecuador.
serrata {Anapepta) Laws, 1935:39, pi. 6, fig. 21.
Miocene, New Zealand.
serrata {Cancellaria) Bronn, 1831:44. Tertiary, Italy.
serrata {Cancellaria) Reeve, 1856:pl. 14, fig. 63. Re-
cent, Indo-Pacific. (= C. crossei Semper, 1861)
serrata {Cancellaria) Olsson, 1964:122, pi. 28, fig. 8.
Neogene, Ecuador. (= C serramata Olsson, 1967)
sheppardi {Admete) Bell, 1919:57, fig. 1; as subsp. of
A. viridula (Fabricius). Tertiary, England.
shilohensis {Cancellaria) Pilsbry & Harbison, 1933:108,
pi. 3, fig. 8. Miocene, New Jersey, U.S.A.
shirleyae {Cancellaria) Olsson, 1967:23, pi. 3, fig. 5.
Pliocene, Florida, U.S.A.
siletzensis {Cancellaria) Anderson in G. D. Hanna,
1924:159; n.n. for C. rotunda Anderson & Martin,
1914, non Dall, 1892. Miocene, Oregon, U.S.A.
silvaerupis {Cancellaria) G. D. Harris, 1897:476, pi.
20, fig. 11; as silvaerupis in text but sylvaerupis,
considered to be correct, on plate caption. Eocene,
Alabama, U.S.A.
silvestris {Cancellaria) Wrigley, 1935:379, pi. 34, fig.
38. Middle Eocene, England.
simiana {Cancellaria) G. D. Hanna, 1924:160; n.n. for
C. crassa Waring, 1917, non Nomland, 1917.
Cretaceous, California, U.S.A.
similaris {Cancellaria) 'Sowerby' - Reeve, 1856:pl. 3,
figs. lOa-b. Error for C. similis Sowerby.
similis {Cancellaria) G. B. Sowerby I, 1833:fig. 38.
Recent, western Africa.
similis {Cancellaria) Aradas, 1846:174, pi. 1, figs.
lla-b. Tertiary, Italy.
similis {Cancellaria) Giebel, 1847:823; 1853:384, pi. 6,
fig. 2. Tertiary, Germany.
similis {Cancellaria) Kaunhowen, 1898:103, pi. 7, fig.
8. Cretaceous, Belgium. (= C. praeevulsa Coss-
mann, 1898)
simplex {Cancellaria) F. M. Anderson, 1905:200, pi.
15, figs. 51-52. Miocene, California, U.S.A.
simplex {Pallidonia) Laseron, 1955:272, figs. 13, 13a.
Recent, Australia.
simplicicostata {Cancellaria) Sacco, 1894:44, pi. 3, fig.
8; as var. of C. serrata Bronn; see Appendix, Note
2. Miocene, Italy.
simplicior {Cancellaria) Sacco, 1894:55, pi. 3, fig. 43;
as var. of C. varicosa (Brocchi); see Appendix,
Note 2. Pliocene, Italy.
simulata {Cancellaria) von Koenen, 1889:107, pi. 17,
figs. 9a-b. Oligocene, Germany.
sinensis {Cancellaria) Reeve, 1856:pl. 8, fig. 35. Re-
cent, Japan.
sinuosa {Cancellaria) Cossmann, 1889:229, pi. 7, fig.
23. Lower Eocene, France.
siouxensis {Cancellaria) Erickson, 1974:217, pi. 20,
figs. 1-5. Cretaceous, North Dakota, U.S.A.
siphonatus {Triton) Reeve, 1844:pl. 18, fig. 81. Recent,
Panamic-Pacific. [Tritonoharpa]
sismondai {Cancellaria) D'Ancona, 1872:102, pi. 13,
figs. 5a-b. Pliocene, Italy.
sismondiana {Cancellaria) 'Grat.' - Peyrot, 1928:225;
ex Grateloup MS. Nomen nudum.
smithfieldensis {Trigonostoma) Oleksyshyn, 1960:101,
figs. 1-2. Miocene, Virginia, U.S.A.
Page 40
THE NAUTILUS. Supplement 1
smithii (Cancellaria) Dall, 1888:70. fig. 292. Recent,
Carolinas to Gulf of Mexico.
smithvillensis {Cancellaria) G. D. Harris, 1895a:65, pi.
6. fig. 2. Eocene, Texas, U.S.A.
smithwickensis (Bonellitia) Hickman, 1980:68, pi. 9,
figs. 2-6. Oligocene, Oregon, U.S.A.
smocki {Cancellaria) Weller, 1907:793, pi. 98, figs.
2-3. Cretaceous, New Jersey, U.S.A.
sobrantensis {Cancellaria) B. L. Clark, 1918:187, pi.
23, fig. 6. Oligocene, California, U.S.A.
solat {Solatia) Jousseaume. 1 887b:222, fig. 3; ex Adan-
son. Recent, western Africa. (?= S. piscaloria
(Gmelin, 1791)).
solida {Cancellaria) G. B. Sowerby I, 1832a:50;
1832b:fig. 6. Recent, Panamic-Pacific.
soriensis {Cancellaria) Eames, 1952:116, pi. 5, figs.
104-105. Eocene, Pakistan.
sotoensis {Cancellaria) Aldrich, 1908:74, pi. 5, fig. 3.
Eocene, Mississippi, U.S.A.
souverbiei {Cancellaria) Crosse, 1868:272, pi. 9, fig. 5.
Recent, ? Indo-Pacific.
sowerbyi {Cancellaria) Bellardi, 1841:9; as sowerby
[sic\\ unnecessary n.n. for C. mitraeformis [sic]
Sowerby, non Brocchi.
sowerbyi {Cancellaria) Crosse, 1861:242; unnecessary
n.n. for C. mitraeformis [sic] Sowerby, non Broc-
chi.
spatiosa {Cancellaria) Nelson, 1870:191. Miocene, Peru.
speciosa {Cancellaria) Deshayes, 1864:100, pi. 73, figs.
1-3. Lower Eocene, France.
speciosa {Caveola) Sohl, 1964a:270, pi. 44, figs. 9-14;
as subsp. of C. acuta (Wade). Cretaceous, Missis-
sippi, U.S.A.
speciosum {Trigonostoma) Vredenburg, 1921:141, pi.
15, figs. 13a-c. Tertiary, Burma.
speciosum {Triton) Bellardi, 1873:226, pi. 14, fig. 12.
Miocene, Italy. [Tritonoharpa]
spectabilis {Cancellaria) Deshayes, 1864:102, pi. 72,
figs. 23-25. Middle Eocene, France.
specularis {Cancellaria) Watson, 1882a:325; 1886:274,
pi. 11, fig. lla-c. Recent, Kerguelen Islands.
spellenbergi {Cancellaria) Riedel, 1932:118, pi. 23,
figs. 13, 13a, 14, 14a, text-fig. 32. Cretaceous,
Cameroon.
spengleriana {Cancellaria) Deshayes, 1830:185. Recent,
Japan.
spengleriana {Cancellaria) 'Kiener' - Sherborn,
1930:6055. Error for C. spengleriana Deshayes.
sphaericula {Cancellaria) Cossmann, 1889:230, pi. 7,
fig. 30. Middle Eocene, France.
sphaeroidea {Admete) Lukovic, 1924:57, pi. 3, figs. 11,
11a. Eocene, Asia.
sphaeroidea {Admete) 'Dukooizen, 1925' - Zoological
Record for 1925. Error for Lukovic, 1924.
sphenoidostoma {Cancellaria) Gardner, 1937:376, pi.
45, figs. 11-12. Miocene, Florida, U.S.A.
spherotopleura {Cancellaria) Gardner, 1937:370, pi. 44,
figs. 14-15. Miocene, Florida, U.S.A.
spinifera {Cancellaria) Grateloup, 1832:342; 1847:pl.
