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Full text of "The Nautilus"

MBL/WHOl 



THE NAUTILUS 



Volume 106 
1992-1993 




AUTHOR INDEX 



Askew, T. M 130 

Bailey, J. F 60 

RiELER, R 15 

Ross, K. J 21 

Emerson, \V. K 39, 147 

eversole, .\. g 119 

Goodfriend, G. a 55 

CioODSELL, J. G. 119 

I I.^HASEWYCH, M. G 1, 43, 130 

Healy, J. M 1 

Hunter, R. U 60 

KooL, S. P 21 



Marshall, B .^ 24 

McLean. J. H 39,115,125 

Oleinik, a. E 137 

Petit, RE 43 

Petuch, E. J 68 

Potter, E, 72 

QuiNN. J. E, Jr 50. 77 

Rehder, ha 123, 127 

Rex, ma 72 

Tucker, J K 76 

Verhecken, a 43 



NEW TAXA PROPOSED IN VOLUME 106 (1992-1993) 



n. sp. 



GASTROPODA 

Pleurotoniariidae 

Pcrotrochtis maureri Harasew\cli & Askew, 1993, n 
C'lypeosectidae 

Pseitdorimula inidatlaniica McLean, 1992 
Trocliidae 

Calliostomii allfrninn Ouiiin, 1992, n. sp. 
Calliostoma argcnlinn Ouiiin, 1992, n. sp. 
CallioHloma atlanluidcs Quinn, 1992, n. sp. 
C'.aUiostoina aiilicurn Quitin, 1992, n. sp. 
CUilliosliima axclolsaoni Quinn, 1992, n. n. 
C'allioslotrm hcnmidcnsc Qninn, 1992, n. sp- 
CaUiofitunia brunncopictum Quinn, 1992, n. sp 
Calliostoma cnidophiluni Quinn, 1992, n. sp. 
Calliostoma coronatum Quinn, 1992, n. sp. 
Calliostoma citbensc Quinn, 1992, n. sp 
Calliostoma debilc Quinn, 1992, n. sp 
Calliostoma denlalum Quinn, 1992, n. sp. 
Calliostoma furosnm Quinn, 1992, n. sp. . . 
Calliostoma gncsti Quinn, 1992, n. sp. ... 
Calliostoma hilarc Quiiui, 1992, n. sp. 
Calliostoma hirlum Quiiui, 1992, n. sp. 
Calliostoma moscaltetlii Quinn, 1992, n. sp. 
Calliostoma purpureiim Quinn, 1992, n. sp. 
Calliostoma rota Quinn, 1992, n. sp. 
Calliostoma rude Quiiwi, 1992, n. sp 
Calliostoma rngosum Quinn, 1992, n. sp. 
CUilliostoma sralrnum Quiiui, 1992, n. sp. 
Calliostoma snirra Quinn. 1992, n. sp. 
Calliostoma semisuavc Quinn, 1992, n sp 
Calliostoma serratulum Quinn, 1992, n. sp. 
Calliostoma tenebrosum Quinn, 1992, n. sp. 
Calliostoma vinosum Quinn, 1992, n. sp 
Calliostoma viscardii Quinn, 1992^ n. sp. 
Solariella cristata Quiiui, 1992, n sp. 
Solariella quadricincta Quinn, 1992, 
Solariella slaminea Quinn, 1992, ii. sp 



130 
115 

96 

103 

102 
97 

105 
86 
85 
83 

102 
93 
90 
87 
95 

106 
78 
97 
87 
95 

100 
83 

103 
92 

110 
85 
78 
87 
96 
85 
52 
50 
53 



Colunibellidae 

Cotonopsis monfilsi Emerson, 1993, n. sp 147 

Faseiolariidae 

Buccinofiisus patuxcntensis Petuch, 1993, n. sp. 165 

Meloiigenidae 

Bustjcotyptis choptankensis Petuch, 1993, n. sp. 166 

Tiirrifulgur manjlandicus Petuch, 1993, n. sp. 166 

Turrifulgur prunicola Petuch, 1993, n. sp. 167 

Thaidinae 

Ecphora (Ecphora) chesapeakensis Petuch, 1992, n. sp. 68 

Ecphora (Ecphora) turneri Petuch, 1992, n. sp. 70 

Ecphora (Trisecphora) scientisensis Petuch, 1992, n. sp. 70 

Ecphorosycon lindajoyceac Petucli, 1993, n. sp. 164 

Muricidae 

Patuxentrophon Petuch, 1993, n gen 165 

Volutidae 

Ftilgoraria (Musashia) novoilpinica Oleinik, 1993, n. sp. 138 

Ftilgoraria (Musashia) genuata Oleinik, 1993, n sp. 140 

Fnlgoraria (Musashia) cordata Oleinik, 1993, n. sp. 140 

Fulgoraria (Musashia) lilitschikensis Oleinik, 1993, n. sp. 140 

Harpidae 

Harpa cabriti Relider, 1992 n. n 124 

Harpa goodtvini Rehder, 1993, n, sp 127 

Gancellariidae 

Trilonoharpa leali Harasew \ch. Petit & 

\ erhecken, 1992, n. sp 45 
Cancellaria petuchi Harasewych, Petit & Verhecken, 

1992, ii..sp ' 47 

Turridae 

Buridrillia dcrmjorum Emerson & McLean, 1992, n. sp. 39 

Calverturris Petuch. 1993, n. gen. 167 

Calverturris schmidti Petuch, 1993, n. sp 168 

Drillia macleani Tucker, 1992, n. n 76 

Transmariaturris Petuch, 1993, n. gen 168 

Helminthoglyptidae 

Hemitrochus bowdcncnsis (Jncidfriend. 1992, n. sp. 55 



rHE NAUTILUS 



Volume 106, Number 1 
February 27, 1992 
ISSN 0028-1344 

A quarterly devoted 
to malacology. 



Dry 






MR 6 my f 

i 

^^' Hole, Mass. J 




EDITOR-IN-CHIEF 
Dr. M. G. Harasewych 
Division of Mollusks 
National Museum of 
Natural History 
Smithsonian Institution 
Washington, DC 20560 

ASSOCIATE EDITOR 
Dr. R. Tucker Abbott 
American Malacologists, Inc. 
P.O. Box 2255 
Melbourne, FL 32902 



CONSULTING EDITORS 
Dr. Rudiger Bieler 
Department of Invertebrates 
Field Museum of 
Natural History 
Chicago, IL 60605 

Dr. Robert T. Dillon, Jr. 
Department of Biology 
College of Charleston 
Charleston, SC 29424 

Dr. William K. Emerson 

Department of Living Invertebrates 

The American Museum of Natural 

History 

New York, NY 10024 

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 Zoology 
Harvard University 
Cambridge, MA 02138 

Ms. Paula M. Mikkelsen 
Harbor Branch Oceanographic 
Institution, Inc. 
Ft. Pierce, FL 33450 

Dr. Donald R. Moore 

Division of Marine Geology 

and Geophysics 

Rosenstiel School of Marine and 

Atmospheric Science 

University of Miami 

4600 Rickenbacker Causeway 

Miami, FL 33149 

Dr. Gustav Paulay 
Marine Laboratory 
University of Guam 
Mangilao, Guam 96923 

Mr. Richard E. Petit 

P.O. Box 30 

North Myrtle Beach, SC 29582 

Dr. Edward J. Petuch 
Department of Geology 
Florida Atlantic University 
Boca Raton, FL 33431 



Dr. David H. Stansbery 
Museum of Zoology 
The Ohio State University 
Columbus, OH 43210 

Dr. Ruth D. Turner 
Department of Mollusks 
Museum of Comparative Zoology 
Harvard University 
Cambridge, MA 02138 

Dr. Geerat J. Vermeij 
Department of Geology 
University of California at Davis 
Davis, CA 95616 



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THEf7NAUTILUS 



CONTENTS 




e 106, Number I 

Februani 27 , 1992 

ISSN 0028-1344 



John M. Healy 
!V1.G. Harasewvch 



Spermatogenesis in Perotrochus quoyanus (Fischer 6i 

Rernardi) (Gastropoda: Pleurotomariidae) 1 



Riidiger Bieler 


Tcnagodiis of Siliquaria'-' L'rira\elini; taxonornic confusion 
in marine "worm-snails iCerithioidea: Sili(iuariidae) 


15 








Silvard P. Kool 


Niirclla Hodinn, 1798 ((Gastropoda: Muricidae) t\pe species 


21 


Kenneth J. Boss 






Bruce A. .Marshall 


A revision ot the Recent species of Eudoliuni Uall, 1889 
(Gastropoda: Toiinoidea) 


24 








VCilliam K. Emerson 
James H. McLean 


Biiridrillia dcroijorurn. new species from the Galapagos 

Islands, a li\inu record of a Neogene Turrid genus 


39 









THE NAUTILUS 106(1):1-14, 1992 



Page 1 



Spermatogenesis in Perot rochus quoyanus (Fischer & Bernardi) 
(Gastropoda: Pleurotomariidae) 



John IVI. Healy 

Departnu'iil of Zoology 
The University of Queensland 
Si. Lueia, Brisbane 
Queensland, ALSTRALIA 4()H7 



M.G. Harasewych 

Department of Invertebrate Zoology 
National Museum of Natural History 
Smithsonian Institution 
Washington, DC: 20560 USA 



ABSTRACT 

The male reproducti\e system, ultrastructure of spermatozoa 
and spermatogenesis are described for the pleurotomariid Pcr- 
otrochus quotjanus (Fischer i Bernardi) CIross morpholog\ of 
the male reproductive system of P. qtiotjanus agrees in all 
essential details with that of Mikadot rochus beyrichii. In all 
features, spermatozoa of Perotrochus quoyanm closely resem- 
ble those of Perotrochus westralis Whitehead, 1987, as well as 
spermatozoa of certain members of the Trochoidea (Trochidae, 
Liotiidae) Spermatozoa of P. quoyanus have a conical acro- 
somal vesicle with a finely ridged anterior layer, a short, rod- 
shaped nucleus with numerous lacunae, a midpiece consisting 
of five (rareK four) mitochondria surrounding a pair of cen- 
trioles, a rootlet cotmecting the centrioles and axonenie to the 
nucleus, and a flagelluni (55-58 ^m long) that is continuous 
with the distal centriole. Investigated species of Haliotidae and 
Scissurellidae {Sinezona sp.) differ from Perotrochus in acro- 
somal substructure, and, in the case of Sinczuna. also in mid- 
piece and nuclear morpholog\ 

Key Words. Spermatozoa, Spermatogenesis, Mollusca, Gas- 
tropoda, Pleurotomariidae, Perotrochus, male reproductive 
tract 



INTRODUCTION 

Living species of the Pleurotomariidae have a host of 
primitive gastropod features including a prominent la- 
bial shell slit as well as paired gills, auricles, osphradia, 
kidneys, and hypobranchial glands (Woodward, 1901; 
Bouvier & Fischer, 1902; Fret'ter, 1966; Hickman, 1984; 
Haszprunar, 1988). The Haliotidae and Scissurellidae, 
which share these features and classically have been as- 
signed to the Pleurotomarioidea, are now considered suf- 
ficiently different from the Pleurotomariidae to warrant 
their placement into separate superfamilies, while the 
Pleurotomarioidea is considered most closely related to 
the Trochoidea based on the shared presence of a glan- 
dular urinogenital duct in females (for discussion see 
Haszprunar, 1988, 1989; McLean, 1989). Basic features 
of pleurotomariid anatomy, including radular morphol- 
ogy, have been known for more than a century (Dall, 



1889; Bouvier & Fischer, 1899, 1902; Pelseneer, 1899; 
Woodward, 1901; Fretter, 1964, 1966), but it is only in 
recent years that the advent of deep-sea submersible craft 
has allowed the biology and habitat of living specimens 
to be studied in detail and in situ (Yonge, 1973; Hara- 
sewych et ai, 1988, 1992). 

