5.,^. Kf . 5-.. ?• r. .*. t\ It. t »: \ V. r. ^ 5^ >
■ ^ ^ ^ f ^- " ^ Hj *♦ V *( |v *j > i . ^ ^
j^ jj r ^ - - V '. ^ ^
. . >• r ji- V I -.
J5 f r > j^ I >5
M' ♦ w- \^ f,r iv
^vrr.
--.. , *"S.i' ft
V KT
*• ^ V
^J-J'i#- i*' J
K-J^^Kri^
fe;> &:
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.
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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.
LITERATURE CITED
B0ggild, O. B. 1930. The shell structure of the mollusks. Det Kongelige Danske Videnskabernes Selskabs Skrifter, Na- turvidenskabelig og Mathematisk .\fdeling Series #9, 2(2): 231-326, pis. 1-15.
Carcelles, A. 1953. Nuevas especies de gastropodos marines de las republicas oriental del L ruguav \ .Xrgeiitiiia Com- nuinicaciones Zoologicas del Museo de Historia Natural de Montevideo 4(70): 1-16.
Castellanos, Z. J .\ 1970 Reubicacion de algunas especies de V'oiutidae del Mar .Argentlno. Neotropica 16(49): 1-4.
Clench, W. J. and R. D. Turner. 1964. The subfamilies Voluti- nae, Zidoninae, Odontocy mbiolinae and Calliotectinae in the western Atlantic Johnsonia 4(43):129-180.
Dell, R. K. 1978 Additions to the New Zealand Recent mol- luscan fauna with notes on Fachymelon (Palomelon). Rec- ords of the National Museum of New Zealand 1(11):161- 176.
Harasewych, M. G. 1987. Tractolira germonae, a new abyssal Antarctic volutid. The Nautilus 101(1 ):3-8.
Kaiser, P. 1977. Beitragezur Kenntnisder Voluten (Mollusca) in argentinisch-brasilianischen Gewassern (mit der Bei- schreibuiig zweier neuer Arten) Mitteilungen aus dem Hamburgischen Zoologischen Museum und Iiistitut 74:1 1- 26.
Klappenbach, M. .\. 1979. \\Vwii\.\on oV Mar^incUa' corderui Carcelles, 1953 to a new genus in the famiK (Xlontocym- biolinae (Gastropoda). The Nautilus 94(4): 133-135.
Kuroda. T. and T Habe. 1950 \olutidae in Japan. In. Ku- roda, T. (ed). Illustrated catalogue of Japanese shells, 5. Malacological Society of Japan Tokyo, p. 31-38.
Liltved, B. and V. Millard 1986. Volutidae of South Africa. The Strandloper 215:1-4.
Moore, E. J. 1984. Molluscan paleontology and biostratigra- ph\ of the Lower Miocene upper part of the Lincoln Creek Formation in southwestern Washington. Contributions in Science, Natural History Museum of Los .\ngeles County 351:1-42
Novelli. R. and .\. L . G. Novelli 1982 .\lgumas consideravoes sobre a subfamilia Zidoninae e notas sobre a anatomia de Adelomelon hra.siliana (Lamarck, 1811), Mollusca, Gas- tropoda, \'olutidae. Atlantica, Rio Grande 5:23-34.
Okutani, T, M. Tagawa, and H. Horikawa. 1988. Gastropods from Continental Shelf and Slope around Japan Japan Fisheries Resource Association, Tokyo. 203 p
Pilsbry, H. A and A. A Olsson 1954 Systems of the \ olu- tidae. Bulletins of .American Paleontology ■35(152):27I- 306.
Ponder, \V. F. 1970, The morphology of Alcithoc arabica (Mollusca: Volutidae). Malacological Review 3:127-165.
Rios, E. C. 1985. Seashells of Brazil. Fundagao Universidade do Rio Grande, Rio Grande, 328 p.
Shikama, T. 1967. System and evolution of Japanese fulgo- rarid Gastropoda. Science Reports of the Yokohama Na- tional Universit\. Biological and Geological Sciences 13: 23-132.
Sluardo, J. and M. N'illarroel 1974. On some living and lo.ssil volutes referred to Miomelon Dall, 1907 and Proscapliella von Ihering. 1907. The Veliger 17(2):139-155.
Weaver, C. S. and J. E. Dupont. 1970. Living volutes. A 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-
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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
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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
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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
LITERATURE CITED
Cernohorsky, W. O. 1970. Systematics of the families Mitri- dae and \'olutomitridae (Mollusca: Gastropoda). Bulletin of the .\iickiaiid Institute and Mu.seum 8;1-I90.