25, fig. 15. Tertiary, France.
spinosa {Admete) Sangiorgi, 1926:104, pi. 7. fig. 22; as
var. of A. nassifurmis (Seguenza). Tertiar%'. Italy.
spinosa {Cancellaria) Grateloup, 1827:21. Tertiary, France.
spinosella {Cancellaria) Sacco, 1894:5; n.n. for "C.
umhilicaris (Br.)" Nyst, 1878:pl. 28, fig. 8; as var.
of C. umhilicaris (Brocchi); see Appendix, Note 2.
Tertiary, Belgium.
spinosissima {Cancellaria) Sacco, 1894:33, pi. 2, fig.
44; as var. of C. calcarata (Brocchi); see Appendix,
Note 2. Pliocene, Italy.
spinosula {Cancellaria) 'Br.' - Borson, 1820:213. Error
for C. spinulosa (Brocchi).
spinulatior {Cancellaria) Sacco, 1894:57, pi. 3, fig. 51;
as var. of C dertovaricosa Sacco; see Appendix,
Note 2. Miocene, Italy.
spinulosa {Voluta) Brocchi, 1814:309, pi. 3, fig. 15.
Pliocene, Italy. [Sveltia]
spirata {Cancellaria) Lamarck, 1822b: 115. Recent, Aus-
tralia.
spirata {Cancellaria) 'Edwards MS' - Newton, 1891:172.
Nornen nudum.
spiratior {Cancellaria) Sacco, 1894:30; n.n. for C.
barjonae Pereira da Costa, 1867:pl. 25, fig. 11
(only); as var. of C. barjonae Pereira da Costa; see
Appendix, Note 2. Miocene, Portugal.
spiratior {Cancellaria) Sacco, 1894:8, pi. 1, fig. 12; as
var. of C. cassidea (Brocchi); see Appendix, Note
2. Pliocene, Italy.
spirifer {Cancellaria) 'von Koenen' - Nyst, 1881:8.
Error for C spinifera Grateloup.
spirulata {Cancellaria) 'Doderiein MS' - Davoli, 1982:36.
Nomen nudum.
springvaleensis {Cancellaria) Mansfield, 1925:31, pi.
2, fig. 12. Miocene, Trinidad.
squarrosa {Cancellaria) Davoli, 1980:228, pi. 1, figs.
la-c, 12a-c; e.x Doderiein MS; as teratological form
of C. serrata Bronn. Tertiary, Italy.
stantoni {Cancellaria) Dickerson, 1913:282, pi. 12, figs.
2a-b. Eocene, California, U.S.A.
staringi {Cancellaria) 'Bosquet' - Staring, 1860:372.
Nomen nudum.
stemerdinki {Babylonella) Janse & Janssen, 1983:134,
pi. 3, figs. 9a-b, text-figs. 6a-b. Miocene, Nether-
lands.
stenomphala {Trigonaphera) Habe, 1961b:432, pi. 24,
fig. 12. Recent, Japan.
stibara {Cancellaria) Gardner, 1937:368, pi. 44. figs.
9-10. Miocene, Florida, U.S.A.
stimpsonii {Cancellaria) Calkins, 1878:250, pi. 8, figs.
4-5. Recent, Florida, U.S.A. (?= Trigonostoma
tenerum (Philippi, 1848))
striatula {Cancellaria) 'Deshayes' - Deshayes, 1835:expl.
to pi. 79; error for C. striatulata Deshayes; cor-
rected on errata page 814.
striatulata {Cancellaria) Deshayes, 1835:503. pi. 79,
figs. 29-30. Middle Eocene, France.
stricta {Admete) Hedley, 1907:295, pi. 54, fig. 10.
Recent, Australia.
R. E. Petit and M. G. Harasewych, 1990
Page 41
strictoturrita {Cancellaria) Sacco, 1894:51, pi. 3, fig.
31; as var. of C. allavillae Libassi; see Appendix,
Note 2. Pliocene, Italy.
strombecki {Cancellaria) Speyer, 1864:267, pi. 40, figs.
2. 2a-b. Oligocene, Germany.
stromboides {Cancellaria) Grateloup, 1832:343. Terti-
ary, France.
strongi {Cancellaria) Shasky, 1961:19, pi. 4, fig. 4.
Recent, western Mexico.
strozzii {Cancellaria) Pecchioli, 1864:503, pi. 5, figs.
3-5. Pliocene, Italy.
stuardoi {Cancellaria) McLean & Andrade, 1982:16,
fig. 53. Recent, Chile.
subaciiminata {Cancellaria) d'Orbigny, 1852:56; n.n.
for C. acuminata Bellardi, 1841, non Sowerby,
1832. Tertiary, Italy.
subalta {Cancellaria) Conrad. 1869:100, pi. 9, fig. 22.
Upper Cretaceous, New Jersey, U.S.A.
subangulata {Cancellaria) 'Scacchi' - Dall, 1889a:131.
? Error for C. coronala Scacchi.
subangiilosa {Cancellaria) Wood, 1842:538. Nomen
nudum..
subangiilosa {Cancellaria) Wood, 1848:66, pi. 7, figs.
20a-bb. Eocene, England.
subangulosa {Cancellaria) 'Weinkauff - Paetel,
1888:331. Error for C. subangulosa Wood.
subanodosa {Brocchinia) Sacco, 1894:69, pi. 3, fig. 83;
as var. of B. mitraeformis (Brocchi). Pliocene, Italy.
subantarctica (Zeadmete) Powell, 1933:236, pi. 20, fig.
7. Recent, Antarctic.
subareticulata {Apbera) Sacco, 1894:67, pi. 3, fig. 78;
as var. of A. bronni (Bellardi). Miocene, Italy.
subaspinosa {Cancellaria) Peyrot, 1928:206, pi. 12, fig.
6; as var. of C. barjonae Pereira da Costa. Miocene,
France.
subasutiirata {Brocchinia) Sacco, 1894:69, pi. 3, fig.
87; as var. of B. mitraeformis (Brocchi). Pliocene,
Italy.
subasuturata {Cancellaria) Sacco, 1894:57, pi. 3, fig.
52; as var. of C dertovaricosa Sacco; see Appen-
dix, Note 2. Miocene, Italy.
subauriculata {Cancellaria) Sacco, 1894:7, pi. 1, fig.
10; as var. of C. taurocrassum Sacco; see Appen-
dix. Note 2. Miocene, Italy.
subbiplicata {Cancellaria) Sacco, 1894:40, pi. 2, fig.
65; as var. of C. dertonensis Bellardi; see Appen-
dix. Note 2. Miocene, Italy.
subcanaliculata {Cancellaria) Grateloup, 1832:343; as
suh-canaliculata; as var. of C. umbilicaris (Broc-
chi). Nomen nudum.
subcanaliculata {Cancellaria) Grateloup, 1847:pl. 25,
fig. 14; as var. of C. umbilicaris (Brocchi). Tertiary,
France.
subcancellata {Cancellaria) d'Orbigny, 1852:54; n.n.
for "C. cancellata (L.)" Grateloup, 1847:pl. 25,
figs. 7, 10. Tertiary, France.
subcarinata {Cancellaria) Bronn, 1831:44. Tertiary,
Italy.
subcylindrica {Cancellaria) von Koenen, 1889:138, pi.
12, figs. 8a-c. Oligocene. Germany.
subevulsa {Cancellaria) d'Orbigny, 1850b:315. Eocene,
Europe.
subevulsopsis {Cancellaria) de Gregorio, 1890:49, pi.