The field of comparative spermatology has, over the 
last twenty years, contributed greatly to the resolution 
of taxonomic and phylogenetic problems in numerous 
phyla (Baccetti & Afzelius, 1976; Wirth, 1984; Jamieson, 
1987), including the Mollusca (Nishiwaki, 1964; Popham, 
1979; Giusti, 1971; Kohnert & Storch, 1984a,b; Koike, 
1985; Healy, 1983, 1986, 1988a; Hodgson et ai. 1988). 
Among the Gastropoda, studies of archaeogastropod (,s-./. ) 
spermatozoa and spermiogenesis (Kohnert & Storch, 1983; 
Azevedo et ai, 1985; Koike, 1985; Hodgson & Bernard, 
1988; Healy, 1988b, 1989, 1990a,b) are becoming in- 
creasingly important since it is from this broad assem- 
blage that origins for the caenogastropod and euthyneu- 
ran groups are sought (Cox, 1960; Ponder, 1973; 
Haszprunar, 1988). The recent discovery of pronounced 
sperm dimorphism in the trochoidean Zaiipais laseroni 
Kershaw, 1955, including a multi-tailed, oligopyrene 
paraspermatozoon (Healy, 1990b), has drawn attention 
to the fact that comparatively little is known of the range 
of sperm morphologies existing in the Vetigastropoda. 
Healy (1988b) provided the first ultrastructural infor- 
mation on spermatozoa of the Pleurotomariidae [Pero- 
trochus westralis Whitehead, 1987,' as Plcurotomaria 
ajricana (Tomlin, 1948)], but, because of limitations im- 
posed by the state of preservation of the testes, was unable 
to trace e\ents of spermatogenesis or give substructural 
detail of certain sperm features. L'sing glutaraldehyde- 
fixed testicular material of Perotrochus quoijanus, we 
present the first ultrastructural study of sperm devel- 
opment in a pleurotomariid gastropod. 



' For a discussion of the nomenclature of this species, see 
Wagner and O>onians (1990). 



Page 2 



THE NAUTILUS, Vol. 106, No. 1 



MATERIAL AND METHODS 

Three male specimens of the pleurotomariid Perot rochus 
quoyanus (Fischer & Bernardi, 1856) were collect ed us- 
ing the research submersible JOHNSON-SEA-LINK II, 
1.03 nautical miles west of Ilets-a-Goyaves. off Basse 
Terre, Guadeloupe, West Indies' (16°10'33"N, 
6r49'00"W) at a depth of 350-360 m. Specimens were 
maintained in refrigerated aquaria for six days prior to 
cracking the shells and excising the testes. For scanning 
electron microscopy (SEM), samples were prepared by 
teasing apart sections of fresh testes in filtered seawater, 
transferring droplets of sperm suspension to coverslips, 
and fixing with glutaraldehyde vapor (25% glutaralde- 
hyde in a covered petri dish). The coverslips were passed 
through a graded acetone series (20-100%), critical-point 
dried, and coated with gold-palladium. The sperm were 
examined using a Hitachi S-570 SEM at an accelerating 
voltage of 10 kv. Measurements are based on SEM pho- 
tographs of sperm and calibration grids of standard size 
(2160 lines/mm at 15,000 X for acrosomes, nuclei, and 
mitochondria, 19.7 lines/mm at 1,500 X for tails). For 
transmission electron microscopy (TEM), l-2mm^ pieces 
of testicular tissue were fixed with 5% glutaraldehyde in 
0.2 M cacodylate buffer and shipped to the senior author. 
Upon arrival, samples were further fixed in cold 3% 0.2M 
cacodylate-buffered glutaraldehyde and washed thor- 
oughly in cacodylate buffer before being placed into a 
1% solution of osmium tetroxide (prepared in 0.2M cac- 
odylate buffer) for two hours. Tissues were again rinsed 
in buffer, then dehydrated using an ascending series of 
ethanols (20-100%). Spurr s epoxy resin was used to em- 
bed the tissues (Spurr, 1969). Ultrathin sections were cut 
with an LKB IV Ultratome, collected on uncoated 200- 
mesh copper grids, and stained using either the double 
lead stain of Daddow (1983) or a single leatl procedure 
(20 minutes uranyl acetate, 10 minutes lead citrate). Sec- 
tions were examined using a Hitachi 300 transmission 
electron microscope operated at 75 kV. Remaining soft 
tissues were fixed in 10% formaldehyde in seawater and 
transferred to 70% ethanol for dissection. Shell fragments 
retained as voucher specimens are housed in the National 
Museum of Natural Historv, Smithsonian Institution 
(USNM 878154). 



RESULTS 

Male Reproductive Sy.stem 

The mustard-colored testis (fig. 1, te) lines the right wall 
of the digestive gland (fig. 1, dg) , and empties into a 
thin-walled testicular duct (fig. 1, td) situated ventral to 
both these organs. This duct becomes tubular along the 
ventral surface of the stomach (fig. 1, sto) and continues 
anteriorly, emptying (fig. 1, ga) into the ureter portion 
of tlie right kidney (fig. 1, u) anterior and to the right 
of the opening (fig. 1, o) of the anterior lobe of the right 
kidne\ (fig. 1, ark), which is situated in the cephalic 
hemocoel. The ureter/ urinogcnital duct (figs. 1. 2, u) 
runs anteriorly along the roof of the mantle cavity to the 



Sto 




rav 



rko 




Fipure§ 1-2. Male reproductive tract of Perotrochus qiioy- 
(titiis (Fischer & liernardi), I. Diagrammatic representation 
of male reproductive system, viewed from right side. Walls of 
pericardium and right kidney removed to reveal contents 2. 
Transverse section midway along paiiial gonoduct, viewed from 
anterior, ark, anterior lobe of right kidney; dg, digestive gland; 
ga, genital aperture; Ik, left kidney; o, opening of anterior lobe 
of right kidney; pc, pericardium; r, rectum; rav. right afferent 
liranchial vessel; rk, right kidney; rko, right kidney opening; 
sto, stomach; td, testicular duct; te. testis; u, urinogenilal duct. 



right of the rectum (figs. 1, 2, r), envelops the right 
afferent branchial vessel (figs. 1,2, rav), and drains into 
the mantle cavity through a transversely oriented right 
kidney opening (fig. 1, rko), approximately 1/4 of the 
distance from the rear of the mantle cavity to the rear 
of the mantle slit The urinogenital ducts of all three 
individuals lacked a glandular lining. 

The testes of two animals were full of mature sper- 
matozoa, while that of the third animal were almost 
entirely spent. .Although onl\ scattered groups of devel- 
oping cells remained, we were able to identif>- basic 
features of spermatogonia, spermatocytes and sperma- 
tids. 

Mature Testic;ular Spermatozoa (SKM observations) 

Spermatozoa of Perotrochus quoyanus consist of a con- 
ical acrosomal complex (fig. 4, a), a rod-shaped nucleus 
(fig 4, ii), a cluster of five, equal-sized, spherical mito- 
chondria (fig. 4, m, fig. 5, mp) at the base of the nucleus, 
and a single 55-58 ^m long Dagellum (figs. 3-4, f, table 



J. M. Healy and M. G. Harasewych, 1992 



Page 3 




Figures 3-5. Perotrochus quoyanus- Mature testicular sperm, SEM. 3. Two spermatozoa including entire Qagella (f). 4,5. 
Acrosome (a), nucleus (n) and mitochondria (m) of two spermatozoa. Nuclear lacuna (nl) and detached midpiece (mp) consisting 
of five mitochondria visible in figure 5. 
Scale bars: 3=10 ^m; 4,5 = 2 /um. 



1). The acrosomal complex (externalK, the acrosomal 
vesicle proper) is approximately 1.15 ^m long, tapers 
slightly at contact with the nucleus, and has a maximum 
diameter of 1.18 //m (figs. 4,5, table 1). The nucleus 
measures 3.7 ^m in length, is broadest posteriorly, with 
a maximum diameter of 1.4 nm. Irregular indentations 
on the nuclear surface (Bg. 5, nl) can be correlated by 
TEM with nuclear lacunae (figs. 6, 7, 11, nl) occurring 
beneath the nuclear and plasma membranes. These in- 
dentations are not, therefore, nuclear pores. Spherical 
mitochondria (diameter 0.8 /im) obscure the attachment 
site of the Dagellum. The flagellum narrows markedly 
towards its insertion point within the midpiece (figs. 
13,15). 

Mature Testicular Spermatozoa (TEM observations) 

Acrosome: The acrosomal vesicle is broadly conical, 
with a rounded anterior surface and flattened basal sur- 
face (fig. 7, av). The vesicle has a length of 0.90-0.93 
nm and the maximum diameter of 1.28 ixm at its base 
is wider than the apex of the nucleus (figs. 7,8, n). A 



deep, narrow invagination extends anteriorly from the 
base of the vesicle and is filled with a diffuse, faintly 
fibrous material (figs. 8, 10, sm). Some sections clearly 
indicate an eccentric, slightly angular alignment for the 
invagination relative to the sperm longitudinal axis (figs. 
8, 10). Beneath the anterior face of the acrosomal vesicle 
is an electron-lucent layer containing fine ridges with a 
periodicity of 12-14nm (figs. 7,9, rl). A similarly electron- 
lucent layer, lacking discernible ridged substructure, 
forms the basal rim of the acrosomal vesicle (fig. 7, br). 
A loose, fibrous deposit of subacrosomal material occupies 
the space between the base of the acrosomal vesicle and 
the nuclear apex (fig. 8, sm). 

Nucleus: The mature nucleus (fig. 6, n) is short (3.7 ^m) 
and almost cylindrical, with a shallow depression ante- 
riorly (figs. 7, 8, n) and five (rarely four) shallow de- 
pressions surrounding a centriolar fossa posteriorly (figs. 
14, 15, n). The anterior depression is associated with 
subacrosomal material (fig. 8, sm), while the posterior 
depressions act as sockets for the midpiece mitochondria 
(fig. 15, m). Dense material linking the proximal and 



Page 4 



THE NAUTILUS, Vol. 106, No. 1 



Table 1. Dimensions of mature spermatozoa from SEM ob- 
servations. Linear measurements in fin\. (n = .30, 10 sperm 
from each of three individuals.) 









Standard 








deviation 




Mean 


Range 


(<7) 


Acrosome 








Length 


Lie 


1.01-1.19 


0.07 


Width 


LOS 


1.01-1.18 


0.05 


Nucleus 








Length 


3.67 


3.52-3.78 


0.08 


Width (anterior) 


0.98 


0.95-1.03 


0.02 


Width (posterior) 


L19 


1.13-1.34 


0.08 


Mitochondria 








Diameter 


0.80 


0.68-0.93 


0.09 


Flagellum 








Length 


56,5 


.52.7-61.1 


3.26 



distal centrioles is continuous with a hollow rootlet (figs. 
14, 15, r), the bulbous end of which fills the centriolar 
fossa. Numerous irregularly shaped lacunae (figs. 6, 7, 
11, nl) occur within the nucleus, some of which open 
underneath the nuclear membranes, though not to the 
plasma membrane or cell surface. Nuclear contents are 
highly electron dense and consist of tightly packed fibers 
(diameter 16 nm) set in a finely granular matrix. 

Midpiece: Five (rarely four) spherical (diameter 0.6- 
0.8 ^m) mitochondria (figs. 12, 15, m), each having 
curved, plate-like cristae, surround the proximal and dis- 
tal centrioles to form the sperm midpiece (fig. 12). The 
centrioles (figs. 14, 15, pc, dc), arranged at a 90° angle 
to each other, are hollow, cylindrical structures composed 
of triplet microtubules and emliedded in a pericentriolar 
matrix (triplets often obscured by matrix, see fig. 15 
inset). Nine satellite fibers (figs. 15, 16, sf) connect the 
distal centriole to an annulus (figs. 15,16, an), a ring- 
shaped deposit of material lining the inner surface of the 
plasma membrane. The flagellar axoneme, therefore, is 
anchored to the midpiece and nucleus via the centrioles 
and rootlet as well as by the radial set of satellite fibers. 

Flagellum: The flagellum measures approximately 55- 
58 yuni ill length and consists of a 9 + 2 axoneme enclosed 
by the plasma membrane (figs. 15, f; 17). Many sper- 
matozoa were observed with an angularly offset flagellar- 
centriolar apparatus (fig. 13). This misalignment could 
be due to tight packing of sperm within the testis or even 
slight immaturity, since our .SEM observations on free 



sperm show a normal, posteriorly projecting flagellum 
(figs. 3-5). Occasionally, a dense body is enclosed with 
the axoneme by the plasma membrane (fig. 17, db). Its 
position along the flagellum could not be determined. 
Further study is required to determine whether this 
structure is a true sperm feature of P. quoyanus or an 
artifact of fixation. In the distal region of the flagellum, 
the 9 + 2 substructure of the axoneme degenerates into 
singlet microtubules (fig. 17, arrow). 