Cernohorsk). W. O. 1976. The Mitridae of the world Part I. The subfamiK Mitrinae. Indo-Pacifif Mollusca 3(17); 273-528,
Clench, \\. J. and R D Turner. 1964. The subfamilies X'oluti- nae, Zidoninae, Odontoc\ mbiolinae and Calliotectinae in the western .\tlantic. Johnsonia 4(43):129-180.
Conrad, T. A. 1840. New fossil shells from N. Carolina. .Amer- ican Journal of Science 39(2):387-388.
Conrad, T. A. 1842. Observations on the Secondar\ and Ter- tiar\ formations of the southern .Atlantic states; b\ James T. Hodge. With an appendix b\ T .A. Conrad. .American Journarof Science 42(21:344-348.
Conrad, T. .A. 1863. Catalogue of the Miocene shells of the .Atlantic slope Proceedings of the .Acadenn of Natural Sciences of Philadelphia 14(10,T2):559-583.'
Cossmann, .A. E. M. 1899. Essais de paleoconchologie com- paree. Vol. 3. Paris, 201 p.
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 Caloosahatchee River. Part I Pulmonate, Opisthobranchiate and Ortho- dont gastropods. Transactions of the Wagner Free Institute of Science of Philadelphia 3(1 1:1-200.
DuBar, J. R. 1958. Stratigraph) and paleontology of the Late Neogene strata of the Caloosahatchee River area of south- ern Florida. The Florida Geological Survey, Geological Bulletin 40:1-267.
DuBar. J. R. 1962. Neogene biostratigraphs of the Charlotte Harbor area in southwestern Florida. The Florida Geo- logical Survey, Geological Bulletin 43:1-83.
Gardner, J. 1937. The molluscan fauna of the .Alum Bluff Group of Florida Part \'I. Pteropoda, Opisthobranchia and Ctenobranchia (in part). United States Geological Sur- vey Professional Paper 142-F:251-435.
Heilprin, A. 1887. Explorations on the west coast of Florida and in the Okeechobee wilderness. With special reference to the geology and zoology of the Floridian Peninsula. Wagner Free Institute of Science of Philadelphia. Phila- delphia, PA, vi + 134 p.
Houbrick. R. S. 1986. Discover) of a new living Cerithioclava species in the Caribbean (Mollusca: Prosobranchia: Ceri- thiidae). Proceedings of the Biological Society of Wash- ington 99(2):257-260.
\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 Secondary Relict Pocket. Proceedings of the Biological 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 of South-Carolina containing descriptions and figures of the PoK paria, Echinodermata and Mollusca. Russell and Jones, Charleston, SC, xvi + 152 p.
Weaver, C. S. and J. E. duPont. 1970. Living volutes. A monograph of the Recent N'olutidae of the world. Dela- 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
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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- lettes au cours de la croissance. Bulletin de I'lnstitvit Fran- fais d'.'Xfrique Noire 17:1149-1156.
Rancurel, P. 1965. Description de la prodissoconque de Te- redo thunisoni Tr\on et de Bankia anechoensis Roch Cahiers ORSTOM Serie Oceanographie 3:101-105.
Rees, C. B. 1950. The identification and classification of la- 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.
Sigerfoos, C. P. 1908. Natviral histor>, organization, and late development of the Teredinidae, or ship-worms. Bulletin of the Bureau of Fisheries 27:191-232.
Sullisan, C. M. 1948. Bivalve larvae of Malpeque Bay, P.E.I. Bulletin of the Fisheries Research Board of Canada 77:1- 36,
Turner, R. D. 1966. A surve\' and illustrated catalogue of the Teredinidae (.MoUusca: Bivalvia). The Museum of Com- parative Zoology, Harvard University, Cambridge, 265 p.
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- ganisation for Economic Co-operation and Development, Paris, p. 17-64.
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- tringham (eds. ). Marine borers, fungi and fouling organ- isms of wood. Organisation for Economic Co-operation and Development, Paris, p. 259-301.
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
|
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30- |
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<u |
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o |
20- |
|
i- |
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|
|
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15- |
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|
D |
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Z |
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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
A quarterly devoted to malacology.
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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
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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
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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
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mmm
mm
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D. G. Reid, 1989
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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
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teg tag ocg
63 62 61 60
sr oag
ocg
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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
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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).
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D. G. Reid, 1989
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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).
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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
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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).
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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.
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D. G. Reid, 1989
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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).
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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].
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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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Mori, K., S. Nishihama and M Tanaka. 1985. Community structure of a rocky shore in Tsuji-shima island. .Amakusa. 111. The analysis of relationships between distribution of organisms and micro-topographical conditions using small quadrat. Publications of the .Amakusa Marine Biological Laboratory 8:43-63.
Morton, B. and J. Morton. 1983. The sea shore ecology of Hong Kong. Hong Kong University Press, Hong Kong. 350 p.