3, fig. 28; as var. of C. tortiplica Conrad. Eocene,
Alabama, U.S.A.
subfusiformis {Admete) Meek, 1873:502. Cretaceous,
Utah, U.S.A.
subgranulatus {Muricites) Schlotheim, 1820:140. Oligo-
cene, Europe. [Turehua]
subhirta {Cancellaria) d'Orbigny, 1852:55; n.n. for "C.
hirta (Br.)" Grateloup, r847:pl. 25, fig. 25. Terti-
ary, France.
sublaevis {Cancellaria) Bellardi, 1840:344. Nomen nu-
dum.
submitroides {Admete) Cossmann & Pissarro, 1905:40,
pi. 16, figs. 17-18. Middle Eocene, France.
suboblitecostata {Cancellaria) Sacco, 1894:30, pi. 2,
fig. 30; as var. of C. hirta (Brocchi); see Appendix,
Note 2. Pliocene, Italy.
subobtusa {Cancellaria) Crosse, 1863:pl. 2, fig. 9; as
var. of C. cumingiana Petit de la Saussaye. Recent,
Panamic-Pacific.
subovula {Uxia) Pchelintsev, 1953:272, pi. 41, figs.
12a-f, text-fig. 44. Cretaceous, U.S.S.R.
subsinensis {Cancellaria) Lobbecke, 1881:12, pi. 2, fig.
3; as var. of C. reeveana Crosse. Recent, Indo-
Pacific.
subspinulosa {Cancellaria) Wood. 1872:49, pi. 6, fig.
10; as var. of C. spinulosa (Brocchi). Tertiary,
England.
subsuturalis {Cancellaria) d'Orbigny. 1852:10; n.n. for
C. suturalis Grateloup, 1832, non Sowerby, 1822.
Tertiary, France.
subtaurinensis {Tritonidea) Vergneau, 1965:79, figs.
7a-b. Oligocene, France. [Loxotaphrus]
subteres {Mataxa) Stephenson, 1941:366, pi. 70, figs.
4-5. Cretaceous, Texas, U.S.A.
subthomasiae {Cancellaria) Dall, 1890:44. pi. 11, fig.
3. Oligocene. Florida. U.S.A.
subtiarophora {Cancellaria) Gardner, 1937:366, pi. 44,
figs. 3-4. Miocene, Florida, U.S.A.
subtilicancellata {Cancellaria) Maury, 1925b: 190, pi.
9, fig. 5. Miocene, Brazil.
subumbilicata {Cancellaria) Grateloup, 1832:340; as
sub-umbilicata; as var. of C. varicosa (Brocchi).
Nomen nudum.
subumbilicata {Cancellaria) Grateloup, 1847:pl. 25, fig.
8; as var. of C. varicosa (Brocchi); see Appendix,
Note 1. Tertiary, France. (= C. subvaricosa d'Or-
bigny, 1852)
suburbana {Sveltella) Wrigley, 1935:363, pi. 32, fig. 9.
Lower Eocene, England.
subvaricosa {Cancellaria) d'Orbigny, 1852:54; n.n. for
"C. varicosa (Br.)" Grateloup. 1847:pl. 25, fig. 8.
Tertiary. France. {- C. subumbilicata Grateloup,
1847, ^.v.)
succineiformis {Cancellaria) Boettger, 1906:51. Mio-
cene, Romania.
suessi {Cancellaria) R. Hoemes, 1875:355. pi. 11, figs.
22a-b, 23a-b. Tertiary, Europe.
Page 42
THE NAUTILUS, Supplement 1
sulcata (Cancellaria) Bellardi, 1840:344. Nomen nu-
dum.
sulcata {Cancellaria) Bellardi, 1841:29, pi. 3. figs. 1-2.
Miocene, Italy.
supercrelacea (Cancellaria) Favre, 1869:91, pi. 10, fig.
17. Cretaceous, Europe.
superstes (Bonellitia) Finlay, 1930b:240, pi. 43, fig. 16.
Recent, New Zealand.
suppar (Cancellaria) Ryckholt, 1862:pl. 36, fig. 25.
Cretaceous, Belgium.
supracosticillata (Cancellaria) Sacco, 1894:70, pi. 3,
fig. 93; as var. of C. parvula Beyrich; see Appen-
dix, Note 2. Miocene, Italy.
supracostulata (Cancellaria) Sacco, 1894:11, pi. 1, fig.
25bis; as var. of C. fenestrata Eichwald; see
Appendix, Note 2. Miocene, Italy.
suprafasciata (Cancellaria) Sacco, 1894:38, pi. 2, fig.
57; as var. of C. cancellata (L.); see Appendix,
Note 2. Pliocene, Italy.
suprafasciolata (Cancellaria) Sacco, 1894:18; as var.
of C. dertoscalata Sacco; see Appendix, Note 2.
Miocene, Italy.
surpacifica (Cancellaria) Olsson, 1967:44; n.n. for C.
pacifica Pilsbry & Olsson, 1941, non Anderson,
1905. Pliocene, Ecuador.
sursalta (Cancellaria) Marks, 1949:461, pi. 78, fig. 4.
Miocene, Ecuador.
suteri (Admete) Marshall & Murdoch, 1920:132, pi. 6,
figs. 5, 5a. Miocene, New Zealand.
sutherlandi (Neadmete) Kanakoff & McLean, 1966:4,
figs. 1-2. Pliocene, California, U.S.A.
suturalis (Cancellaria) G. B. Sowerby I, 1822:fig. 4;
in text as C. suturalis; on plate as C. hiplex.
Eocene, France.
suturalis (Cancellaria) Grateloup, 1832:343. Tertiary,
France. (= C suhsuturalis d'Orbigny, 1852)
sylvaerupis (Cancellaria) G. D. Harris, 1897:476, pi.
20, fig. 11 ; as silvaerupis in text and sylvaerupis on
plate caption; we consider sylvaerupis to be the
intended spelling. Eocene, Alabama, U.S.A.
synchrona (Cancellaria) Ryckholt, 1862:pl. 33, fig. 2.
Cretaceous, Belgium.
tabatai (Cancellaria) Yokoyama, 1926a:265, pi. 32, fig.
12. Pliocene, Japan.
tabulata (Admele) G. B. Sowerby II, 1875:128, pi. 24,
fig. 3. Recent, Arctic Ocean.
tabulata (Admele) 'Friele' - Bell, 1919:58. ? Error for
A. contahulata Friele.
tabulata (Cancellaria) Gardner & Aldrich, 1919:23, pi.
1, fig. 9. Pliocene, Virginia, U.S.A.
taeniata (Cancellaria) G. B. Sowerby II, 1849a:137;
1849b:445, pi. 95, figs. 75-76. Recent, unknown
locality. (?= C. coronata Scacchi, 1835)
taiwanensis (Cancellaria) Nomura, 1935:132, pi. 6,
figs. 61a-b [not 60a-b]. Pliocene, Taiwan.
tampaensis (Trigonostoma) Petit, 1967:218; n.n. for C.
depressa Dall, 1915, non Tuomey & Holmes, 1856.
Oligocene, Florida, U.S.A.
tapeina (Cancellaria) Woodring, 1970:335. pi. 51, figs.
8-9. Pliocene, Panama.
tasmanica (Cancellaria) Tenison-Woods, 1876:150. Re-
cent, Tasmania.
tatei (Aneurystoma) Cossmann, 1899a:24; n.n. for C.
gradata Tate, 1889, non Homes, 1854. Miocene,
Australia.
tauracuta (Cancellaria) Sacco, 1894:12, pi. 1, fig. 26;
as var. of C. fenestrata Eichwald; see Appendix,
Note 2. Miocene, Italy.
taurangulifera (Cancellaria) Sacco, 1894:24, pi. 2, figs.
12a-b; see Appendix, Note 2. Miocene, Italy.
laurangolosa (Cancellaria) Sacco, 1894:pl. 3, fig. 64;
plate caption error for C. taurangulosa Sacco.
taurangulosa (Cancellaria) Sacco, 1894:62, pi. 3, figs.