Spermatogenesis 

Spermatogenic cells present within the testis consisted 
principally of isolated clumps of spermatocytes and sper- 
matids (fig. 18, spc, spt). Most of the testis space in ripe 
males was found to be almost totally occupied by tightly 
packed mature spermatozoa. To some extent the process 
of reconstructing events of spermatogenesis was ham- 
pered by the occurrence of many abnormally developing 
spermatocytes and spermatids. The morphology and pos- 
sible significance of these cells is treated in the final 
section of these results. 

Spermatogonia: Spermatogonia (fig. 19) were only rare- 
ly observed. They can be distinguished from spermato- 
cytes and spermatids by their oblong, usualK lobulate 
nucleus (fig. 19, n; length 6.0 ^m, breadth 4.0 /um), prom- 
inent nucleolus (fig. 19, nc; diameter 0.7 nm), well-de- 
veloped nuclear pores (fig. 19, arrows), numerous small 
mitochondria (fig. 19, m; diameter 0.3-0.4 ^m), and more 
extensive cytoplasm. Endoplasmic reticular cisternae, 
where visible, are scattered and poorly developed. The 
presence of centrioles and Golgi complex could not be 
confirmed in the limited number of observed cells. 

Spermatocytes: Spermatocytes (fig. 20) have a spherical 
to ovoid nucleus (fig. 20, n; diameter 4.0-4.5 ^m) that 
appears to lack either a nucleolus or prominent nuclear 
pores. The small electron-dense patches visible in many 
cells (fig. 20, arrowheads) ma\ be sites of s\naptinemal 
complexes, although these structures are more easily dis- 
cerned in moribund spermatocytes that have partially 
lost nuclear contents (fig. 38, arrows). Mitochondria (fig. 
20, m; diameter 0.6-0.75 jum) markedK larger than those 
of spermatogonia are pressed slightK into the surface of 
the nucleus. Highly electron-dense proacrosomal vesicles 
(fig. 20, pav; diameter 0.1-0.2 ^l^\) of Golgian origin are 
foimd throughout the cytoplasm. The axoneme (fig. 21, 
ax) de\elops intracellularly from one of a pair of or- 
thogonally arranged centrioles (fig. 21, pc, dc) positioned 
close to the concave face of the Golgi complex (fig. 21, 
G). Even at this early stage in axoneme formation, sat- 
ellite fibers (fig. 21, sf) are associated with the future 



Figures 6-17. Perotrochus quoyanus. Mature testicular sperm, TEM 6. Acrosome (a), nucleus (n), nuclear lacunae (nl), and 
mitochondria (m) of two spermatozoa. 7,8. Acrosomal vesicle (av) showing ridged layer (rl), basal rim (br), subacrosomal material 
(sm), and apex of nucleus (n) with nuclear lacuna (nl). 9. Detail of ridged laser (rl) in acrosome 10. Transverse section throng!) 
acrosomal vesicle showing subacrosomal material (sin). I 1. Transverse section through nucleus showing nuclear lacuna (nl). 12. 
Transverse section ihrougli midpiece, five mitochondria (m) surround the distal centriole (dc) 1.3. ,\ngularl\ offset centriolar (c) — 
flagellum (f) apparatus of a spermatozoon. 14. Detail of centriolar fossa and attached rootJcl \r), proximal (pc) and di.stal centrioles 




•^i -^ 






15 



(dc), and mitochondria (m), 15. Base of nucleus (n), rootlet (r), proximal (pc) and distal (dc) centrioles, satellite fibers (sf ), annulus 
(an), flagelluni (f), and mitochondria (m). Inset: triplet microtubules of proximal centriole (arrowheads). 16. Oblique section 
showing distal centriole and three of nine satellite fibers (sf) attached to annulus (an). 17. Transverse section through flagella. Note 
distal region (right) and dense bod> (db) (left). .Arrow indicates singlet microtubules in distal region of flagelluni. 
Scale bars: 6 = 1 Mm; 7,8,10-17 = 0.25 mi"; 9 = 0. 1 m'". 



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THE NAUTILUS, Vol. 106, No. 1 



distal centriole. Endoplasmic reticular cisternae are poor- 
ly developed. 

Spermatids (Speriiiiogenesis): Spermatids can be di- 
vided into three categories based on the condensed state 
of the nucleus: early cells, middle-stage cells, and ad- 
vanced spermatids. 

In early spermatids the nucleus (fig. 22, n) is spherical 
with pale-staining, fibrous contents. Middle-stage sper- 
matids (figs. 23,24) are distinguished from earlier cells 
liy a marked increase in the electron densit\ of the nu- 
clear fibers, and by a tendency of the mitochondria and 
centrioles to move toward the incipient posterior pole of 
the nucleus. Although multiple proacrosomal vesicles are 
still apparent within the c\ toplasm of middle-stage sper- 
matids (figs. 24, 25, pav), it is during this phase of sper- 
miogenesis that the definitive acrosomal vesicle is formed 
by fusion of proacrosomal vesicles. In advanced sper- 
matids, mitochondria and the acrosomal vesicle come to 
lie in shallow depressions of the nucleus, while the nu- 
cleus itself becomes oblong and its constituent fibers more 
condensed (figs. 26,27,36,37). In addition, the acrosomal 
vesicle undergoes pronounced changes in shape and sub- 
structure. Initially, the acrosomal vesicle is round and 
underlain by a thin disjointed layer of subacrosomal ma- 
terial (fig. 26, sm). As seen in figure 26, the site of first 
contact between the definitive acrosomal vesicle and nu- 
cleus may occur close to where the mitochondria are 
situated. Following attachment of the acrosomal vesicle 
to the condensing nucleus, vesicle contents become dif- 
ferentiated into a cluster of coarse granules (fig. 27, g) 
and a more extensive homogeneous portion (fig. 27, h). 
These granules become partitioned into two deposits that 
occupy anterior and posterior depressions in the homo- 
geneous portion (fig. 28). Subsequently, an invagination 
of the homogeneous portion, but not the acrosomal mem- 
brane, begins to form anteriorly (fig. 29, arrowhead). 
The anterior cluster of granules transforms into a finely 
ridged layer (figs. 28-34, rl). As this layer grows, it ex- 
tends into a deepening invagination of the homogeneous 
portion (figs. 29-31, arrowhead). The posterior cluster of 
granules forms the electron-lucent basal rim of the ac- 
rosomal vesicle. A thin deposit of dense material defines 
the basal region of the acrosomal membrane (figs. 28,29, 
dm). Late in spermiogenesis, the basal invagination of 
the acrosomal vesicle develops and is filled with suba- 
crosomal material (fig. 31, sm). The anterior invagination 
of the homogeneous portion, which is not an invagination 
of the vesicle membrane, and the basal invagination of 
the vesicle are distinct and unconnected structures. The 
anterior invagination ultimately disappears, perhaps by 
a process of eversion, leaving the ridged electron-lucent 
layer (figs. 31,34,35, rl) and a small electron-lucent i)late 
(figs. 31,34,35, asterisk). 

Nuclear lacunae, so clearly apparent in mature testic- 
ular spermatozoa, only become evident in the very last 
stage of spermiogenesis. These spaces are not in contact 
with the exterior of the spermatid. The ccntriolar fo.ssa 
(fig. 36, arrowhead) forms through invagination ol the 
nuclear extension that lies between the posteriorly po- 



sitioned mitochondria Origins of the pericentriolar ma- 
trix and centriolar rootlet were not determined. Presum- 
ably the centrioles play some role in the growth of these 
structures. 

Aberrant spermiogenic cells: In addition to spermat- 
ogonia, spermatocytes and spermatids, the testes also con- 
tained numerous abnormally developing spermatocytes 
and spermatids. Some of these spermatocytes appear 
moribund (fig. 38). The spermatids, however, are clearly 
recognizable by their angular shape, evidently the result 
of cytoplasmic pressure from adjacent cells (figs. 18, spt; 
39). Nuclear condensation and proacrosomal vesicle pro- 
duction seem to proceed as in normally developing cells. 
Gradually, however, the nucleus becomes oblong then 
angular and ultimately irregular in shape (figs. 39-42). 
Like normal spermatozoa, mature nuclei of the abnor- 
mal, presumably abortive, lines have numerous lacunae 
(fig. 42, nl) and a fibro-granulate substructure (figs. 40- 
42). The proacrosomal vesicles, rather than forming a 
definitive acrosomal vesicle, remain as a clump of un- 
fused entities (Figure 40, inset, pav). Mitochondria, lo- 
cated in depressions of the nucleus, and axonemal profiles 
are often observed in developing and mature aberrant 
spermatozoa (figs. 40, m; 41, ax). The position and num- 
ber of centrioles was not determined. 



DISCUSSION 

Reproductive System: The morphology of the male re- 
productive system of Perotrochus quoyanus agrees in all 
major features with that of Mikadotrochus bcyrichii. the 
only other species of pleurotomariid for which the male 
reproductive system has been documented (Woodward, 
1901). The female reproductive system of pleurotoma- 
riids differs from the male reproductive s\stem in that 
the pallial portion of the right kidney, the urinogenital 
duct, is glandular. To date, only M. beyrichii (Wood- 
ward, 1901) and Perotrochus midas (Fretter, 1966) are 
confirmed to have glandular female urinogenital ducts. 
The duct of the holot\ pe of Perotrochus amabilis (Bayer, 
1963), an "immature' female on the basis of gonadal 
sections, lacked glandular elements, prompting Fretter 
(1964:179) to suggest that this was a young individual 
that had never spawned, and that glands may develop 
in the walls of this duct only as the gonad becomes 
mature. Examination of the shells of more than a dozen 
specimens collected in the intervening decades reveals 
that the holotype of M. arnahilis is among the larger 
specimens known of this species. It is therefore unlikely 
that the holotype is an immature individual. Gonadal 
development of several western Atlantic pleurotomariids 
varies with season (Harasewych, unpublished observa- 
tions), suggestitig that the glandular lining of the uri- 
nogenital duct of females may develop and diminish 
cyclically. As evidenced by the three specimens used in 
this stud) , the urinogenital ducts of male pleurotomariids 
arc not glandular, even during the spawning season. Nev- 
ertheless, absence of a glandular urinogenital duct may 



18 






•*•• 19 









• /' 



s^^ 










'M 








Figure§ 18-24. I'crotrvchus qiioyanus. Spermatogenesis, 18. Survey section of testis showing spermatozoa (spz), developing 
spermatocytes (spc), and advanced spermatids (spt). Arrows indicate aberrant spermatids. 19. Spermatogonium, Note lobulate 
nucleus (n), large nucleolus (nc), nuclear pores (arrows), and numerous small mitochondria (m), 20. Spermatocytes. Note mito- 
chondria (m), nucleus (n), presence of proacrosomal vesicles (pav), and putative synaptinemal complexes (arrowheads). 21. Sper- 
matocyte Golgi complex (G) close to proximal and distal centrioles (pc, dc) and axoneme (ax). Note satellite fibres (sf) associated 
with distal centriole. 22. Early spermatids (spt) with homogeneously granular nuclei (n) and proacrosomal vesicles (pav). 23. 
Middle stage spermatids with very electron dense fibrillar nuclei (n) and mitochondria (m). 24. Middle stage spermatid showing 
pair of centrioles (c), proacrosomal vesicles (pav), and mitochondria (m). 
Scale bars: 18 = 10 ^m; 19,20.22-24 = 1 Mm; 21 = 0.5 Mm. 



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THE NAUTILUS, Vol. 106, No. 1 



y. a; 




FiguiT., 2.> .>.i (.;. Ini:. quuy.iiui.; \k icwiiiif (Ifvelupmeiil. 25. (iruup of proacTo.sonial vesicles (pav) near mitochondrion 

(m) and axoneme (ax). 26. Spermatid with acrosomal vesicle (av) contacting nucleus (n) near mitochondrion (m) Subacrosomal 



J. M. Heal) and M. G. Harasewych. 1992 



Page 9 



not be a sufficient criterion for identifying male speci- 
mens. 