Mowry, R, W. 1956. .Alcian blue techniques for the histo- chemical study of acidic carboh\drates. Journal of His- tochemistry and C\lochemistr\ 4409-410,
Nevill, G. 1885. Hand list of Mollusca in the Indian Museum, Calcutta, Part 2. Indian Museum, Calcutta, 306 p.
Ohgaki, S. 1981. Spawning activity in Nodilittorina exigua and Peasiella roepstorffiana (Littorinidae, Gastropoda). Publications of the Seto Marine Biological Laboratory 26: 437-446.
Ohgaki, S 1985. Distribution of the famiK Littorinidae (Gas- tropoda) on Hong Kong rock) sliores. In: Morton, B. and D. Dudgeon (eds.). The malacofauna of Hong Kong and southern China II, \ ol. 2:457-464.
Oyama, K. and Y. Takemura. 1961. The molluscan shells, Vol. 5. Resources Exploitation Institute, Tokyo.
Pallary, P. 1926. Explication des planches de J.C. Savigny. Memoires presentes a I'lnstitute d Egypte 11:1-138.
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Preston, H. B. 1908. Descriptions of new species of land, marine and freshwater shells from the .Andaman Islands. Records of the Indian Museum 2:187-210.
Reeve, L. A. 1862. Monograph of the genus Trochus. Con- chologia Iconica, \'ol. 13 Lovell Reeve & Co., London.
Reid, D. G. 1986. The littorinid molluscs of mangrove forests in the Indo-Pacific region: the genus Littoraria. British Museum (Natural Histor) ), London. 228 p.
Reid, D, G, 1988. The genera Bembicium and Risellopsis (Gastropoda: Littorinidae) in .Au.stralia and New Zealand. Records of the .Australian Museum 4091-150.
Reid, D. G. 1989. The comparative morphology, phylogen) and evolution of the gastropod family Littorinidae. Philo-
D. G. Reid, 1989
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sophical Transactions of the Royal Society, London. Series B, 324:1-110.
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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,
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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 |
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THE NAUTILUS, Vol. 103, No. 2
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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.
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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. Kraus, Vienna. [36], 442[17] p., 18 pis.
Buonanni, F. 1681. Ricreatione dell' occhio e della mente neir osservation' delie C>hiocciole, proposta a' curiosi delle opere della natura, &iv. N'arese, Rome, [xiv], 384[1.5]. 109 pis.
Chemnitz, J. H. 1780. Neues systematisches Conchylien Cab- inet. Rafpe, Nijrnberg. volume 4. [.\.\iv], 344 p., pis. 122- 159. [Although the plates were published in 1780, Born had a copy of the manuscript prior to publication]
Uall, W. H. 1885. On Turbinella pyrum. Lamarck, and its dentition. Proceedings of the Lnited States National Mu- seum 8:345-348, pi 19
Dall, \V H. 1915. .\n index to the Museum Boltenianum. Smithsonian Institution Special Publication 2360: 64 p.
Gualtieri, \ 1742 Index Testarum Conchyliorum ciuae ad- servantur in Museo N. Gualtieri . . . et methodice distri- butae exhibenlur tabulis CX. (Introductio ad historiam Testaceorum a ... P. Tournefortio in codice MS. relicta.) Albizzini, Florence, p. xxiii, 110 pis., with descriptive let- terpress, 1 portrait.
Haller, B. 1888. Die morphologic der Prosobrancher, gesam- melt auf einer Erdumsegelung durch die kiinigl. italie- nische Korvette "Vetor Pisaiii ". Morphologisches Jahr- buch. Fine Zeitschrift fur Anatomie und Entwicke- lungsgeschichte 14(1):54-169.
Hanley, S. 1855. Ipsa Linnaei Concli\lia. The Shells of Lin- naeus, determined from his manuscripts and collection. Also, an exact reprint of the Vermes Testacea of the "Sys- tema Naturae and 'Mantissa . Williams and Norgate, Lon- don, 556 p., 5 pis.
HarasewNch. M G. 1987 .\ re\isi()n of the genus Berithovo- lula w ith notes on the evolution of the subfamil) Plycha- tractinae (Prosobranchia: Turbinellidae). The Nautilus 101(4):166-181.
Hollister, S. C. 1958. \ review of the genus Busycon and its allies — Part I. Palaeontograpliica .\mericana 4(28):47-126, pis. 8-18.
International Commission on Zoological Nomenclature. 1985. International Code of Zoological Nomenclature. Third edi- tion, XX, 338 p.
Kesteven, H. L. 1904. The anatomy of Megalatractus. Mem- oirs of the Australian Museum 4:419-449, pis. 39-42.