64, 64bis; as var. of C. lyrata (Brocchi); misspelled
as taurangolosa on plate caption; see Appendix,
Note 2. Miocene, Italy.
taurapertum (Cancellaria) Sacco. 1 894: 1 1 ; error for C.
taurocrassum, corrected in errata on p. 73.
taurelegens (Cancellaria) Sacco, 1894:46, pi. 3, fig. 13;
as var. of C. evulsa (Solander); misspelled as
tauroelegans on plate caption; see Appendix, Note
2. Miocene, Italy.
taurinensis (Pollia) Bellardi, 1873:175, pi. 12, fig. 13.
Miocene, Italy. [Loxotaphrus]
taurinia (Cancellaria) Bellardi, 1840:343, 344.'
taurinia (Cancellaria) Bellardi, 1841:11, pi. 1, fig. 16;
as var. of C. varricosa (Brocchi).' Tertiary, Italy.
taurinia (Cancellaria) Bellardi, 1841:14; as var. of C.
lyrata (Brocchi).' Tertiary, Italy.
taurinia (Cancellaria) Bellardi. 1841:17, pi. 2, figs.
15-16; as var. of C. uniangulata Deshayes.' Terti-
ary, Italy.
taurinia (Cancellaria) Bellardi, 1841:20, pi. 2, figs.
3-4; as var. of C. nodulosa Lamarck.' Tertiary,
Italy.
taurinia (Cancellaria) Bellardi, 1841:24, pi. 3, figs.
15-16; as var. of C. honelli Bellardi.' Tertiary, Italy.
taurinia (Cancellaria) Bellardi, 1841:25, pi. 2, figs.
17-18; as var. of C. evulsa Sow. [sic]} Tertiary,
Italy.
taurinia (Cancellaria) Bellardi, 1841:27, pi. 3, figs.
19-20; as var. of C. cancellata Lam. [sic]} Tertiary,
Italy.
taurinia (Cancellaria) Bellardi, 1841 :pl. 3, figs. 9-10;
as var. of C. contorta Basterot.' Tertiary. Italy.
taurinia (Cancellaria) Bellardi, 1841:35, pi. 4, figs.
13-14; as var. of C. ampullacea (Br.).' Tertiary,
Italy.
Mn 1840 Bellardi used laurinia as a specific name for a Cancellaria
and also as a vandal name for nine olher species. All of these 1840
usages are nomina niida. In 1841 he used laurinia as a varielal name
for nine different species o( Cancellaria. Sacco (1894:58) elevated one
of these, the "var." of C. varicosa (Brocchi), lo species level.
R. E. Petit and M. G. Harasewych, 1990
Page 43
tauroacosticillata {Cancellaria) Sacco, 1894:41, pi. 2,
fig. 69; as var. of C. dertonensis Bellardi; sec
Appendix, Note 2. Miocene, Italy.
tauroaspina (Cancellaria) Sacco, 1894:56, pi. 3, fig.
48; as var. of C. varicosa (Br.); see Appendix, Note
2. Miocene, Italy.
tauroaspinosa {Cancellaria) Sacco, 1894:31, pi. 2, fig.
35; as var. of C. harjonae Pereira da Costa; see
Appendix, Note 2. Miocene, Italy.
tauroaspira (Cancellaria) Sacco, 1894:21, pi. 1, figs.
60a-b; as var. of C. bernardii Mayer; see Appendix,
Note 2. Miocene, Italy.
taurobispinosa (Cancellaria) Sacco, 1894:62, pi. 3, fig.
65; as var. of C. lyrata (Br.); see Appendix, Note
2. Miocene, Italy.
taurobliqiiata (Cancellaria) Sacco, 1894:40, pi. 2, fig.
66; as var. of C. dertonensis Bellardi; see Appen-
dix, Note 2. Miocene, Italy.
taurocaudata (Cancellaria) Sacco, 1894:21, pi. 1, figs.
59a-b; as var. of C bernardii Mayer; see Appendix,
Note 2. Miocene, Italy.
taurocompressa (Cancellaria) Sacco, 1894:41, pi. 2,
fig. 70; as var. of C. dertonensis Bellardi; see
Appendix, Note 2. Miocene, Italy.
tauroconnectens (Cancellaria) Sacco, 1894:34, pi. 2,
fig. 48; as var. of C. calcarata (Br.); see Appendix,
Note 2. Miocene, Italy.
tauroconnectens (Cancellaria) Sacco, 1894:11, pi. 1,
fig. 22; as var. of C. gradata Homes; see Appen-
dix, Note 2. Miocene, Italy.
tauroconvexula (Cancellaria) Sacco, 1894:47, pi. 3,
figs. 22a-b; see Appendix, Note 2. Miocene, Italy.
taurocostatior (Cancellaria) Sacco, 1894:20, pi. 1, fig.
57; as var. of C. obsoleta Homes; see Appendix,
Note 2. Miocene, Italy.
taurocosticillata (Cancellaria) Sacco, 1894:7, pi. 1, fig.
8; as var. of C. scabra Deshayes; see Appendix,
Note 2. Miocene, Italy.
taurocrassum (Cancellaria) Sacco, 1894:7, pi. 1, figs.
9a-b; see Appendix, Note 2. Miocene, Italy.
tauroelegans (Cancellaria) Sacco, 1894:pl. 3, fig. 13;
plate caption error for taiirelegans Sacco.
tauroelongata (Cancellaria) Sacco, 1894:39, pi. 2, fig.
61; as var. of C cancellata (L.); see Appendix.
Note 2. Miocene, Italy.
tauroelongata (Cancellaria) Sacco, 1894:11, pi. 1, fig.
24; as var. of C. fenestrata Eichwald; see Appen-
dix, Note 2. Miocene, Italy.
taurofasciata (Cancellaria) Sacco, 1894:23, pi. 2, fig.
8; as var. of C. geslini Basterot; see Appendix,
Note 2. Miocene, Italy.
taurofaveolata (Cancellaria) Sacco, 1894:18, pi. 1, figs.
51a-c. Probably an error for taurofaveolata, as in
the text reference is made to faveolata [sic] Sow-
erby; see taurofaveolata; see Appendix, Note 2.
Miocene, Italy.
taurofoveolata (Cancellaria) Sacco, 1894:pl.l, figs. 51a-
c; plate caption only; in text and index as tauro-
faveolata, q.v.; see Appendix, Note 2 Miocene,
Italy.
taurolaevigatum (Cancellaria) Sacco, 1894:12, pi. 1,
figs. 28a-c; see Appendix, Note 2. Miocene, Italy.
taurolaevior (Cancellaria) Sacco, 1894:53, pi. 3, fig.
36; as var. of C. callosa Partsch; see Appendix,
Note 2. Miocene, Italy.
taurolatior (Cancellaria) Sacco, 1894:20, pi. 1, figs.
56a-b; as var. of C. obsoleta Homes; see Appendix,
Note 2. Miocene, Italy.
tauroparva (Brocchinia) Sacco, 1894:68, pi. 3, fig. 82;
as var. of B. mitraefarmis (Br.). Miocene, Italy.
tauroparvula (Cancellaria) Sacco, 1894:24, pi. 2, figs.
13a-b; as var. of C westiana Grateloup; see
Appendix, Note 2. Miocene, Italy.
tauroparvula (Cancellaria) Sacco, 1894:10, pi. 1, fig.
19; as var. of C. ampullacea (Br.); see Appendix,
Note 2. Miocene, Italy.
tauropercostata (Cancellaria) Sacco, 1894:10, pi. 1, fig.
20; as var. of C. ampullacea (Br.); see Appendix,
Note 2. Miocene, Italy.
tauropercostata (Cancellaria) Sacco, 1894:52, pi. 3, fig.
33; as var. of C. deshayesiana Desm. [sic]; see
Appendix, Note 2. Miocene, Italy.
taurospinulosa (Cancellaria) Sacco, 1894:65, pi. 3, fig.
75; see Appendix, Note 2. Miocene, Italy.
taurotransiens (Cancellaria) Sacco, 1894:23, pi. 2, fig.