Spermatogenesis: Despite the complicating factor of 
moribund and abnormally developing cells within the 
testis, spermatogenic stages of Perotrochus quoyanus re- 
semble those reported for the Trochoidea (Kohnert & 
Storch, 1983; Azevedo et a/., 19S5; Koike, 1985; Healy, 
1989). Using museum-preserved tissues, Healy (1988b) 
was able to determine that acrosomal development in 
Perotrochus westralis involved the production of mul- 
tiple proacrosomal vesicles. Fusion of proacrosomal ves- 
icles into a definitive acrosomal vesicle has been dem- 
onstrated in many bivalves (Longo & Dornfeld, 1967; 
Kubo, 1977; Bernard & Hodgson, 1985; Hodgson & Ber- 
nard, 1986; Eckelbarger et ai, 1990), and, outside the 
Mollusca, in groups as disparate as the Polychaeta (Fran- 
zen, 1987) and Echinodermata (Dan & Sirakami, 1971; 
Chia & Bickell, 1983). In contrast, acrosome develop- 
ment in patelloidean gastropods centers on the produc- 
tion of a single, electron-lucent vesicle to which small 
vesicles from the Golgi cisternal edges fuse and contrib- 
ute (Hodgson & Bernard, 1988). Our study has discov- 
ered details of acrosome development previously unde- 
scribed in the Vetigastropoda, including the 
differentiation of anterior and posterior extremities of 
the vesicle and formation of fine ridges in the anterior 
electron-lucent layer. There are reasons for believing that 
these events also occur in the Trochoidea. Mature ac- 
rosomes of trochids frequently show anterior and pos- 
terior electron-lucent layers (the anterior layer with ridg- 
es: HeaK & Daddow unpublished). In spermatids of 
Omphalius pfeifferi (Philippi, 1846), the definitive ac- 
rosomal vesicle (Koike, 1985:plate 3D) closely corre- 
sponds to the stage illustrated herein for Perotrochus 
quoyanus (fig. 30). The origin of the subacrosomal ma- 
terial in P. quoyanus and in Giblntla umbilicalis (da 
Costa, 1778) (see Azevedo et ai, 1985) is unknown. A 
Golgian source seems unlikely, as this secretory organelle 
has migrated posteriorly by the time the definitive ac- 
rosomal vesicle has formed (the stage when subacrosomal 
material becomes visible). Possibly, the acrosomal vesicle 
itself is capable of organizing the accretion or polymer- 
ization of extravesicular materials within the cytoplasm. 
Takaichi & Dan (1977) proposed a similar origin for 
subacrosomal material in the pulmonate Euhadra hick- 
onis (Kobelt, 1879). An interesting feature of spermio- 
genesis in Perotrochus quoyanus is the often distant po- 
sitioning of the nuclear-contacted acrosomal vesicle 



relative to this vesicle's final position at the nuclear apex 
(see fig. 26). A comparable situation occurs in the trochid 
Calliotropis glyptus (Watson, 1879) (see Healy, 1989) 
and evidently in the turbinid Lunella granulata (Gmelin, 
1791) (see micrographs of Koike, 1985). In Perotrochus, 
Calliotropis, and Lunella, however, the mature acro- 
.somal vesicle lies at the nuclear apex, indicating that by 
some means, perhaps via nuclear shape change late in 
spermiogenesis or acrosomal movement, the vesicle at- 
tains its final position. 

The pattern of nuclear condensation in Perotrochus 
quoyanus differs from that occurring in the Trochoidea 
in two respects: (1) the heterochromatin forms a ho- 
mogeneous network of dense fibers, whereas the hetero- 
chromatin forms distinct granules in Trochoidea; and (2) 
nuclear lacunae appear only at the last stage of spermatid 
development, whereas the lacunae are well developed 
and visible at earlier stages in Trochoidea (Kohnert & 
Storch, 1983; Azevedo et a!., 1985; Koike, 1985; Healy, 
unpublished data). Unfortunately, no comparative in- 
formation exists on nuclear condensation or, in fact, on 
any aspect of spermiogenesis, in the Haliotidoidea, Scis- 
surelloidea, or Fissurelloidea. Initially thecentriolar fossa 
of Calliotropis glyptus spermatids resemble the mature 
fossa of Perotrochus spp., but late in spermiogenesis, the 
solid rootlet and attached centrioles of C. glyptus become 
drawn into a greatly expanded fossa (Healy, 1989). 

Incorporation of the future flagellar axoneme within 
the cytoplasm of spermatocytes and spermatids in Per- 
otrochus warrants some comment. The same phenom- 
enon can be seen in published micrographs of developing 
spermatids in the trochid Monodonta turbinata (Born, 
1778) (see Kohnert & Storch, 1983) and in the turbinid 
Lunella granulata (see Koike, 1985). Unfortunately, nei- 
ther Kohnert and Storch (1983) nor Koike (1985) offer a 
discussion of this positioning of the axoneme. In sper- 
matids of the caudofoveate Chaetoderma sp., the prox- 
imal and distal centrioles each give rise to an axoneme 
within the cytoplasm (Buckland-Nicks & Chia, 1989). 
Of these two axonemes, only that associated with the 
future distal centriole survives into the mature sperma- 
tozoon. A similar situation has been reported by Eckel- 
barger et al. (1989) in paraspermatozoan development 
of the abyssal sea urchin Phrissocijstis multispina, with 
the exception that both axonemes survive in the mature 
cell. Given the large number of abnormally developing 
spermatids observed in the ripe testes of our specimens 
of Perotrochus quoyanus, it cannot be ruled out that the 
intracellular axoneme in spermatids of this species may 



material (sin) is thin. Arrowheads indicate axonemal profiles. 27. Acrosomal vesicle showing granule cluster (g) and homogeneous 
portion (h) of vesicle contents. Note also subacrosomal material (sm ), 28. Granules (g) distributed in anterior and posterior depressions 
of homogeneous portion (h). Note ridged layer (rl) and basal rim defined by dense material (dm). 29. Beginning of invagination 
(arrowhead) of homogeneous portion (h). 30. Penetration of ridged layer (rl) into deepening invagination (arrowhead) of homo- 
geneous portion. 3 1 . Acrosome of late spermatid showing developing basal invagination of acrosomal vesicle as well as invagination 
of homogeneous portion (arrowhead). Asterisk indicates electron-lucent plate 32-34. Sequence of ridged layer (rl) development 
shown in detail Granule cluster (g). .'Vsterisk indicates electron-lucent plate. 35. Nearly mature acrosome. Electron-lucent plate 
indicated h\ asterisk. Subacrosomal material (sm). 
Scale bars: 25,26 = 0.5 nm. 27-35 = 0.25 ^m. 



Page 10 



THE NAUTILUS, Vol. 106, No. 1 



be an aberrant rather tlian normal feature. Examination 
of testes from animals collected at the commencement 
of the reproductive season should resolve this question. 

Aberrations in spermatogenesis: Few ultrastructural 
studies have dealt w itli the incidence of spermatogenic 
abnormalities in niollusks. Takaichi (1979) detailed ra- 
diation-induced malformations of the mitochondrial 
sheath and nucleus and duplication of the axoneme in 
spermatids of the pulmonale Euhadra hickonis. Dorange 
and Le Peiinec (1989) noted binuclear spermatids and 
angularly dislocated axonemes in late spermatids of Pec- 
ten maximus (Linne, 1758) and regarded these features 
as true aberrancies. O'Foighil (1985) suggested that an- 
gular dislocation of the axoneme in testicular sperm of 
the bivalve Lasaea suhviridU Dall, 1899 could be due to 
slight immaturity. In Perotrochus quoyanus we have 
observed numerous spermatocytes and spermatids that 
were undergoing a form of development clearly different 
from normal spermatogenesis. Leaving aside the phe- 
nomenon of sperm dimorphism (a well-documented and 
'normal' occurrence in many Caenogastropoda — see 
Healy, 1988a for discussion), the irregular shape of the 
condensed nucleus (pressed into shape by abutting cells), 
and the apparent inability of proacrosomal vesicles to 
fuse into a single acrosomal vesicle, strongly suggest that 
these are abnormal cells. Bearing in mind that a certain 
background level of spermatogenic abnormality proba- 
bly exists in many if not most animal species (Bryan & 
Wolosewick,1973; Baccetti & Afzelius, 1976), we believe 
the appearance of aberrant cells in P. quoyanus is prob- 
ably a normal event heralding the end of the annual 
reproductive phase in this species. We base this view on 
the fact that all three males examined were either spent 
or contained principally mature spermatozoa in the testis 
(with isolated pockets of developing and aljnormal sper- 
matogenic stages). 

Spermatozoa: Healy (1988b) has previously drawn at- 
tention to the structural similarities between spermatozoa 
of Perotrochus westralis and those of the Trochoidea, 
particularly Trochidae. Our observations on glutaral- 
dehyde-fixed testis sperm of P. quoyanus have enabled 
us not only to confirm these similarities but also to expand 
on details of the Perotrochus spermatozoon as recon- 
structed by Healy from sea-water formalin/ethanol-pre- 
served material. 

The electron-lucent anterior layer of the Perotrochus 
acro-somal vesicle contains regularly spaced ridges. Sim- 
ilar ridges have elsewhere been observed in the Trochi- 



dae [A!/.s7ro('Of/j/ra roni^ric^a (Lamarck, 1822), Banhivia 
australis (Menke, 1830); Healy & Daddow unpublished] 
and in the liotiid Liotina peronii (Kiener, 1839) (Healy 
& Ponder unpublished). It is interesting to note that the 
acrosomal vesicle of other pleurotomarioidean (s./.) fam- 
ilies (Haliotis spp. — Haliotidae, Lewis et al., 1980; Sakai 
et al., 1982; Sinezona sp. — Scissurellidae, Healy, 1990a) 
lack an electron-lucent anterior layer, whereas in the 
fissurellids Scutiis antipodes Montfort, 1810 and Mont- 
fortuhi conoidea Reeve, 1842, a layer is present but 
exhibits no discernible ridged substructure (Healy, un- 
published). The acrosomal complex in Haliotis and in 
Sinezona also differs from that of Perotrochus b\ having 
an extensive subacrosomal deposit similar to that seen in 
spermatozoa of some fissurellids (Scutus antipodes, 
Montforttila conoidea — see Healy, 1990a for illustra- 
tions) and many bivalve species (see references in Po- 
pham, 1979). The difference in appearance of subacro- 
somal material between Perotrochus westralis (rod-like) 
and P. quoyanus (diffuse, with some evidence of fibrous 
texture), may be due to use of different fixation methods 
(P. westralis — sea water formalin/ethanol; P. quoyan- 
us — glutaraldehyde in cacodylate buffer). Azevedo et al. 
(1985) state that exposure of spermatozoa of Gibbula 
unihilicalis to sea water for five minutes resulted in a 
clearly defined rod (or perforatorium), deri\ed from a 
formerly diffuse subacrosomal substance. It therefore 
seems possible that the subacrosomal rod of P. westralis 
may also be an end product of prolonged exposure to 
sea water. The dense layer of material visible within the 
subacrosomal material in the vicinity of the nuclear apex 
(see figs. 7, 8) may also be involved in rod formation. 
This layer was observed by Healy (1988b) in sea water- 
formalin/ethanol fixed sperm of P. westralis and inter- 
preted as the possible remnants of nuclear membranes. 
Our observations, based on glutaraldehyde-fixed sperm 
ot P. quoyanus, show that such material truly lies outside 
the intact nuclear and acrosomal membranes, and there- 
fore constitutes part of the subacrosomal material. 