Linne. C. 1758 Caroli Linnaei ... Systema Naturae per Reg- na tria Naturae, secundum classes, ordines. genera, species, cum characteribus, differentiis, synonymis, locis . . . Editio decima reformata. Impensis Direct. Laurentii Salvii: Hol- miae, Tom 1. .^nimalia. p. [Iv], 823.
l.iiine, C. 1764. Museum S:ae R:ae M:tis Ludovicae Ulricae Reginae Svecorum, Gothorum, Nandalorumciiie ... in quo .\nimalia rariora, exotica, imprimis Insecta it (^onchilia dcicribuntur in determinatur Prodromi instar editum. 6iC. Holmiae. Lileris ii impensis Direct. Laur. Salvii. vi -I- 720
+ [2] P
Linne. C. 1767. Caroli a Linne . . . Systema Naturae per Regna tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis. locis . . . Editio duodecima reformata. Impensis Direct. Lau- rentii Salvii Holmiae, Tom 1. Regnum .\nimale Pars II. Classis \'-VI. Nomina Generica. .Nomina trivialia Papi- liniiuni and Phalaenarum. Synon\ma, Termini .\rtis. .Ap- pendix S\non\ morum, .\ddenda. Errata, p. 5:33-1327. [36].
Roding, P. F. 1798. Museum Boltenianus sive Catalogus ci- meliorum e tribus regnis naturae quae olim collegerat. Pars Secunda continens Conchylia sive Testacea univalvia, bivalvia & multivalvia Trappi. Hamburg, viii + 199 p.
Rumphius, G. E. 1705. D Amboinsche Rariteitkamer, behel- zeiide eene beschrv\inge van allerhande zoo weeke als harde schaalvisschen, le weeten raare Ivrabben. Kreeften, en diergeKke Zeedieren, als mede allerhande Hoorntjes en Schulpen, die men in d .\mboinsclie Zee vindt: daar beneven zommige Mineraalen, Gesteenten, en soorten van Aarde, die in d'.\mboinsche, en zommige omleggende Ei- landen gevonden worden. Verdeelt in drie booken, &c. Fr. Halma, Amsterdam, p. [xx\iii]. 340 [43], 60 pis., 1 portrait.
Schroter, J, S. 1783. Einleitung in die Coiich\lieii-Kentniss nach Linne. J. J. Gebauer. Halle. X'olume 1 xxxii. 860, [2] p.. pis. 1-3,
Thicle, J. 1929 Handbuch der SNSteiualisclieii W'eichtier- kunde. G. Fischer, Jena. l(l):l-376.
Wenz, W. 1943. Gastropoda. In: O. H. Schindewolf (ed.). Handbuch der Paliiozoologie. Band 6. Gebriider Born- traeger, Belin. Teil 6:1201-1506.
W inckwortli, K. 1945. The types of the Boltenian Genera. Proceedings of the Malacological Society of London 26(4 & 5):136-148.
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THE NAUTILUS
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Volume 103, Number 3 December 28, 1989 ISSN 0028-1344 |
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EDITOR-IN-CHIEF Dr. M. G. Harasew ych Division of Mollusks National Museum of Natural History Smithsonian Institution Washington, DC 20560
ASSOCIATE EDITOR Dr. R. Tucker Abbott American Malacologists, Inc. P.O. Box 2255 Melbourne, FL 32902
CONSULTING EDITORS Dr. Riidiger Bieler Department of Malacology 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
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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
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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
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Dr. Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138
Ms. Paula M. Mikkelsen Harbor Branch Oceanographic Institution, Inc. Ft. Pierce, FL 33450
Dr. Donald R. Moore
Division of Marine Geology
and Geophysics
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Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138
Dr. Geerat J. V'ermeij Department of Geology University of California at Davis Davis, CA 95616
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T H Ef7N AUT I L U S
CONTENTS
Marine Biological Laboratory LIBRARY
JAN 8 1990
Woods Hole, Mass.
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
92
Carl M. Way Andrew C. Miller Barrv S. Pavne
The influence of physical factors on the distribution and abundance of freshwater mussels (Bivalvia: Unionidae) in the lower Tennessee River
96
W. B. Saunders Paul N. Bond Lee C. Hastie David Itano
On the distribution of Nautilus pompilius in the Samoas, Fiji and Tonga
99
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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 |
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Bradford H. Burnham Paul E. Fell |
Distribution of Melampus bidentatus (Say and Succinea wilsoni (Lea) within a tidal marsh in eastern Connecticut |
109 |
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Alan R. Kabat |
The "Gray Catalogues" [Mollusca] of the British Museum |
113 |
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INews and Notices |
116 |
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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.
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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
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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 ).
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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,
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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-
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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,
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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
News and Notices
MONOGRAPHS OF MARINE MOLLUSC A CHANGES HANDS
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THE NAUTILUS
Volume 103, Number 4 April 18, 1990 ISSN 0028-1344
A quarterly devoted to malacology.