9; as var. of C. geslini Basterot; see Appendix,
Note 2. Miocene, Italy.
taiiroturris (Cancellaria) Sacco, 1894:41, pi. 2, fig. 68;
as var. of C. dertonensis Bellardi; see Appendix,
Note 2. Miocene, Italy.
tauroturrita (Cancellaria) Sacco, 1894:53, pi. 3, fig.
37; as var. of C. callosa Partsch [sic]; see Appen-
dix, Note 2. Miocene, Italy.
tauroturritula (Cancellaria) Sacco, 1894:49, pi. 3, fig.
26; as var. of C. contorta Basterot; see Appendix,
Note 2. Miocene, Italy.
tegalense (Trigonostoma) Oostingh, 1938:108, pi. 6,
figs. 116a-c. Pliocene, Indonesia.
telemba (Cancellaria) Olsson, 1964:121, pi. 21, fig. 4.
Miocene, Ecuador.
tenera (Cancellaria) Philippi. 1848:24. Recent, Carib-
bean.
tenuilineata (Uxia) Wrigley, 1935:375, pi. 34, fig. 30.
Upper Eocene, England.
tenuiplica (Cancellaria) 'Edwards MS' - Newton,
1891:173. Nomen nudum.
tenuiplicata (Sveltia) Wrigley, 1935:360, pi. 32, fig. 5.
Lower Eocene, England.
tenuis (Cancellaria) A. Adams, 1855:123. Recent, "China
Seas". (?= C. scalata Sowerby, 1832)
tenuis (Cancellaria) Reeve, 1856:pl. 16, figs. 75a-b; ex
Adams MS. Recent, ? Indo-Pacific.
tenuispiralis (Turehua) Beu & Maxwell, 1987:22, PI.
1, figs, i, o-p. Oligocene, New Zealand.
tenuistriata (Cancellaria) von Koenen, 1865:471, pi.
15, fig. 1. Oligocene, Germany.
tera (Cancellaria) de Gregorio, 1890:46, pi. 3, figs.
12-13. Eocene, Alabama, U.S.A. (?= C. babylonica
Lea, 1833)
Page 44
THE NAUTILUS, Supplement I
teramachii (Trigonaphera) Habe, 1961a:73, Appendix
p. 27, pi. 36, fig. 4; 1961b:436, pi. 24, fig. 8.
Recent, Japan.
terebrans (Cancellaria) von Koenen. 1889:137, pi. 8,
figs. 4a-c. Oligocene, Geimany.
teres {Zeadmete) Laws, 1940:54, pi. 6, fig. 21. Pleisto-
cene, New Zealand.
terrareginensis {Cancellaria) Etheridge, 1907:327, pi.
60, fig. 1 1 . Lower Cretaceous, Australia.
terryi (Cancellaria) Olsson, 1942:62, pi. 8, fig. I.
Neogene, Panama.
teschi {Cancellaria) Koperberg, 1931:67. Tertiary, Indo-
nesia.
tessella (Trigonosioma) Garrard, 1975:30, pi. 3, fig. 18.
Recent, Australia.
tessellata {Cancellaria) G. B. Sowerby I, 1832a:51;
1832b:figs. 20. 20*. Recent, Panamic-Pacific.
tessellata {Cancellaria) I. Lea, 1833:140, pi. 5, fig.
138. Eocene, southeastern U.S.A. (= C. leai Crosse,
1861; ?= C. alveata Conrad, 1833)
testaventricosa {Cancellaria) Grateloup, 1847:pl. 25,
fig. 16; see Appendix, Note 1. Tertiary, France.
textilis {Cancellaria) Kiener, 1841:10, pi. 7, fig. 1.
Recent, Indo-Pacific.
thiemeana (Cancellaria) Geinitz, 1874b: 175, pi. 31,
figs, lla-b. Cretaceous, Germany.
thisbe (Trigonosioma) Olsson, 1964:126, pi. 22, fig. 6.
Pliocene, Ecuador.
tholoensis (Cancellaria) Ladd, 1934:230, pi. 41, fig. 2.
Miocene, Fiji.
thomasiae (Cancellaria) 'Crosse' - Dall, 1890:44. Error
for C. thomasiana Crosse.
thomasiana (Cancellaria) Crosse, 1861:231; n.n. for
"C. scalarina Lam." Reeve, 1856:pl. 6, fig. 25.
Recent, Indo-Pacific.
thuringiae (Cancellaria) Giebel, 1853:384. Tertiary,
Germany.
thysthlon (Trigonostoma) Petit & Harasewych, 1987:79,
figs. 5, 8-13. Recent, Philippines.
timorensis (Cancellaria) Koperberg, 1931:68; as subsp.
of C. nassoides Schepman. Tertiary, Indonesia.
tjibaliungensis (Cancellaria) K. Martin, 1895:50, pi. 7,
figs. 116a-c. Tertiary, Indonesia.
toroensis (Cancellaria) Olsson, 1922:84, pi. 6, fig. 4.
Miocene, Panama.
torquayensis (Cancellaria) Chapman, 1922:16, pi. 3,
fig. 25. Oligocene, Australia.
torquilla (Cancellaria) Zekeli, 1852:81, pi. 14, fig. 11.
Cretaceous, Europe, [not a cancellariid]
tortipUca (Cancellaria) Conrad, 1865a:32. Nomen nu-
dum.
lortiplica (Cancellaria) Conrad, 1865b: 145. Eocene,
Texas, U.S.A.
tortoniana (Cancellaria) Sacco, 1894:46; n.n. for "C.
hellardii Michelotti" Homes, 1854:pl. 34, fig. 18
only. See C. saccoi Hoemes & Auinger, 1890; see
Appendix, Note 2. Miocene, Austria.
torula (Cancellaria) Weisbord, 1962:398, pi. 40, figs.
9-11. Tertiary, Venezuela.
tosaensis (Fusiaphera) Habe, I961a:72, Appendix p.
28, pi. 35, fig. 21. Recent, Japan.
tosaensis (Trigonaphera) Habe, 1961a:72, Appendix p.
26, pi. 35, fig. 17. Recent, Japan.
tournoueri (Svelria) Peyrot, 1928:224, pi. 13, figs. 11,
12; ex Benoist list. Miocene, France.
trachyostraca (Cancellaria) Brown & Pilsbry, 1911:345,
pi. 23, figs. 1-2; as var. of C. dariena Toula.
Miocene, Panama.
trailii (Cancellaria) 'Hutton' - Garrard, 1975: 44. Error
for C. trailii Hutton.
trailii (Cancellaria) Hutton. 1873:26. Recent, New
Zealand.
tranhirta (Cancellaria) Sacco, 1894:pl. 2, fig. 22; plate
caption error for transhirta Sacco.
transennus (Ficus) Suter, 1917:13, pi. 3, fig. 9. Mio-
cene, New Zealand. [Tatara]
transhirta (Cancellaria) Sacco, 1894:27, pi. 2, fig. 22;
as var. of C. piscatoria (L.) [sic]: see Appendix,
Note 2. Pliocene, Italy.
trapezium (Cancellaria) Borson, 1820:214. Pliocene,
Italy.
traverseensis (Tromina) Clarke, 1961:365, pi. 2, fig. 8.
Recent, South Atlantic. [Iplunopsis]
traversensis (Tromina) 'Clark, 1959' - Bouchet &
Waren, 1985:261. Error for T. traverseensis Clarke,
1961.
trema (Cancellaria) Olsson, 1932:162, pi. 15, figs.
11-12. Miocene, Peru.
triangularis (Cancellaria) Nelson, 1870:191, pi. 6, fig.
10. Miocene, Peru.
tribulis (Cancellaria) Nomland, 1917:238, pi. 12, figs.
1, la; as var. of C. fernandoensis Arnold. Pliocene,
California, U.S.A.
tribulus (Voluta) Brocchi, 1814:310, pi. 3, fig. 14.