The close resemblance of the crypt-like nuclear fossa 
of Perotrochus spp. (HeaK, 198Sb; this study) to the 
spermatid fossa of Calliotropis glyptus (Healy, 1989) has 
already been mentioned. In most vetigastropods and the 
Patellogastropoda, the centrioles are only superficially 
attached to a shallow nuclear invagination. In Haliotis, 
the proximal centriole itself sometimes occupies the shal- 
low fossa (Lewis et al, 1980; Sakai et al.. 1982), while 
in Sinezona (Scissurellidae) and Calliotropis (Trochidae) 
the centriole(s) and proximal portion of the a.xoneme are 



Figures 36-42. Perotrochus quoyanus '.ib.'.M. IJcvt'lopiiig iiucleu.s (n), iiiidpiece niitoclidndria (m). axoneme (ax), centrioiar 
fossa (arrowhead), and proxiiiiu! (pel and (iLslai (("iitrioies ulc) of advanced spermatids. 36 Inset. Triplet microtubules of centriole 
in advanced spermatid 38. Morihund spermatoc\te sliowing s\ iiaptinemal complex (arrows), mitocliondria (m). proacrosomal 
vesicles (pav), and axoneme profiles (arrowheads). 3'>. .\herrant spermatit! Note angular shape of cell and its condensing nucleus 
(n), as well as the presence ot proacrosomal vesicles and milocliondria (m) M). Mature' aberrant spermatozoon wedged between 
early, prohabb normal spermatids. Note mitociiondria (ni) in depressions al base of nucleus (n) Inset. Detail of unfused proacrosomal 
vesicles (pav) from aberrant spermatozoon. 41. Fully "condensed' nucleus (n) of aberrant spcini showing irregular shape and 
multiple axonemal profiles (ax). 42. Nuclear lacunae (nl) of aberrant spermatozoon 
Scale bars: 36,37,40 Inset,4l,42 = 0.25 tim. 38-40 = 1 m"i 



J. M. Healy and M. G. Harasewych, 1992 



Page 11 




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THE NAUTILUS, Vol 106, No. 1 



actually contained within the fossa (Healy, 1989, 1990a). 
The ball-and-socket fitting of rootlet and centriolar fossa 
of Perot rochtis spp. is unusual among gastropods, al- 
though a similar configuration occurs in the shipworm 
bivalve Lyrodus bipartita (Jeffreys, 1860) (see Figure 4 
of Popham, 1974). Examination of other genera {Mi- 
kadotrochtts. Entemnotrochus) may show this type of 
nuclear fossa to be a feature of all Pleurotomariidae. 

Nuclear lacunae are widely reported in spermatozoa 
of externally fertilizing mollusks, polychaetes, brachio- 
pods, echinoderms, as well as of some internally fertil- 
izing groups (e.g. some teleosts, Homo) (Baccetti & Afzel- 
ius, 1976). Their occurrence or degree of development 
seems to be more closely linked with the mode of nuclear 
condensation than with the degree of modification of 
nuclear shape occurring during spermiogenesis. For ex- 
ample, in the trochoid Zaiipais lascroni. the euspermatid 
nucleus undergoes marked elongation during conden- 
sation (fibro-granular pattern), but retains lacunae that 
ultimately fuse to form an axial tube within the mature, 
filiform nucleus (Healy, 1990b). Lacunae are usually not 
observed where nuclear condensation proceeds through 
either or both longitudinal fibrillar and lamellar phases 
(see Kaye, 1969; Horstman, 1970, Maxwell, 1983; Koh- 
nert & Storch, 1984b; Koike, 1985). 

The midpiece and satellite fiber/centriole complex of 
Perotrochus spp. are essentially as observed in the ma- 
jority of Vetigastropoda and Patelloidea (Koike, 1985; 
Hodgson & Bernard, 1988; Healy, 1990a; Healy & Dad- 
dow unpublished), the Bivalvia (for references see Po- 
pham, 1979), Scaphopoda (Dufresne-Dube et al, 1983) 
and Caudofoveata (Buckland-Nicks & Chia, 1989). The 
same arrangement of these organelles, clearly one asso- 
ciated with sperm tail attachment and stability, also oc- 
curs in spermatozoa of many other externally fertilizing 
animal species (for major references see Baccetti & Afzel- 
ius, 1976; Wirth, 1984). 

The flagellum consists of an axoneme (9 + 2 microtu- 
bular substructure) sheathed by the plasma membrane. 
Our scanning electron micrographs reveal that the fla- 
gellum is narrower in diameter close to the nucleus. TEM 
observations suggest that this is probably the result of a 
more closely applied plasma membrane in this region of 
the flagellum, although slight narrowing of the axoneme 
does occur near the distal centriole (see figs. 4, 15). At 
present we caruiot clarify the origin of the dense body 
.sometimes observed within the flagellum (see fig. 17). It 
was not observed in longitudinal sections through the 
immediate post-nuclear region of the flagellum and could 
yet prove to be an artifact of fixation. 

Systematic Considerations: If spermatozoa of Perotro- 
chus spp. are representative of the Pleurotomariidae, 
then a closer relationship between this family and the 
Trochoidea (particularly Trochidae) than with the other 
pleurotomarioidean {s.l.) families Haliotidae and Scis- 
surellidae seems evident. This conclusion accords both 
with Ilas/.prunar's (1988, 1989) finding that no synapo- 
morphies exist to unite the Pleurotomarioidea (s.l.), and 
with his decision to place the Haliotidae and Scissurel- 



lidae into separate superfamilies within the Vetigastro- 
poda. The question as to whether ancestral vetigastropods 
were more like scissurellids than pleurotomariids (see 
Haszprunar, 1988, 1989 for discussion) cannot yet be 
resolved using sperm data alone because too many sig- 
nificant taxa (including the new hydrothermal vent 
groups) remain unstudied. Based on the present evi- 
dence, however, we suspect that spermatozoa of any stem 
vetigastropod would have resembled more closely the 
unmodified type of Perotrochus (Healy, 1988b; this pa- 
per) than the modified type of Sinezona (Healy, 1990a). 
Vetigastropoda, Patellogastropoda and Neritimorpha 
can be distinguished on the basis of sperm features (es- 
pecially acrosomal and nuclear) and features of sper- 
miogenesis (dimorphic in the case of the Neritimorpha; 
rarely so in the Vetigastropoda) (Koike, 1985; Healy, 
1988a, 1990a, b). It will be interesting to determine 
whether the cocculinids — once included in the Vetigas- 
tropoda (Salvini-Plawen, 1980) but since removed to a 
separate archaeogastropod suborder, Cocculiniformia 
(Salvini-Plawen & Haszprunar, 1987) — also show char- 
acteristic sperm and spermiogenic features. 

ACKNOWLEDGMENTS 

We are grateful to the crews of the Johnson-Sea- Link 
II submersible and the R/V Seward Johnson for their 
assistance in collecting and maintaining the specimens 
upon which this study is based. Thanks are are extended 
to Professor G. Grigg of the Department of Zoology, 
University of Queensland for providing access to TEM 
facilities and to Mrs. L. Daddow and Mr. T. Gorringe 
(also Department of Zoology) for assistance with TEM 
and photography. Financial support for the work has 
been provided by a Queensland Museum Postdoctoral 
Research Fellowship (to J.M.H.). This study represents 
contribution number 284 of the Smithsonian Marine Sta- 
tion at Link Port, and contribution number 881 of the 
Harbor Branch Oceanographic Institution. 

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THE NAUTILUS 106(l):15-20, 1992 



Page 15 



Tenagodus or Siliqiiaria? Unraveling Taxonomic 
Confusion in Marine "Worm-Snails" 
(Cerithioidea: Siliquariidae) 



Riidiger Bieler 

Department of Zoology 
Field Museum of Natural History 
Roosevelt Road at Lake Shore Drive 
Chicago, IL 60605, L'SA 



ABSTRACT 

The nomenclatural history and availability of some genus- and 
family-group names in the marine "worm-snair' family Sili- 
quariidae are discussed: (1) Tenagodus Guettard, 1770 (type 
species; Serpula anguina Linne, 17.5iS) has priority over its 
objective junior synon\m Siliqiiaria Bruguiere, 1789. Tena- 
godes. Tenagoda and Silicaria are unjustified emendations 
Siliqiiaria Schumacher, 1817 (also subsequently emended to 
Silicaria) is a junior homonym described in the Bivalvia. (2) 
Siliquarius Montfort. 1810 (type species: Siliqitarim angiiiliis 
Montfort, 1810) is an available name. (3) Angiiinaria Schu- 
macher, 1817, is preoccupied by Angninaria Lamarck, 1816 
(Bryozoa). (4) Montfortia Delia Campana, 1890 (non Mont- 
forlia Recluz, 1843), and its replacement name Hcmitcnago- 
diis Rovereto, 1899, are based on the type species Tenagodus 
bernardii Morch, 1860. (.5) Agaihirscs Montfort, 1808, is based, 
b\ original designation, on (he t\pe species .A^af/ilr,sc.s/iir('<'//i(.s 
Montfort, 1808 (a senior synon\ni of Siliqiiaria spinosa La- 
marck, 1818). Agathyrsus Herrmannsen, 1846, is an unjustified 
emendation; "Agathinus," "Agathirsis" and 'Agathirsus ' are 
incorrect subsequent spellings. (6) The family name Siliquarii- 
dae Anton, 1838, has priority over its objective junior synonym 
Tenagodidae Gill, 1871 

Key uords: Ta\<iniini\ , Nomenclature. Synonym) . (laslrnpoda 



INTRODUCTION 

Marine "worm-snails ' are now understood to be mem- 
bers of three different families, the Vermetidae, genus 
Vertincularia in the Turritellidae, and Siliquariidae (e.g., 
Morton, 1951, 1953; Bieler, 1990) The siliqiiariids, often 
(but not always) recognizable by longitudinal shell slits 
or series of holes, are an enigmatic group. Little is known 
about their biology and geographical distribution, and 
the published literature provides seldom more than de- 
scriptions of empty shells or shell fragments. Consider- 
able confusion exists about the synonymy and relation- 
ships of nominal taxa such as Tenagodus and Siliqiiaria, 
or Tenagodidae and Siliquariidae. In the aftermath of 
more than 200 years of misspelling, emending and re- 



placing certain names, today s worker is left to wonder 
about how many "real ' taxa are hidden behind Tena- 
godtis-Tenagodes-Tenagoda, Siliqiiaria -Silicaria -Sili- 
quarius, or Agathirses-Agathijrsiis-Agathinus. This is 
complicated further by family names that were later 
based on some of these names. 

This paper will not discuss phylogenetic relationships, 
a topic that demands detailed anatomical studies that 
are in progress (Bieler, in preparation). No interpretation 
of subjective synonymy between nominal genera in the 
Siliquariidae will be made before that study is complete. 
However, taxonomic problems often can be solved by a 
careful study of existing literature and application of 
ICZN rules. This contribution is meant as a taxonomic 
"house cleaning, ' addressing problems of type species, 
priority and availability of various generic and family 
names for this group. A main goal is to identify objective 
synonymies (and consequently reduce the number of 
nominal taxa in future publications). 



TAXONOMY 

Genus-group names 

Tenagodus Guettard, 1770 

Type species: Serpula anguina Linne, 1758. 

The name Tenagodus was first made available by 
Guettard (1770:128). Several later authors {e.g., Herr- 
mannsen. 1849:540; Delia Campana, 1890:139; Coss- 
mann, 1912:146) refer to an earlier work by Guettard 
(usually cited as "1760") for the introduction of Tena- 
godus or "Tenagoda." However, that publication, "An- 
nee 1760" of the Histoire de T Academie Royale des 
Sciences (1766a, b), contains only vernacular names in 
text and figure captions. Guettard (1770) did not indicate 
a type species. The first subsequent designation of a type 
species appears to be that of Adams and Adams (1854: 
360-361), who selected "T[enagoda]. anguina, Linnae- 
us" [Serpula anguina Linne, 175S]. 



Page 16 



THE NAUTILUS, Vol. 106, No. 1 



Two different eiiu'iidatioiis of Tenagodus (unjustified 
in the sense of ICZN Art. 33) were introduced: 

Tenagodes P. Fischer, 1885 (p. 692); Sacco, 1896:17; Cossmann, 
1912:146. 

Tenagoda Agassiz, 1848 (p. 1049); Herrmannsen, 1849:540; 
Adams & Adams, 1854:360, corrected to Tenagodus in 
1858:656; Chenu, 1859:321; Paetel, 1869:9. 

Siliquaria was first introduced by Bruguiere (1789:xv). 
The type species is Serpula angiiina Linne, 1758, by 
subsecjuent monotypy (Lamarck, 1799:79). Bruguiere's 
Siliquaria was later erroneously credited to Lamarck (e.g.. 
Schumacher, 1817:262; Rang, 1829:187). Silicaria, an 
emendation of Siliquaria Bruguiere or "Lamarck' (un- 
justified in the sense of ICZN Art. 33), was introduced 
and used by some authors (e.g., Bosc, 1802:157; Daudin, 
1800:32; Audouin, 1829b). 