Marine Biological Labor^'- LIBRARY ^
my 2 mo
EDITOR-IN-CHIEF Dr. M. G. Harasewych Division of Mollusks National Museum of Natural History Smithsonian Institution Washington, DC 20560
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The American Museum of Natural
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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:
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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-
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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.
LITERATURE CITED
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Abbott, R. T. 1974. American seashells, 2nd ed. Van Nostrand Reinhold, New York, vii + 663 p., text figs., 24 pis
Baker, F. C. 1919. Mollusca of the Crocker land expedition to Northwest Greenland and Grinnell Land. Bulletin of the American Museum of Natural History 4:479-517.
Binney, W. G. 1870. Report on the Invertebrata of Massa- chusetts. Second Edition, comprising the Mollusca. Wright & Potter, Boston, v + 525 p., 12 pis.
Broderip, W. G. and G. B. ,Sowerby 1829 Observations on new or interesting Mollusca, contained, for the most part, in the museum of the Zoological Society. Zoological Journal 4:359-366.
Chemnitz, J. H. 1781. NeuessystematischesConchylien-Cab- inet. Vol. 5. Gabriel Nicolaus Raspe, Nurnberg, [20] -I- 34 p., pis. 160-19,3.
Couthouy, J. P. 1838 Descriptions of new species of Mollusca and shells, and remarks on several polypi found in Mas- sachusetts Bay. Boston Journal of Natural History 2:53- 111.
Dillwyn, L. W. 1817. A descriptive catalogue of recent shells. Vol. 2. John & Arthur Arch, London, p. 581-1092.
Fabricius, O. 1780. Fauna Groenlandica. J. G. Rothe, Co- penhagen, xvi -I- 452 p.
Forbes, E. and S. Hanley. 1853. A history of British mollusca and their shells. Vol 2. John Van Vorst, London, viii -t- 557 p., pis. 33-69
Fretter, V. and A Graham. 1977. The prosobranch molluscs of Britain and Denmark, Part 2 — Trochacea The Journal of Molluscan Studies, Sup|)lement 3, p. 39-100
Galkin, la. I. 1955. Trochid gastropod mollusks of the far eastern northwestern seas of the USSR. (Family Trochi- dae). Analytical Fauna U.S.S.R. 57:1-131 [in Russian J
Gmelin, J. F. 1791. Linneaus, Systema Naturae. Edition 13, Vol. 1(6). Leipzig, p. .3021-3910.
Gould, A. A. 1841. Report on the invertebrata of Massachu- setts. Cambridge, xiii + 373 p, 15 pis
Gray, J. E. 1826. On Terelnatula custata dud TurlK/ carneus. Zoological Journal 2:567.
Hagg, R 1905. Mollusca und Brachiopoda gessammelt von der schwedi.schen zoologischen Polarexpedition nach Spits- bergen, dem nordostlichen Gronland und Jan Ma\t'n i j 1900, II. Scapho poda. Gastropoda. Placophora und zwei vorhcr nicht erwahnte Lamellibranchiata .Arkiv for Zoolo- gi 2(13):1-1.36.
Hancock, A. 1846. A list of shells dredged on the west coast of Davis Strait; with notes and descriptions of eight new species. Annals and Magazine of Natural History 18:323- 338.
Jeffreys, J. G. 1865. British conchology. Vol. 3 John \an Vorst, London, 393, p., 9 pis.
Jeffreys, J. G. 1877. New and peculiar Mollusca of the Patelli- dae and other families of Gastropoda procured in the \'al- orous' expedition. Annals and Magazine of Natural His- tory (ser. 3) 19:231-243.
Johnson, C. W. 1915. Fauna of New England, 3. List of the Mollusca Occasional Papers of the Boston Society of Nat- ural History 7:1-231.
Johnson, C^. VV 1934 List of the marine Mollusca of the Atlantic Coast from Labrador to Texas. Proceedings of the Bo.ston Society of Natural History 40(1): 1-203.
Kelly, K. L. and D. B. Judd. 1965. The ISCC-NBS method of designating colors and a dictionary of color names. National Bureau of Standards Circular 553:v -t- 158 p. [Reprint; includes supplement ISCC-NBS color name charts illustrated with centroid colors].
Lamarck, J P B M de. 1804. Suite des Memories sur les fossiles des environs de Paris. .-Xnnales du Museum National d'Histoire Naturelle 446-55, 105-115, 212-222, 289-298, 429-436.
Leach, W. R. 1819. Description des nouvelles especes d ani- maux decouvertes par le vaisseau Isabelle dans un voyage au pole boreale. Journal de physique, de chimie, d'hi.storie naturelle et des arts 88:462-477.