Pliocene, Italy. [Sveltia]
tribulus (Cancellaria) 'Nomland' - G. D. Hanna,
1924:160. Error for C. tribulis Nomland.
tributus (Cancellaria) 'Br.' - Cossmann, 1899a: 13.
Error for C. tribulus (Brocchi).
tricarinata (Narona) Traub, 1979:114, pi. 17, figs. 5a-b,
6a-b. Paleocene, Austria.
tricincta (Cancellaria) von Koenen, 1885:10, pi. 1. figs.
5a-b. Paleocene, Denmark.
tricostata (Cancellaria) Sacco, 1894:17, pi. 1, figs.
46a-b; as var. of C. dertoscalata Sacco; see Appen-
dix, Note 2. Miocene, Italy.
trigonostoma (Delphinula) Lamarck, 1822a:231. Recent,
Indo-Pacific. [?= Trigonostoma scalare (Gmelin,
1791)]
trilineata (Cancellaria) Handmann, 1882:263. Miocene,
Austria.
trinodosa (Cancellaria) Chenu, 1859:274, fig. 1812.
?Recent, locality unknown. (?= C. tuberculosa
Sowerby, 1832)
tripUcata (Admefe) Namias, 1898:39. Pliocene, Italy.
triplicatus (Cantharus) Roding, 1798:133. Recent, local-
ity unknown. [- Buccinum pyrozonias Gmelin,
1791; ?= Cancellaria cancellata (Linne, 1767)]
R. E. Petit and M. G. Harasewych, 1990
Page 45
tritonidea (Cancellaria) Gabb, 1866:11, pi. 2, fig. 18.
Pliocene, California, U.S.A.
tritonis {Cancellaria) G. B. Sowerby I, 1832b:fig. 15.
Recent, Indo-Pacific. (?= C. spenglcikina Deshayes,
1830)
triumpha {Trigonostoma) Olsson, 1964:126, pi. 22, figs.
7, 7a. Pliocene, Ecuador.
trochilia (Cancellaria) Olsson, 1964:125, pi. 22, figs.
5. 5a-b. Pliocene, Ecuador.
trochlearis (Cancellaria) Faujas de Saint Fond, 1817:197,
pi. 10, figs. 2a-b. Tertiary, France.
trochlearis (Cancellaria) Lamarck, 1822b: 11 6. Tertiary,
France. (?= C. trochlearis Faujas de Saint Fond,
1817)
truncata (Cancellaria) G. B. Sowerby II, 1849a: 137;
as var. of C. undiilata Sowerby. Recent, ? Philip-
pines.
liiberculata (Cancellaria) 'Sowerby' - G. B. Sowerby
n, 1849b:457; et al. Error for C. tuberculosa G. B.
Sowerby 1.
tuberculata (Fasciolaria) Giebel, 1861:40. Oligocene,
Gemiany. ITiirchua]
tuberculifera (Anapepta) Laws, 1935:38, pi. 6, fig. 19.
Miocene, New Zealand.
tuberculosa (Cancellaria) G. B. Sowerby I, 1832a:51;
1833:fig. 36. Recent, Panamic-Pacific.
tuberculosa (Cancellaria) Sacco, 1894:24; n.n. for "C.
westiana Grat." Almera & Bofill, 1884:pl. B, figs.
1-3; as var. of C e.xwestiana Sacco; see Appendix,
Note 2. Tertiary, Spain.
tumescens (Cancellaria) von Koenen, 1889:123, pi. 10,
figs. 5a-c, 6a-c, 7a-c. Oligocene, Germany.
tumida (Cancellaria) von Koenen, 1889:125, pi. 9, figs.
9a-b. Oligocene, Germany.
tumida (Bonellitia) Hickman, 1980:71, pi. 8, figs.
20-23. Oligocene, Oregon, U.S.A.
tumida (Notlwadmete) Oliver, 1982:16, figs. 1-5. Re-
cent, Antarctic.
turbiformis (Paladmete) Stephenson, 1953:191, pi. 43,
figs. 30-31. Cretaceous, Texas, U.S.A.
turbiniformis (Paladmete) 'Stephenson' - Sohl,
1964a:271. Error for f* . turbiformis Stephenson.
turgidula (Cancellaria) Cocconi, 1873:166, pi. 3, figs.
32-33; as var. of C. boneltii Bellardi. Tertiary, Italy.
turgidula (Cancellaria) 'Guidotti MS' - Cocconi,
1873:168. Nomen nudum.
turneri (Cancellaria) White, 1889:25, pi. 3, figs. 6-7.
Cretaceous, California, U.S.A.
turricula (Cancellaria) Lamarck, 1822b: 116. Tertiary,
Italy. (?= C. Ivrata (Brocchi,1814))
turriculata (Cancellaria) Tate, 1889:156, pi. 10, fig.
14. Eocene, Australia.
turriculata (Hilda) Newton, 1922:29, pi. 4, figs. 24-25.
Eocene, Nigeria. [Varicohilda]
turriculatum (Triton) Deshayes, 1835:608, pi. 80, figs.
7-12. Eocene, France, [non Triton turriculatum
Deshayes, 1833; = Plesiotriton deshayesianus Beu
& Maxwell, 1987]
turns (Cancellaria) 'Edwards MS' - Newton, 1891:173.
Nomen nudum.
turrita (Cancellaria) 'Bell' - Michelotti, 1838:396;
Bellardi, 1840:344. Nomen nudum.
turrita (Cancellaria) Bellardi, 1841:20, pi. 2, figs.
11-12; as var. of C. nodulosa Lamarck. Tertiary,
Italy.
turrita (Cancellaria) G. B. Sowerby II, 1874:721, pi.
59, fig. 12. Recent, unknown locality.
turritella (Cancellaria) Hoeninghaus, 1831:145. Nomen
nudum.
turritior (Cancellaria) Sacco, 1894:9, pi. 1, fig. 17; as
var. of C. ampullacea (Br.); see Appendix, Note 2.
Pliocene, Italy.
turritissima (Cancellaria) Meyer, 1886:73, pi. 1, fig.
15. Eocene, Alabama, U.S.A.
turritovula (Cancellaria) Sacco, 1894:45; n.n. for "C.
evulsa (Sol.)" Speyer, 1867:pl. 11 [sic; error for pi.
16], fig. 1; as var. of C. evulsa (Solander); see
Appendix, Note 2. Tertiary, Germany.
turritula (Cancellaria) Sacco, 1894:32, pi. 2, fig. 40;
as var. of C. doderleini Mayer; see Appendix, Note
2. Miocene, Italy.
turrituloides (Cancellaria) Sacco, 1894:53, pi. 3, fig.
39; as var. of C. deperdita Michelotti; see Appen-
dix, Note 2. Oligocene, Italy.
tyosenensis (Cancellaria) Hatai & Nisiyama, 1940:128,
pi. 5, fig. 3. Miocene, Japan.
typica (Cancellaria) 'Krause' - Paetel, 1888:332. No-
men nudum.
U
uaquala (Cancellaria) Mansfield, 1935:27, pi. 2, figs.
7-8; as subsp. of C. agria Mansfield. Miocene,
Florida, U.S.A.
ulmula (Cancellaria) G. D. Harris, 1895a:66, pi. 6, fig.
6. Eocene, Texas, U.S.A.
umbilicaris (Valuta) Brocchi, 1814:312, pi. 3, figs.
10-11. Pliocene, Italy. [Ventrilia]
umbilicata (Cancellaria) Lesson, 1842:203. Recent,
western Mexico. Nomen dubium.
umbilicata (Admete) Hickman, 1980:77, pi. 9, fig. 13.
Oligocene, Washington, U.S.A.
umbilicatior (Admete) Sacco, 1894:71; n.n. for "C.
subangulosa Wood" von Koenen, 1889:pl. 12, fig.