Siliquaria was also described as a genus of bivalves, 
by Schumacher (1817:43, 129). Schumacher knew of the 
conflict with the earlier name, and conserved his nominal 
taxon by dismissing the senior homonym. For the latter 
he introduced the new worm-snail genus Anguinaria (see 
below): "Je n'ai pu conserver le nom de ce genre donne 
par Mr. Delamarck savoir: Silicaire (Silicaria), comme il 
y a un genre parmi les bivalves qui porte un nom presque 
semblable " (1817:262). Siliquaria Schumacher was also 
subsequently emended to Silicaria (e.g., Paetel, 1875: 
191). 



Discussion of the type species 

Linne's (1758:787) Serpula anguina was originally 
based on more than one species, and the Linnean col- 
lection contains specimens of several forms, with no clear 
type specimen indicated (Hanley, 1855:448). Linne had 
adopted the species name from the non-binominal "Solen 
anguinus' of Rumphius (1705:125) and referred to fig- 
ures in that work (Rumphius, 1705: pi. 41, fig. H) and 
in Argenville (1742: pi. 29, fig. H). Rumphius' figure 
shows a spiny specimen of Tenagodus, Argenville's il- 
lustration is that of a member of the Vermetidae and 
was later used by Linne (1767:1266) as reference to Ser- 
pula arenaria Linne, 1758. The type locality for S. an- 
guina was given as "India " (Linne, 1758:788). Linne also 
listed several references to a variety "/3 " {e.g., Rumphius, 
1705: pi. 41, fig. 2; Gualtieri, 1742: pi. 10, fig. Z). Most 
of these latter illustrations show a spineless Mediterra- 
nean form of Tenagodus later described as Anguinaria 
ohtusa Schumacher, 1817 (see below). 

Born (1780:440), possibly misled by Linne's statement 
in the Museum Ulricae (1764:701) that the variety had 
a spiny shell, then confused the issue by applying the 
name Serpula anguina to the smooth form and intro- 
ducing the name Serpula rnuricata for the true anguina. 
Lamarck (1801:98) based his interpretation of S. anguina 
on yet another illustration of a smooth form, by Davila 
(1767: pi. 4, fig. E), later described ds Siliquarius anguilus 
Montfort, 1810 (see below). Manv subsequent autliors 
(e.g., Lamarck, 1818:337; Philippi, 1836:173; Chenu, 



1843:1; Cossmann, 1912:146) followed and sometimes 
defended (Weinkauff, 1868:330) Born's misconception. 
Although the problem was discussed at several occasions 
[Hanley, 1855:448; M6rch, 1860b:403; Tryon, 1886:189; 
Sowerby, 1876:(1)], the nsune anguina remained mistak- 
enly in use also for the smooth Mediterranean form, 
especially in paleontological works (e.g., Strausz, 1966: 
122). 

When Lamarck (1799:78) and Adams & Adams (1854: 
361) employed the name anguina in fixing the type 
species of Siliquaria and Tenagodus, respectively, it was 
in this misidentified fashion. Lamarck ( 1799:78-79) men- 
tioned a shell without spines for Siliquaria, and his later 
works (e.g., 1801, 1818) clearly identify the Mediterra- 
nean form. Adams and Adams (1854:361) mentioned that 
the "typical species is found in the Mediterranean ' and 
their illustrations in the same work (pi. 39, fig. 5) show 
smooth Mediterranean specimens. 

To best serve stability in nomenclature, and in agree- 
ment with current usage (e.g.. Wenz, 1939:679; Mala- 
testa, 1974:200; Squires, 1990:286), the nominal species 
named in fixation are here accepted as type species, 
regardless of misidentification. A referral to the Inter- 
national Commission on Zoological Nomenclature, as 
specified in ICZN Article 70(b), seems unnecessary in 
this case. 

The synonym} of Tenagodus and Siliqiuiria has long 
been known (e.g.. Chenu, 1843: 1; Herrmannsen, 1849: 
540; Paetel, 1869:xiii, 56; Keferstein, 1862-66:1056), and 
the priority of Tenagodus was stressed by some authors 
(e.g.. Strausz, 1966:122). Other workers, however, re- 
jected Tenagodus as non-binominal (e.g.. Daii, 1889:259; 
Gould, 1966:3; Abbott, 1974:96), and both names have 
been used in parallel for the past 200 years. 

The genus-group name Tenagodus is available and 
has priority over Siliquaria (a junior objective synonym), 
for reasons given below: 

(1) The name Tenagodus was used as a scientific name 
by the author when published [ICZN Art. 11(b)]. 
Guettard s description (1770:128) begins "Genre X. 
Tenagodus. Tenagode [the French vernacular]. Ca- 
ractere generique. ..." 

(2) The name Tenagodus was introduced in a work pub- 
lished before 1931 and containing uninominal genus- 
group names without associated nominal species. It 
can be accepted "as consistent with the Principle of 
Binominal Nomenclature in the absence of evidence 
to the contrary" [ICZN Art. ll(c)(i)]. 

(3) The original work contains a description, illustra- 
tions, and bibliographic references to additional il- 
lustrations, thus fulK satisf\ing ICZN Art. 12 (a new 
scientific name pui)lished before 1931 must have been 
accompanied by a description, definition or indica- 
tion). 

(4) The work has not been placed on the Official Index 
of Rejected and Invalid Works [ICZN Art. 78(h)]. 

(5) Other genus-group names introtluced in the same 
work in similar fashion are in current use, such as 



R. Bieler, 1992 



Page 17 



Kuphus, Uperotiis and Brechitcs, which were ac- 
cepted by later workers (e.g.. Turner, 1966; Smith, 
1971) for same reasons. A suppression of Guettard s 
work by plenary power of ICZN would create new 
taxonomic problems. 

(6) The name Tcnagodus (in various spellings) has been 
used frequenti\ {e.g., Morcii, lS6()a, 1); Fischer, 1885; 
Sacco, 1896; Martin, 1899; Simroth, 1906; Clossmann, 
1912; Thiele, 1929; Wenz, 1939; Strausz, 1966; Mala- 
testa. 1974; Boss, 1982). It does not qualify as an 
"unused senior synonym" [ICZN Art. 79(c)]. 

(7) Siliqiiaria Bruguiere, 1789, and Tcnagodus Guet- 
tard, 1770, have (by subsequent designations) the 
same type species; they are objective synonyms. 

Siliquarius Monlfort, 1810 

Type species: Siliquarius anguilus Montfort, 1810. 

Siliquarius was introduced by Montfort (1810:39). The 
type species, by original designation, was given as Sili- 
quarius anguilus (Montfort, 1810). The description was 
accompanied by a drawing of a siliquariid shell without 
spines, and Montfort made reference to a figure by Davi- 
la (1767: pi. 4, fig. E) and to Siliquaria anguina sensu 
Lamarck, 1801 (who had based his concept of S. anguina 
on Davila's figure). 

Siliquarius was used by some later authors [e.g., Morch, 
1860b) because Siliquaria was considered "preoccupied 
for a genus of plants by Forskal (1775). The name Sili- 
quarius Montfort is available for nomenclatural purposes. 

Anguinaria Schumacher, 1817 

[preoccupied] 

Schumacher (1817:262) introduced Anguinaria and in- 
cluded two new species: Anguinaria obtusa. referring to 
"Serpula anguinaria [err. pro anguina]" sensu Born 
(1780:440, pi. 18, fig. 15), and Anguinaria rubra, referring 
to Serpula anguina, sensu Martini (1769:50, pi. 2, figs. 
13-14). No subsequent type species designation was lo- 
cated. 

The name Anguinaria Schumacher, 1817, is not avail- 
able for nomenclatural purposes, because it is preoccu- 
pied by Anguinaria Lamarck, 1816 (p. 142; Bryozoa). 

Hemitenagodus Rovereto, 1899 

Type species: Tenagodus bernardii Morch, 1860. 

Delia Campana (1890:139-140) introduced the genus- 
group name Montfortia as a subgenus of Tenagodus (p. 
139) or new genus (p. 140). The name Montfortia was 
based, by original designation, on Tenagodus (Siliquari- 
us) bernardii Mcirch, 1860 (1860a:368, a sponge-dwelling 
Recent form from unknown type locality). Later workers 
(e.g., Wenz, 1939:680) erroneously considered the fossil 
species Montfortia ligustica Delia Campana, 1890, as 
type species by monotypy. 

Rovereto (1899:108) recognized Montfortia Delia 
Campana, 1890, as preoccupied by Montfortia Recluz, 
1843, and introduced the replacement name Hemitena- 



godus. Thus, Hemitenagodus is also based on Tenagodus 
bernardii Morch. 



Agathirses Montfort, 1808 

Type species: Agathirses furcellus Montfort, 1808. 

Montfort (1808:399) introduced Agathirses with A. 
furcellus as type species by original designation ( 'Espece 
servant de type au genre") for an Eocene fossil. The 
species name was borrowed from "furcelle (furcella), ' 
listed as a "singulier tube testacee multioculaire " of un- 
certain systematic placement by Lamarck (1801:104). 
Montfort referred to tiie 'Siliquaire de Grignon " as il- 
lustrated by Faujas-St.-Fond (1803: pi. 3). Lamarck (1818: 
338) eventually also named the Grignon fossil, as Sili- 
quaria spinosa. Sherborn (1930:6087) erred when he 
credited the name Siliquaria spinosa to G. Fischer (1807: 
244), who had only used the French vernacular "Sili- 
quaire epineuse." Later authors erroneously cited the 
junior synonym, Siliquaria spinosa Lamarck, as type spe- 
cies of Agathirses (e.g., Herrmannsen, 1846:25; Coss- 
mann, 1912:148; Wenz, 1939:680). 

At least one intentional emendation of Agathirses (un- 
justified in the sense of ICZN Art. 33) was introduced: 

Agothijrsiis Herrmannsen, 1846 (p. 25), Agassiz (1848:29); Coss- 
mann, 1912:148, 

"Agathinus Mft." Paetel (1888:499), "Agathirsis Mft." 
Paetel (1875:5) and "Agathirsus Cossmann, 1912" Wenz 
(1939:680) are here considered incorrect subsequent 
spellings without nomenclatural bearing [ICZN Art. 33(c)]. 

Family-group names 

Siliquariidae 

[based on Siliquaria Bruguiere, 1789] 

Siliquariidae was first introduced by Anton (1838:xiii), 
as "Siliquariacea." The name was frequently credited to 
later authors, such as Chenu, 1859; it was also indepen- 
dently proposed by several later workers (e.g., Morton, 
1951:40) unaware of its earlier introduction. This is Sil- 
iquariadae Schaufuss in Paetel (1869:9) and "Siliquari- 
dae" auct. (e.g., Paetel, 1869:9). 

Tenagodidae 

[based on Tenagodus Guettard, 1770] 

Tenagodidae was first proposed by Gill (1871:8). It was 
also independently introduced by later authors who over- 
looked the earlier introduction (e.g., Malatesta, 1974: 
200). 

Although the genus-group name Tenagodus has pri- 
ority over Siliquaria, the family-group name Siliquari- 
idae has priority over Tenagodidae: 

(1) Both generic names have been the basis of family 
names. 

(2) The family-group names Siliquariidae and Tenago- 
didae are objective synonyms, because their name- 



Page 18 



THE NAUTILUS, Vol. 106, No. 1 



bearing genera are objective synonyms [ICZN Art. 
61(c)(ii)]. 
(3) As the senior s\ iionym, Siliquariidae can continue in 
use altliough Siliqiiaria is a junior synonym, because 
the Principle of Priority also applies to family-group 
names concerned [ICZN Art. 40(a)(i)]. 