Linne, C 1758. Systema Naturae. Tenth Edition. \'ol 1, Laurentius Salvus, Stockholm, 824 pp.
Lowe, R. T. 1825. Descriptions of some shells, belonging principally to the genus Chiton, observed on the coast of Argyleshire in the summer of 1824. Zoological Journal 2: 93-i08.
Moller, H. P. C. 1842. Index Molluscorum Groenlandiae. Kroyer's Naturhi.storisk Tidsskrif t 4:76-97. (Also published as separate.)
Morch, O. A. L. 18.57. Fortegnelse over Gronlands Bloddyr. In: Rink's Gronland Geographisk og Statistik Beskrivet, Tillaeg No. 4:75-100.
Nordsiek, F". 1968. Die europaischen Meeres-Gehauses- chnecken (Prosobranchia). Gustav Fischer Verlag, Stutt- gart, viii -I- 273 p., 31 pis.
Odhner, N. 1912. Northern and Arctic invertebrates in the collection of the Swedish State Museum (Rijksmuseum), V. Prosobranchia, I. Diotocardia Kunglige Svenska Vetenskapsakademiens Handlingar 48:(10):l-93, 7 pis.
Odhner, N. 1915. Die Molkiskenfauna des Eisfjords Zoolo- gische Ergebnisse der Schwedischen Expedition nach Spitsbergen, 1908, Pt 2, Kunglige Svenska N'etenskap- sakademiens Handlingar 54:1-274, 1 pi., maps.
Philippi, R. A. 1846-56. Die Kreiselschnecken oder Tro- choideen. SystematischesConchylien-Cabinet von Martini und Chemnitz, Vol. 2(3): 1-372,' 49 pis
Pilsbrv, H. A. 1889-90. Trochidae. Manual of Conchology, Vol 11, 519 p., 67 pis.
Posselt, H. J. 1895. Ostgrbnlandske Mollusker. Meddelelser cm Gronland 19:61-93.
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Reeve, L. 1842. Conchologica Systematica, Vol, 2. Longman,
Brown, Cireen and Longmans, London, 337 p., pis. 130-
300. Reeve, L. 1855. Account of the shells collected by Captain
Sir Edward Belcher, C.B., north of Beechey Island. In:
Belcher, E., The last of the arctic voyages, Vol. 2, p. 392-
399, pis, 22-23. Rehder, HA. 1981. The Audubon Society field guide to North
American sea shells. Alfred A. Knopf, New York. 894 p.,
705 figures. Sars, G. O. 1878. Mollusca Regionis Articae Norvegiae Chris-
tiania .\iii + 3 + 466 p , 18 pis . 1 map. Sneli, J. A. 1975. Mollusca Prosobranchia Forgjellesnegler
Nordiske Marine Arter. Universitetsforlaget, Trondheim.
55 p., 60 figures.
Sowerby, G. B. 1838a. A descriptive catalogue of the species of Leach's genus Margarita. The Malacological and Con- chological cal Magazine 1 :23-27.
Sowerby, G. B. 1838b A catalogue of all the species hitherto known of Leach's genus Margarita. Conchological illus- trations, pts. 132-134, 3 pis.
Sowerby II, G. B. 1878. Monograph of the genus Margarita. Conchologica Iconica, Vol. 20, pis. 1-3.
Thorson, G. 1941. Marine Gastropoda: Prosobranchiata. The Zoology of Iceland 4(60): 1-150.
Thorson, G. 1944. Marine Gastropoda: Prosobranchiata. The East Greenland Meddelelser om Gronland 121(13)1-181.
Winkworth, R 1932. The British marine Mollusca journal of Conchology 19:211-252.
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-
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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
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\
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.
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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
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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.
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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.
LITERATURE CITED
Beu, A. G. 1985. .\ classification and catalogue of li\ ing world liaiifllidac (=( a nialiidae) and Bursidae. .\nierican Con- rliologist (Bulletin Concliologists of America) 13(4):.55-66.
Beu, A. G. 1987. Taxonomy of gastropods of the families Ranellidae (=Cymatiidae) and Bursidae. Part 2. Descrip- tion of 14 new modern Indo-West Pacific species and sub- species, with revisions of related taxa New Zealand journal of Zoology l:^(;5):27.'5-.3.55.
Beu. .\ G 1988 Ta\onom\ of gastropods of the iamilies Ranellidae (=(a matiidae) and Bursidae. Part .5. EarK his- tor\ of the families, with four new genera and recognition
W. K. Emerson and W. E. Sage, III, 1990
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of the family Personidae. In: Grant-Mackie, J. A., K. Ma- suda, K. Mori, and K Ogasawara. (eds ) Professor Tain io Kotaka C^ommemorative Volume on Molluscan Paleon- tology The Saito Gratitude Foundation, Sendai, Japan, p. 69-96.