14; as var. of A. minuta (Braun); see Appendix,
Note 2. Tertiary, Germany.
umbilicina (Cancellaria) Sacco, 1894:20, pi. 1, fig. 58;
as var. of C. doliolaris Basterot; see Appendix,
Note 2. Miocene, Italy.
unalaschkensis (Admete) 'DalF - Kobelt, 1887b: 103.
Error for A. unalashkensis (Dall).
unalashkensis (Cancellaria) Dall, 1873:58, pi. 2, fig.
1. Recent, Aleutian Islands, Alaska, U.S.A.
undata (Admete) Leche, 1878:47; n.n. for "C. viridula
(Fab.)" Middendorff, 1849:pl. 10, figs. 3-4; as var.
of A. viridula (Fabricius). Recent, North Atlantic.
(?= Admete viridula (Fabricius, 1780))
undatocostata (Admete) Verkrtizen, 1875:237; as undato-
costata. Recent, North Atlantic. (?= A. viridula
(Fabricius, 1780))
Page 46
THE NAUTILUS, Supplement 1
undiilata (Cancellaria) G. B. Sowerby 11, 1849a: 136;
1849b:443, pi. 92, fig. 12; pi. 95, fig. 79. Recent,
Tasmania.
uniangulata (Cancellaria) Deshayes, 1830:181. Plio-
cene, Italy.
unicosticillata {Cancellaria) Sacco, 1894:43, pi. 3, fig.
2; as var. of C. bonclli Bellardi: see Appendix,
Note 2. Miocene, Italy.
unidentata {Cancellaria) 'Sowerby' - H. & A. Adams,
1854:276. ? Error for C. indentata Sowerby.
unifasciata {Cancellaria) 'Orbigny' - Dull, I909b:281.
Nomen nudum.
uniplicata {Cancellaria) G. B. Sowerby I, 1832b:fig.
13. Recent, western Mexico. (?= C. mitriformis
Sowerby, 1832)
urceolata {Cancellaria) Hinds, 1843:47; 1844b:41, pi.
12, figs. 7-8. Recent, Panamic-Pacific.
urcianensis {Cancellaria) d'Ancona, 1872:117, pi. 12,
figs. 4a-c. Pliocene, Italy.
urumacoensis {Cancellaria) H. K. Hodson in Hodson
& Hodson, 1931:45, pi. 18, figs. 2-3. Miocene,
Venezuela.
V
valida {Mataxa) Stephenson, 1941:365, pi. 70, figs. 1-3.
Cretaceous, Texas, U.S.A. (?= M. elegans Wade,
1916)
varicifera {Cancellaria) Tenison-Woods, 1879:231, pi.
21, fig. 12. Miocene, Australia.
variciferus {Phos) Tate, 1888:169, pi. 11, fig. 3.
Miocene, Australia. [Loxotaphrus]
varicosa {Cancellaria) 'Brocchi' - error for, or emenda-
tion of. Valuta varricosa Brocchi, 1814. Spelled as
varicosa by almost all authors after Brocchi.
varicosa {Cancellaria) Bellardi, 1840:344; as var. of
"C bonellii Bellardi". Nomen nudum.
varicosa (Merica) Shuto, 1969:172, pi. 11, figs. 4-5,
15-16; as subsp. of M. asperella (Lamarck). Neogene,
Philippines.
varicosissima {Cancellaria) Handmann, 1882:264. Mio-
cene, Austria.
varicosus {Cantharus) Tate, 1888:166, pi. 8, fig. 10.
Eocene, Australia. [Semitriton]
variculosa {Merica) Peyrot, 1928:212, pi. 12, figs.
42-43. Miocene, France.
variecosticillata {Cancellaria) Sacco, 1894:38; as "anom."
of C cancellata pluricosticillata Sacco; see Appen-
dix, Note 2. Tertiary, Italy.
varricosa {Valuta) Brocchi, 1814:311, pi. 3, fig. 8.
Pliocene, Italy. \Sveltia]
vectensis {Uxia) Wrigley, 1935:378, pi. 34, fig. 37; pi.
35, fig. 53; as var. of U. elongata (Nyst). Eocene,
England.
venatrix {Cancellaria) 'Edwards MS' - Newton,
1891:173. Nomen nudum.
venatrix (Bonellitia) Wrigley. 1935:370, pi. 33, fig. 23.
Middle Eocene, England.
veneranda {Admete) Beets, 1946:93, pi. 4, figs. 32-36;
pi. 6, fig. 1. Pliocene-Pleistocene, Netherlands.
venezuelana (Cancellaria) H. K. Hodson in Hodson &
Hodson, 1931:45, pi. 23, figs. 1, 4. Miocene,
Venezuela.
ventricosa (Cancellaria) Hinds. 1843:47; 1844b:41, pi.
12, figs. 11-12. Recent, Panamic-Pacific.
ventricosa (Cancellaria) Grateloup, 1847:2, pi. 1, fig.
16; as var. of C. geslini Basterot. Tertiary, France.
ventricosa (Admete) Friele, 1879:275; as var. of A.
viridula (Fabricius). Recent, North Sea.
ventrilia (Ventrilia) Jousseaume. 1887a: 164, text-fig. 2.
Recent, Caribbean. (?= Trigonostoma tenerum
(Philippi, 1848))
venusta (Cancellaria) Tuomey & Holmes, 1856:144, pi.
28, fig. 18. Pliocene, South Carolina, U.S.A.
vera (Cancellaria) Beyrich, 1856:557; as var. of C.
evulsa (Solander). Tertiary, Germany.
verauxii {Cancellaria) 'Kiener' - H. Adams & A.
Adams, 1854:276. Error for C. verreauxii Kiener.
verbeeki (Cancellaria) K. Martin, 1895:49, pi. 7, figs.
115, 115a. Pliocene, Indonesia.
verrauxii (Cancellaria) 'Kiener' - G. B. Sowerby II,
1849b:450. Error for C. verreaitxii Kiener.
verreauxii (Cancellaria) Kiener, 1841:17, pi. 8, fig. 3.
Recent, Indo-Pacific.
vespertina (Cancellaria) F. M. Anderson, 1905:200. pi.
16, figs. 77-78. Tertiary, California, U.S.A.
vetusta (Cancellaria) Gabb, 1866:12, pi. 2. fig. 19.
Tertiary, California, U.S.A.
vexillata (Tritonoharpa) Ball, 1908:320, pi. 8, fig. 7.
Recent, Panamic-Pacific.
vicarii (Cancellaria) d'Archiac, 1850:291. Nomen nu-
dum, [described as Buccinum vicaryi d'Archiac &
Haime, 1854; not a cancellariid]
vidali (Cancellaria) Philippi. 1887:64. pi. 7, fig. 5.
Miocene, Chile.
vigneauxi (Daguinia) Magne. 1966:127, fig. 1. Mio-
cene, France.
vincenti (Uxia) Gilbert. 1938:120. pi. 4, fig. 10;,
text-fig. 35. Oligocene, Belgium.
vindobonensis (Cancellaria) Sacco, 1894:62; n.n. for
"C. lyrata (Br.)" Homes. 1854:pl. 34, fig. 5; see
Appendix, Note 2. Miocene, Austria.
vinnulum (Trigonostoma) Iredale, 1925:263, pi. 42, fig.
18. Recent, Australia.
viridula (Tritonium) Fabricius, 1780:402. Recent, North
Atlantic. [Admete]
vivus (Plesiotriton) Habe & Okutani, 1981:194, figs.
2-3. Recent, Philippines.
vokesae (Coluhraria) Allen, 1970:71, pi. 2, figs. 10-11.
Eocene, Louisiana, U.S.A. [Plesiotriton]
vokesae (Olssonella) Petit, 1970:84. pi. 1. figs. la-b.
Oligocene, North Carolina, U.S.A..
volutanus (Murex) Risso, 1826:198. Tertiary, Europe.