ACKNOWLEDGMENTS 

This paper required access to some rare and obscure 
works and would not have been possible without the 
excellent library facilities at the Academy of Natural 
Sciences of Philadelphia (ANSP), the Delaware Museum 
of Natural History (Wilmington), the Field Museum of 
Natural History (Chicago), the Museum of Comparative 
Zoology (Harvard University, Cambridge [MCZ]), and 
the National Museum of Natural History (Smithsonian 
Institution, Washington, D.C. [USNM]). Special thanks 
are due to Dr. Kenneth J. Boss, Mrs. Marion Britz, Dr. 
Silvard Kool (MCZ), Dr. R. Tucker Abbott (Melbourne, 
Florida), Dr. Robert Robertson (ANSP), Dr. M. G. Har- 
asewych (USNM), and Dr. Alan Kabat (formerly MCZ), 
who helped with the literature search. Dr. James F. Quinn 
(Florida Marine Research Institute, St. Petersburg) ex- 
pertly commented on an earlier draft of the Tenagodus- 
Siliquaria discussion, and Mr. Richard E. Petit (North 
Myrtle Beach) kindly reviewed the manuscript. Two 
anonymous reviewers provided valuable critique and 
suggestions. 



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THE NAUTILUS 106(1 ):21-23, 1992 



Page 21 



Nucella Roding, 1798 (Gastropoda: Muricidae): Type Species 



Silvard P. Kool 
Kenneth J. Boss 

Moliusk Ueparlmt-iit 
Museum of Comparative Zoology- 
Harvard I'liiversit) , Cambridge, MA, 
0213.S 



ABSTRACT 

Bticciniim filosum Gmelin, 1791 [junior synonym: Nucella 
tlu'obronui Roding, 1798; senior synonym Biiccinurn lapillus 
Linnaeus, 1758; now known as Succlla lapillus (Linnaeus)], is 
the type species of Succlla Roding 1798 In subsequent des- 
ignation of Stewart (1927:386, footnote #260), 

Key words: Nucella lapillus. type species; taxonomy; Murici- 
dae, 



INTRODUCTION 

Roding (1798:130-131) introduced the genus Nucella 
and included five names: iV, reticulata, N . nioschatellina 
with Buccinum laeve Gmelin, 1791 listed as a synonym; 
N. macina; N. lapillus with Buccinum rusticurn Gmelin, 
1791 as a synonym; and N. theohroma with Buccinum 
filosum Gmelin, 1791 as a synonym. Roding did not 
specify a ty pe species. Because the type species was not 
fixed in the original publication, only the subsequent 
designation of a type species from the nomina enumer- 
ated above is valid in accordance with Article 69 (Code 
of the International Commission on Zoological Nomen- 
clature [hereafter I.C.Z.N.] (1985)). Several authors have 
either incorrectly designated a type species for Nucella 
or discussed efforts to designate a type. This confusion 
was partly caused by the fact that the binomen Nucella 
lapillus R(")ding is a secondary homonym, but not a syn- 
onym, of Buccinum lapillus Linnaeus, 1758. 

TAXONOMIC HISTORY: A COMEDY OF ERRORS 

Dall (1906; 1909), Clench (1947), and, in more detail, 
Rehder (1962) discussed the species introduced by Rod- 
ing (1798) in Nucella. The first species, N. reticulata, is 
the cancellariid Cancellaria reticulata (Linnaeus, 1767); 
the second, N. moschatellina is a nomen dubium; the 
third, N. macina is a nomen nudum, the fourth, N. 
lapillus, not to be confused with and not the same as 
Buccinum lapillus Linnaeus, 1758, had in its synonymy 
Buccinum rusticum Gmelin, 1791 and is identified as 
Latirolagena smaragdula (Linnaeus, 1758); the fifth spe- 
cies, Nucella theobroma, with Buccinum filosum Gme- 



lin, 1791 listed as a synonym, is a subjective junior syn- 
onym of Buccinum lapillus Linnaeus, 1758, now referred 
to as Nucella lapillus (Linnaeus, 1758) (Kool, 1989). 

Dall (1906), after prompting acceptance of Roding's 
names, acknowledged that tletermining the correct type 
species for Nucella would "recjuire special study to un- 
ravel." In a later work, Dall (1909) provided not one, 
but three different binomina for the type species des- 
ignation, without mentioning the word "type. " 

First, in his list of subgenera within the genus Thais. 
he (1909:46-47) cited "Purpura filosa Menke = P. la- 
pillus L." after Nucella. Wenz (1941:1123) mistakenly 
interpreted this as a species designation and credited Dall 
(1909) with the designation of "N. filosa (Gmelin) [Buc- 
cinum] = lapillus (Linne) [Buccinun^]." Whatever the 
inappropriateness of this invalid designation, Dall left no 
doubt that he recognized these specific nomina as syn- 
onyms by placing "P. lapillus L." in synonymy with 
Purpura filosa "Menke " [= Buccinum filosum Gmelin]. 
It should be mentioned that Dall s attribution of filosa 
to Menke (1830:62) was an erroneous concession to the 
custom of the nineteenth century where some authors 
credited themselves for a specific name after having al- 
tered the generic placement and, thus, the binominal 
combination, in this case, from Buccinum filosum to 
Purpura filosa. 

Secondly, in the same work, Dall (1909:48) stated ". . . 
thus fixing the name Nucella on tlie well-known Purpura 

lapillus by which action he obviously meant to 

restrict, by subsequent designation, the concept of Nu- 
cella, but he again did not mention the necessary word 
"ty pe " 

Thirdly , in another listing of subgeneric and sectional 
taxa, Dall (1909:50) designated many type species, and 
he placed "T. lapillus (Linn) ' right after Nucella s.s., 
probably meaning this to be a type designation. But 
although Nucella is on that list, he again failed to use 
the word "ty pe. Therefore, v\ hatever his intentions, Dall 
(1909) provided three different, ambiguous, indefinite 
and invalid subsequent designations. 

Later works that treated Nucella include Suter (1913: 
425), who listed "Purpura lapillus L. ' as type species of 
Nucella, an invalid designation, because lapillus Lin- 



Page 22 



THE NAUTILUS, Vol. 106, No. 1 



naeus was not included among the species of Nucella in 
Rfiding (1798), and i.s therefore not available (I. CI. Z.N. , 
Article (59). 

Iredale (1915:472), without realizing that Dall had not 
designated a valid type species for Nucella, reprinted 
Dall's (1909:50) list and added the word "Type," but 
misquoted Dall by listing "T lapillus Lam." instead of 
"T. lapillus (Linne)." 

Dall (1915:557) discussed the type species designation 
oi Nucella and stated: "The type of the subgenus is the 
solitary North Atlantic species of N. lapillus of Linnaeus, 
..." However, although Dall mentioned the word 
"type, lapillus Linnaeus is not available because it was 
not a name included under Nucella by Roding (1798). 

Stewart (1927:386) wrote in a footnote (#260): "[Nu- 
cella:] Type species Buccinunt filusuni Gmelin = B. la- 
pillus h." Stewart was the first author to designate a valid 
type species for Nucella, without realizing this himself; 
he continued: "It is probably safe to say that Dall fixed 
the t\pe in 1909. The species Buccinuni filosum, orig- 
inally used by Roding [as a synonym ot Nucella theo- 
broma Roding] in Nucella, is nomenclatorially available 
for the type designation. Stewart also recognized that B. 
filosum was synonymous with B. lapillus Linnaeus. 

Treating Nucella as a separate genus with four sec- 
tions, Thiele (1929:298) mentioned Nucella (Nucella) 
lapillus (Linne) as an example; he did not, however, use 
the word "type," and none of the available nomina was 
cited. 

Grant and (;ale (1931:716), following Stewart (1927), 
also credited Dall (1909) as the subsequent designator 
and gave the type as "Nucella lapillus Linnaeus." How- 
ever, aside from not being available, this binomen was 
not one of the three combinations cited by Dall as possible 
types. 

In his list of the type species of marine mollusca of 
the British Isles, Winckworth (1932:229) listed N. theo- 
lyronia Roding, 1798 as the "genotype" of Nucella, how- 
ever, Stewart s (1927) type designation preceded this one. 

Winckworth (1945:141), as he had done in his 1932 
paper, again cited N. theobroma Roding as type species 
of Nucella and correctly noted that it was a junior syn- 
onym of Buccinuiu lapillus Linnaeus. However, he er- 
roneously attributed the designation to Dall (1909), who 
never used the name theubroma. 

Clench (1947:86) in an ambiguous treatment of lapil- 
lus Linnaeus, placed the species in the geiuis Thais (Riid- 
ing, 1798) and used the subgeneric name Polytropa 
Swainson, 1840 [type species Buccinuni lapillus Linnaeus 
by subsequent designation, j.E. Gray, 1847], rather than 
Nucella. (bench's argument for not using Nucella was 
that Roding never intended to include Buccinuni lapillus 
Linnaeus in his genus Nucella. Glench arrived at this 
conclusion after comparing the hgures in Martini (1777: 
pi. 121, figs. 1 1 1 1-11 12) tor lapillus Limiaeus and those 
ioT filosum Gmelin (pi. 121, figs. 1 113-1 1 14). According 
to Clench, the figures for Riiding's Nucella theubroma 
and its synonym Buccinum filosum Gmelin, are unrec- 
ognizable, whereas the figures of i^(")ding's Nassa rudis 



and its associated synonym, "Buccinum lapillus Gme- 
lin," are those of Buccinum lapillus Linnaeus The latter 
portion ot this argument is true, but the former, as Reh- 
der (1962) pointed out, is not tenable: the "unrecogniz- 
able" figures oi Nucella theobroma in Martini (1777: pi. 
121, figs. 1113-1114) also clearly represent Nucella la- 
pillus Linnaeus, in one of its many variations. Rehder 
(1962, figs. 1-2) illustrated a specimen oi lapillus from 
France that closely matched Martinis (1777: pi. 121, figs. 
1113-11 14) illustrations. It has long been known that the 
species displays a high degree of variation in shell mor- 
phologv and color (Martini, 1780: pi. 122, figs. 1124- 
1 125, 1128-1 129; Forbes & Hanley, 1851; Crothers, 1985). 
In his discussion of Nucella, Clench did not mention the 
attempted designations of Dall (1915), Suter (1913), 
\Ninckworth (1932), and the valid designation of Stewart 
(1927). 

Clench's usage of Polytropa instead of Nucella is not 
followed herein. Nucella is a valid taxon to be used for 
Linnaeus lapillus. Interestingly enough. Clench cited 
"Purpura filosa Menke" and "Buccinun} filosum Gme- 
lin" in his synonymy of "Thais {Polytropa) lapillus 
Linne," which contradicts his own argument that Roding 
never intended to include Buccinum lapillus Linnaeus 
in his genus Nucella. 

Finally, in an attempt to resolve some of the irregu- 
larities and inconsistencies discussed so far, Rehder (1962: 
110) stated that Dall (1909:50) intended to designate a 
type and that another possible attempt at a designation 
(Dall, 1909:46) is invalid because Dall cited fi/o.sa Menke, 
no\. filosa Gmelin. Further, Rehder attributed the earliest 
type designation to Winckworth (1932:229) who listed 
Nucella theobroma and pointed out that Clench (1947) 
had overlooked this type designation. 

SUMMARY 

In conclusion, the correct tv pe species for Nucella Rod- 
ing, 1798 is Buccinum filosum Gmelin, 1791, a subjective 
junior synonym of Buccinum lapillus Linnaeus, 1758, 
and an objective senior synonym of Nucella theobroma 
Roding, 1798, by subsequent designation of Stewart (1927: 
386, footnote #260). 

ACKNOWLEDGMENTS 

The authors thank Mr. Richartl I Johnson for a critical 
re\ iew of an earlier draft of this paper, and are indebted 
to two anoiiN nious reviewers. 

LITERATURE CITED 

clench, W.J 1947. Tlic geiu'ra Purpura and Thais in the 
western Atlantic, Joliii.soiiia 2(23):61-91, pis. 32-40. 

(Mothers, J.H. 1985 Dog-whelks: an iiitrodiictioii to the bi- 
ology of Nucella lapillus. Field Studies 6:291-360 

D.ill, VV.n. 1906. Early history of the generic name Fiisus. 
Journal of Coiichology 1 1( l'o):289-297. 

Dall. W.H. 1909. Contrihtitions to the Tertiar\ paleontology 
(il till- Pacilic coast. 1. The Miocene of .Astoria and Coos 



S. p. Kool and K. J. Boss, 1992 



Page 23 



Bay, Oregon United States Geological Survey Professional 
Paper 59, 278 pp., 23 pis, 14 figs. 