Clench, W. J. 1945. Some notes on the life and e.xplorations of Hugh Cuming. Occasional Papers on Mollusks, Cam- bridge, M.\ l(3):lT-28.
Dance, S. P. 1986. A history of shell collecting E J Brill, Leiden, i-xv, 1-265 p.
Emerson, W K. and E L. Puffer 1953. A catalogue of the molluscan genus Di.storsio (Gastropoda, Cymatiidae). Pro- ceedings of the Biological Society of Washington 66(16): 93-108.
Fulton, H. C. 1938. Description and figures of new Japanese marine shells. Proceedings of the Malacological Society of London 23(l):55-57.
Gray, J. E. 1854. On the division of ctenobranchous gaster- opodus Mollusca into larger groups and families. Proceed- ings of the Zoological Society of London, [for 1853] 21: 32-44.
Iredale, T. 1913 The generic name to be used for Murex tritonis Linne. The Nautilus 27(5):55-56.
Lamarck. J. B. .\. M de. 1803. Suite des Memoires sur les fossiles des environs de Paris. Annales Museum Histoire Naturelle, Paris 2:217-227.
Lamarck, J B. A. M. de. 1816. Tableau encyclopedique et Methodique des Trois Regnes de la Nature, Paris, Liste des objects representes. Atlas 3, pis. 391-488.
Lewis, H. 1972. Notes on the genus Distorsio (Cymatiidae) with descriptions of new species. The Nautilus 86(2-4): 27-50.
Linne, C. von. 1758. Systema naturae per regna tria naturae. Editio decima, reformata. Stockholm 1:1-824.
Manoja, R. E. 1989. More on genus Distorsio Roding, 1798. La Conchiglia, Rome 21(242-245):28, 29.
Nickles, M. 1950. Mollusques testaces marines de la Cote occidentale d'Afrique. In: Paul Lechevalier (ed.) Manuels Oest-Africains 2:i-x, 1-269 p
Oyama, K 1958. The molluscan shells. Science and Photog- raphy Club, Kokvo, II, Distorsio. pi. 1, figs. 1-12; pi. 2, figs. i-12.
Parth, M. 1989. Brief notes on the genus Distorsio Roeding 1798, and description of a new species. La Conchiglia, Rome 21(233-236):52-57.
Peluch. E. J. 1987. New Caribbean molluscan faunas. The Coastal Education and Research F"oundation, Charlottes- ville, \'A, 154 p.. Addendum 4 p.
Puffer, E. L. 1953. Distorsio reticulata vs. Distorsio rlathrata in the West Indies. Proceedings of the Biological Society of Washington, 66(17): 109-124.
Reeve, L. A. 1844a. Monograph of the genus Triton. Con- chologica Iconia:, or illustration of the shells of molluscous animals. Reeve Brothers, London, 2, Triton text and 20 pis. [pi. 12, May 1844].
Reeve, L. A. 1844b. Descriptions of new species of Tritons, collected chiefly by H. Cuming, Esq. in the Philippine Islands. Proceedings of the Zoological Society of London 12(137):110-122[Dec., 1844].
Roissy de, F. 1805, In: Buffon, G. L. L. and C. S. Sonnini, Histoire Naturelle, Generale et Particuliere, Des Mol- lusques, Animaux sans Vertebres et a Sang Blanc. Paris, 6: 1-480 [pages 56, 57, reprint of description of Murex can- cellinus Lamarck (1803:225)].
Roding, P. F. 1798. Museum Boltenianum . . . pars secunda continens Conchylia. Hamburg, i-vii, 1-199 p.
Springsteen, F. J. 1981. The genus Distorsio Roeding, 1798 in the Philippines. Carfel Philippine Shell News, Manila, 3(4):1, 3, 6-10.
Springsteen, F. J. 1984. Distorsio update. Carfel Philippine Shell News, Manila 6(1 ):5-ll.
Springsteen, F. J. 1985. Distorsio decipiens (Reeve, 1844) a valid biospecies rediscovered. Carfel Philippine Shell News, Manila 7(5):3-5.
Springsteen, F. J. and F. M. Leobrera. 1986. Shells of the Philippines. Carfel Seashell Museum, Manila, 377 p.
Tryon, G. W., Jr. 1880[-1881]. Family Tritonidae, Manual of Conchology. Philadelphia, Series 1, 3(5-6):l-97 [p. 1- 64, 1880; 6.5-128, 1881].
Valenciennes, A. 1832. Coquilles univalves de I'Amerique Equinoxiale, recueillies pendant le voyage de MM. de Humboldt et Bonpland. In: von Humboldt, F. H. A. and A. J. A. Bonpland (eds). Voyage aux regions equinoxiales du comparee, Paris, 2:263-339, pi. 57.