[?= Cancellaria spinulosa (Brocchi, 1814)]
volutella (Cancellaria) Lamarck, 1803:63. Eocene, France.
volvutella (Cancellaria) 'Lamarck' - Sohl, 1960:128.
Error for C. volutella Lamarck.
vossi (Admetida) Petit, 1976:39, pi. 1, fig. 5. Recent,
Caribbean.
R. E. Petit and M. G. Harasewych, 1990
Page 47
W
waikaiaensis (Trigonostomd) Finlay, 1924b:466, pi. 49,
figs. 4a-b. Oligocene, New Zealand.
waltonensis (Milra) Aldrich, 1910:121. pi. 11, fig. 1.
Miocene, Florida, U.S.A. [Aplwra]
waltoniana {Cancellaria) Gardner, 1937:366, pi. 44,
figs. 5-6. Miocene, Florida, U.S.A.
wannonensis (Cancellaria) Tate, 1889:156, pi. 8, fig.
11. Miocene, Australia.
washingtonensis (Cancellaria) Weaver, 1916:51, pi. 5,
figs. 77-78. Oligocene, Washington, U.S.A.
watanabei (Admete) Shikama, 1962:47, pi. 2, figs.
12a-b. Recent, Japan.
watsoni (Zeadmete) Petit, 1970:86; n.n. for Cancellaria
carinata Watson, 1882, non Briart & Comet, 1877.
Recent, Kerguelen Islands.
wattebledi (Trigonosloma) Peyrot, 1928:255, pi. 14,
figs. 15-17; ex Benoist MS. Neogene, France.
waynensis (Cancellaria) Mansfield, 1940:207, pi. 27,
fig. 19. Oligocene, Mississippi, U.S.A.
weaveri (Cancellaria) Etherington, 1931:108, pi. 14,
figs. 1, 3, 17. Miocene, Washington, U.S.A.
werenfelsi (Cancellaria) Jung, 1965:552, pi. 75, figs.
9-11. Miocene, Venezuela.
westi (Cancellaria) "Bast.' - Mayer, 1872:505. Nomen
nudum.
westiana (Cancellaria) 'Grateloup' - M. Homes,
1854:325; Crosse, 1861:248; et al. Error for, or
emendation of, C. westziana Grateloup.
westralia [Tritonoharpa] Beu & Maxwell, 1987:47, pi.
14, figs, a-g, 1. Recent, Australia.
westralis (Cancellaria) Garrard, 1975:5, pi. 1, figs. 2-3;
as subsp. of C. melanostoma Sowerby. Recent,
Australia.
westziana (Cancellaria) Grateloup, 1847:pl. 1, figs. 18,
21. Tertiary, France.
wigginsi (Cancellaria) Emerson & Hertlein, 1964:362,
figs. 5d-e. Pleistocene, Mexico.
wildi (Cancellaria) Mayer, 1872:505. Nomen nudum.
williamsi (Olssonella) Petit, 1976:36, pi. 2, figs. 2a-b.
Pliocene, Virginia, U.S.A.
wilmeri (Cancellaria) G. B. Sowerby II, 1881:637, pi.
56, fig. 2. Recent, Andaman Islands, India.
withrowi (Trigonaphera) Petit, 1976:39, pi. 2, fig. 3.
Recent, Senegal. (?= Scalptia scala (Gmelin, 1791))
woodii (Admete) Sacco, 1894:72; n.n. for "C costellif-
era Sow." Wood, 1848:pl. 7, fig. 21; as var. of A.
costellifera (Sowerby); see Appendix, Note 2. Terti-
ary, England.
woodringi (Trigonostoma) Jung, 1965:557, pi. 76, figs.
1-2. Miocene, Venezuela.
woodworthi (Admete) Ball, 1905:123. Recent, Califor-
nia, U.S.A.
wouweri (Admete) Harmer, 1918:410, pi. 39, fig. 50.
Pliocene, Netherlands.
wynoocheensis (Cancellaria) 'Weaver' - Etherington,
1931:107. Error for C. wynoochensis Weaver, 1916.
wynoochensis (Cancellaria) Weaver, 1916:50, pi. 4,
figs. 51-54; as wynootchensis in text and wynooch-
ensis on plate explanation; we consider wynoochen-
sis to be the intended spelling. Miocene, Washing-
ton, U.S.A.
wynootchensis (Cancellaria) Weaver - Error for v^ynooch-
ensis, q.v.
X
xavieri (Colubraria) Campbell, 1961:141, pi. 10, figs.
7-8. Recent, Panamic-Pacific. [?= Tritonoharpa vex-
illata Ball, 1908]
xenia (Cancellaria) Olsson, 1964:119, pi. 21, fig. 9.
Miocene, Ecuador.
Y
yokoyamai (Admete) Oyama in Taki & Oyama, 1954:24,
pi. 3, fig. 5; n.n. for "Admete viridula Fabricius"
Yokoyama, 1920:45, pi. 2, fig. 5. Pliocene, Japan.
(= Lora cancellata Otuka, 1937, <7.v.)
yolandia (Cancellaria) Pilsbry & Olsson, 1941:21, pi.
4, fig. 3. Pliocene, Ecuador.
yonabaruensis (Cancellaria) MacNeil, 1960:98, pi. 5,
fig. 4. Miocene, Okinawa.
ytenensis (Cancellaria) 'Edwards MS' - Newton,
1891:173. Nomen nudum.
ytenensis (Bonellitia) Wrigley, 1935:369, pi. 33, fig. 18;
pi. 35, fig. 47. Middle Eocene, England.
Z
zahni (Cancellaria) Bose, 1910:239. pi. 13, fig. 16.
Tertiary, Tehuantepec, Mexico.
zapoteca (Cancellaria) Bose, 1910:240, pi. 13, fig. 17.
Tertiary, Tehuantepec, Mexico.
zetes (Trigonostoma) Kautsky, 1925:141, pi. 10, figs.
10-11. Miocene, Germany.
ziervogeliana (Cancellaria) Lamarck, 1822b: 11 5. [=
Voluta zien'oyelii Gmelin, 1791; Mitridae]
zitteli (Cancellaria) Wanner, 1902:139, pi. 19, figs.
12-13. Cretaceous, northem Africa.
Page 48
THE NAUTILUS, Supplement 1
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APPENDIX
Note 1. Grateloup. In his Atlas of 1847, Grateloup listed
varieties for a number of species of Cancellaria. Most
of the varietal names appear in polynominal form. The
index to the Atlas, valid in its own right, lists varieties
only in binary form. Many of Grateloup's varietal names
were utilized by later authors although the validity of
dating some of them from Grateloup is questionable. The
Atlas, although dated 1840, was not published earlier than
1847. The plates are numbered consecutively within each
of the families represented, with one to five plates for
each family or major group of families, as though each
set of plates was to be accompanied by a monograph.
The plate captions for most plates have a second number
given in parentheses which indicates the position of the
plate in the Atlas. The index uses these numbers. In some
citations, the Cancellaria plate is cited as "Plate 1",
while in other citations it is shown as "Plate 25", the
number given in the index.
Note 2. Sacco. In his monograph of 1894 Sacco utilized five
genera in Cancellariidae and included 14 subgenera
within the genus Cancellaria. In his text, nomina are
treated as if the subgenera are actually genera. As an
example, page 4 of his work is headed "Gen. Cancel-
laria" with a subheading showing Trigonostoma as a
subgenus. The species he includes in the subgenus
Trigonostoma are listed in such a manner that the genus
would appear to be Trigonostoma (e.g., Trigonostoma
oligolongiim instead of Cancellaria (Trigonostoma) oli-
golonga). This same format is utilized for all other
subgenera. Taxa are also listed in this manner in the
index, although the text is clear as to the relative
positions of the genus-group taxa. In this compilation,
new taxa are listed under the genus in which they were
originally proposed. We note that monographers and
cataloguers of other families have treated Sacco's taxa
similarly.
215
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