Wagner, R. L. and R. T. Abbott. 1978. Wagner and Abbott's Standard Catalog of Shells. American Malacologists, 3rd ed Family Cymatiidae Iredale. 1913, 12-801, 802.
Wolfe, C. 1976. Sorting out two look-alike Distorsios. Hawai- ian Shell News 24(9):12.
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,
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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
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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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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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TH E f7 N AUTI LUS
Supplement 1
(Issued with Volume JOS)
March 6, 1990
ISSN -0028- 1344
Wlarine Biological Laboratory ► LIBRARY [
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.
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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.
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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.
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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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Hoeninghaus, F. W. 1831. Versuch einer geognostischen Eintheilung seiner Versteinerung-Sammlung. Jahrbuch fiir Mineralogie, Geognosie, Geologie und Petrefaktenkunde, Jahrgang"2: 132- 17^3.
Hoemes - also see Homes.'
Hoemes, R. 1875. Die Fauna des Schliers von Ottnang. Jahrbuch der Kaiseriich-Koniglichen Geologischen Reich- sanstalt 25:333-400. pis. 10-15.
Hoemes, R. and M. Auinger. 1879-91. Die Gasteropoden der Meeres-Ablagerungen der ersten und zweiten Miocanen Mediterran-Stufe in der Osterreichisch-Ungarischen Mon- archic. Abhandlungen der Kaiseriich-Koniglichen Geolo- gischen Reichsanstalt 12:1-382, pis. 1-50. [1-52, pis. 1-6, 1879; 53-112, pis. 7-12, 1880; 113-152, pis. 13-16, 1882 153-192, pis. 17-22, 1884; 193-232, pis. 23-28, 1885 233-282, pis. 29-36, 1890; 283-330, pis. 37-42, 1891 331-382, pis. 43-50, 1891.]
Holzapfel, E. 1888. Die Mollusken der Aachener Kreide. Palaeontographica 34:29-180, pis. 4-20.
Hdlzl, O. 1958. Die Mollusken-Fauna des oberbayerischen Burdigals. Geologica Bavarica 38:1-348, pis. 1-22.
Homes - also see Hoemes.'
Homes, M. 1848. Verzeichniss der Fossil-reste aus 135 Fundorten des Tertiar-Beckens von Wien. Wilhelm Braumiiller, Wien. 43 p.
Homes, M. 1851-56. Die Fossilen Mollusken des Tertiar- Beckens von Wien. 1. Univalven. Abhandlungen der Kaiseriich-Koniglichen Geologischen Reichsanstalt 3:1- 736, pis. 1-52. [1-42, pis. 1-5, 1851; 43-184, pis. 6-15, 1852; 185-296, pis. 16-32, 1853; 297-384, pis. 33-40, 1854; 385-460, pis. 41-45, 1855; 461-736, pis. 46-52, 1856.]
Hutton, F. W. 1873. Catalogue of the marine Mollusca of New Zealand, with diagnoses of the species. Colonial Museum and Geological Survey Department. Wellington, xx + 116 p., 1 pl.
Hutton, F. W. 1877. Descriptions of some new Tertiary Mollusca from Canterbury. Transactions and Proceedings of the New Zealand Institute 9:593-598, pl. 16.
Hutton, F. W. 1885. Descriptions of new Tertiary shells. Transactions and Proceedings of the New Zealand Insti- tute 17:313-332, pl. 18.
Icke, H. and K. Martin. 1907. Over tertiaire en kwartaire vormingen van het eiland Nias. Sammlungen des Geolo- gischen Reichs-Museums in Leiden 8(3-4):204-252, pis. 14-18.
Ihering - see von Ihering
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Iredale, T. 1929a. Strange molluscs in Sydney Harbour. The Australian Zoologist 5:337-352, pis. 37-38. [Introductory remarks and plates reprinted in Sydney Harbour Trust Officers' Joumal 5(2):20-24, pis. A & B, July, 1929.]
Iredale, T. 1929b. Queensland moUuscan notes, no. 1. Mem- oirs of the Queensland Museum 9(3):261-297, pis. 30-31.
Iredale, T. 1930. Queensland molluscan notes, no. 2. Memoirs of the Queensland Museum 10(l):73-88, pl. 9.
Iredale, T. 1936. Australian molluscan notes: No. 2. Records of the Australian Museum 19:267-340, pis. 20-24.
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'Moriz Homes and Rudolph Hoemes. although father and son, used different orthography for their surname. This difference is consistent in all their published work which we have seen.
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Mayer, K. 1861b. Die Faunula des marinen Sandsteines von Kleinkuhren bei Konigsberg. Vierteljahrsschrift der Nalur- forschenden Gesellschafl in Zurich 6:109-123.
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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.
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