Dall, \V H. 1915. Notes on the species of the niollnscaii sub- 
genus Mticclhi inhabiting the northwest coast of America 
and adjacent regions. Proceedings of the L'niteii States 
National Museum 49:557-572, pis. 74-75. 

Forbes, E. and S. Hanle>. 1851. A histor\ of British Mollusca. 
and their shells. Volume 3. London: John van X'oorst, Pa- 
ternoster Row. [3] vi-\, [1] pp. 2-616. 

(wnelin, J F 1791 Caroli a Linne, Svstema Nat\irae per Reg- 
na Tria Naturae . Editio Decima Tertia, Aucta, Refor- 
mata. Tomus 1, Pars 6. Beer, Leipzig., pp. 3021-3910 [for 
a full citation and discussion of the tv\'0 printings of this 
work see Kabat and Petit (1988)]. 

Grant, I'S., IV and H.R. Gale. 1931. Catalogue of the marine 
Pliocene and Pleistocene Mollusca of Galifornia and ad- 
jacent regions. Memoirs of the San Diego Society of Nat- 
ural History 1:1-1036, 15 figs., 32 pis. 

Grav, J.E. 1847 A list of the genera of Recent Mollusca, their 
sv nonvma and tv pes Proceedings of the Zoological Societv 
of London 15:129-219 

International ("onnnission on Zoological Nomenclature. 1985 
International Code of Zoological Nomenclature Third 
Edition, i-xx -t- 338 pp. International Trust for Zoological 
Nomenclature/British Museum (Natural Historv), Lon- 
don; LIniversity of California Press, Berkclev anil Los .An- 
geles. 

Iredale, T 1915 Art XLVTI — A commentary on Suter s 
"Manual of the New Zealand Mollusca. Transactions and 
Proceedings of the New Zealand Institute for the year 
1914, 47:417-497. 

Kabat. A.R. and RE. Petit. 1988 The two printings of J F. 
('•mvUu's. Syslcma Naturae, 13th Edition (1788-96). The 
Nautilus 102(4) 164-166. 

Kool, S.P. 1989. Phylogenetic analysis of the subfamily Thai- 
dinae (Prosobranchia: Neogastropoda; Muricidae). Ph.D. 
Dissertation, The George Washington L'niversity, Wash- 
ington,!) C, i-xiii -I- 342 pp. 



Liimaeus, C. 1758. Caroli Linnaei . . Systema Naturae per 

Regna tria Naturae, secundum classes, ordines, genera, 

species, cum characteribus, differentiis, synonymis, locis 
Editio decima reformata Impensis Direct Tom 

l.Animalia. Salvii: Holmiae, 824 pp 
Martini, F H.W. 1777. Neues systematisches Conchylien- 

('abinet. Volume 3. Raspe, Niiremberg, i-vi + 434 pp , 

pis. Ixvi-cxxi. 
Martini, F.H.W. 1780. Neues systematisches Conchylieii- 

Cabinet. Volume 4, Raspe, Nuremberg, [24] -I- 344 pp., 

pis. cxxii-clix 
Menke, K.T. 1830. Synopsis methodica inolluscorum gener- 

um omnium et specierum earum Uslar, Pyrmonti, 

i-xvi -I- 169 pp. 
Rehder, H. A. 1962. The status of Nucclla Roeding. The Nau- 
tilus 75(3): 109-1 1 1. 
Roding, P.F. 1798. Museum Boltenianum . : pars secunda 

continens Conchylia . Hamburg (Trapii), i-viii -I- 199 

pp 
Stewart, R.B. 1927. Gabb's (California fossil type gastropods 

Proceedings of the Academy of Natural Sciences of Phil- 
adelphia. 78:287-447. pis. .XX-XXXII. (January 3. 1927 

teste [Table of] Contents). 
Suter. H. 1913. Manual of the New Zealand Mollusca With 

an atlas of Quarto Plates. Mackay, Government Printer, 

Wellington, N. Z., i-xxiii 4-1120 pp. 
Swainson, W. 1840. A treatise on malacologv, or shells and 

shell-fish. London. 419 pp. 
Thiele, J 1929. Handbuch der Systematischen Weichtier- 

kunde. Volume 1. F'ischer, Jena, 376 pp. 
Wenz. W, 1941. Prosoliranchia. [in] OH. Sehindewolf (ed). 

Handbuch der Paliiozoologie Volume 6, part 5 Gebriider 

Borntr;iger. Berlin, pp 96M-1200. 
Winckvvorth. R. 1932. The British marine Mollusca. Journal 

of Conchology 19(7):21 1-252. 
Winckvvorth, R. 1945. The types of the Boltenian genera. 

Proceedings of the Malacological Societv of London 26(4/ 

5): 136-148. 



THE NAITILUS 106(1 ):24-.38, 1992 



Page 24 



A Revision of the Recent Species of Eudolium Dall, 1889 
(Gastropoda: Tonnoidea) 



Bruce A. Marshall 

National Museum of New Zealand 
P.O. Box 4(i7, Wellington, New 
Zealand 



ABSTRACT 

The tonnid genus Eudolium contains three Recent species: E. 
crosseanum Monterosato, 1869 (= thornpsoni McGinty, 1955), 
£. hairdii X'errill & Smith (= solidior Dautzenberg & Fischer, 
1906 = lineata Schepnian, 1909 = inflatum Kuroda & Habe, 
1952 = kuroharai .Azuma, 1960) and £. pyriformc Sowerbv, 
1914. Eudolium crosseanum and E. pyrijorme respectively 
have amphiatlantic-Mediterranean and Indo-Pacific distribu- 
tions, while £. hairdii occurs widely in the Atlantic, Pacific and 
Indian Oceans. Genus and species group synonymies and dis- 
tributions are discussed, and shells, radulae and male external 
anatomies are illustrated. 

Key words: Mollusca; prosobranch gastropods; Tonnidae; Eu- 
dolium. systematics; zoogeography 



INTRODUCTION 

The taxonomy of the species of the prosobranch family 
Tonnidae is poorly imderstood. Atlantic species were re- 
vised by Turner (1948), but there has been no critical 
modern revision of the Indo-Pacific species. The present 
contribution is the resuh of an attempt to determine the 
identity of two species of the genus Eudolium Dall, 1889 
obtained off New Zealand in recent years. 

Since Dail's (1889a) review, Eudolium bairdii (Verrill 
& Smith, 1881) has been treated as a synonym of E. 
crosseanum (Monterosato, 1869) by the majority of au- 
thors. From examination of numerous specimens, how- 
ever, it transpires that not only are they distinct species, 
but that five subsequently introduced taxa are synonyms. 
Moreover, £. crosseanum is more closely related to the 
Indo-Pacific £. pyriforrne (Sowerby, 1914) than to £. 
bairdii. 



ABBREVIATIONS AND TEXT CONVENTIONS 

AMS — Australian Mihseum, Sydney. 
MCZ — Museum of (Comparative Zoology, Harvard. 
MNHN — Museum National d'Hi.stoire Pslaturelle, Pari.s. 
NMNZ — National Museum of New Zealand, Wellington. 
NMP — Natal Museum, Pietermaritzburg. 
USNM — National Museum of Natural History, Wash- 
ington, DC. 



In captions to illustrations shell height dimension pre- 
cedes diameter. 



SYSTEMATICS 

Class Gastropoda 

Superfamily Tonnoidea Suter, 1913 
Family Tonnidae Suter, 1913 
Genus Eudolium Dall, 1889 

Doliopsis Monterosato, 1872(not Vogt, 1852, nor Conrad, 1865): 
8. Type species (by monotypy): Dolium crosseanum Mon- 
terosato, 1869; Recent, Mediterranean 

Eudolium Dall, 1889a:232. Substitute name for Doliopsis Mon- 
terosato (preoccupied). 

Galeodolium Sacco, 1891:4. Type species (by subsequent des- 
ignation of Voices, 1986:178): Cassidaria mutica Michelot- 
ti. 1861; Oligocene, Italy 

■'Tuberculodolium Sacco, 1891:9. Type species (by subsequent 
designation of Vokes, 1986:178): Eudolium antiquum Sac- 
co, 1890; Oligocene, Italy 

?Simplicodolium Sacco, 1891:13. Type species (here designat- 
ed): Pyrula fasciata Borson, 1821; Pliocene, Italy. 

Remarks: The shells of Eudolium species differ from 
those of Tonna Brunich, 1771 in having more narrowly 
tapered bases, consistently narrow, widely spaced spiral 
cords with narrow secondary spirals, and fine axial sculp- 
ture on all teleoconch whorls. The two groups are cer- 
tainly closely related, with similar shells, radulae, jaws 
and external anatomies. An operculum is lacking in both 
groups, at least in the adults. Eudolium species are most 
commonly encountered at 200-600 m depth, while few 
Tonna species range much deeper than 50 m. Although 
Eudolium has been placed as a subgenus of Tonna by 
some authors {e.g.. Dall, 1889a, b; Vredenburg, 1919; 
Thiele, 1929; Kilias, 1962), 1 prefer to treat it at generic 
level because of the distinctive shell facies and deeper 
center of bathymetric distribution. 

As here interpreted the genus Eudolium contains three 
Recent species: E. crosseanum (Monterosato, 1869), £. 
bairdii (Verrill & Smith, 1881), and £. pyriforrne (Sow- 
erby, 1914). Of other species that have been referred 



B A Marshall. 1992 



Paae 25 



here, Dolium [Eudolium) verrilli Dall, 1889 has been 
transferred to Hadroocorys Quinn, 19S0 iQuinn. 1980). 
while £. aulacodes Tomlin. 1927 belongs in Oocorys 
Fischer. 1883 (Kilbum. 1986). Parvitonna perselecta Ire- 
dale, 1931 was referred to Eudolium by Kilias (1962). 
but the holotype (.\MS C. 57790) is clearK a species of 
Tonna (sensu stncto). Tonna tessellatum (Bruguiere. 
1789). T. fasciatum (Bruguiere. 1789) and T. zonatum 
(Green. 1830) were referred to Tonna {Eudolium ) by 
Vredenburg (1919). but the> too belong in Tonna {sensu 
stricto). 

\okes (1986' interpreted Cossmann's 1 190-'3:lo9! state- 
ment "types de Galeodolium et de Tuberculodolium: E. 
antiquum Sacco, Cassid. mutica Michelotti. d'apres les 
types communiques par M. Sacco" as the first valid sub- 
sequent designation of t\pe species for Galeodolium and 
Tuberculodolium. Cossmann. however. inad\ertentK re- 
versed the order of the ""t\p>e sp>ecies and thus selected 
species that were not originalK included (ICZN .\rt. 69a). 
Although N'okes simpK corrected the order. Cossmann 
must be considered to have associated the names re- 
speciicely, so it is concluded that Cossmann's selection 
is invalid. .AccordingK. X'okes vl986) is considered to be 
the first subsequent designator of the typ>e species of these 
ta.xa. 

N'okes (1986) used Galeodolium as a subgenus of Eu- 
dolium for species with strong nodules and a refle.xed. 
denticulate outer lip. The differences between the t\pe 
si>eciesof Eudolium and Galeodolium Michelotti. 1S61: 
pi. 13. fig. 16: Sacco. 1S91: pi. 1. fig. li. however, are 
simpK matters of degree, because E. bairdii exhibits a 
smooth morphological transition bet^veen the extremes 
in its infraspecific \ariabilit\ (see below). Placement of 
Tuberculodolium and Simplicodolium in synon\mv is 
tentative pending study of the typ>e material, which was 
not available to me. Judging from published photo- 
graphs, the undoubtedK juvenile holotype of Eudolium 
antiquum Sacco. 1S90 .Ferrero Mortara et al.. 1984) 
seems narrower than confirmed species of Eudolium, 
while the lectotype of Pyrula fasciata Borson. 1821 (Pa- 
via, 1976) may be a juvenile Tonna. Because Galeodoli- 
um and Eudolium are regarded as synonyms, and as 
Eudolium and Tonna are undoubtedK confamilial. Ga- 



leodoliidae Sacco, 1891 is treated as a synonym of Ton- 
nidae Suter. 1913 (1825) (Tonnidae is conserved using 
ICZX Art. 40b). 

Eudolium sjjecies occur in temp)erate and tropical seas 
on soft substrata on continental shelves and slopes at 17- 
823 m depth. Undoubted Eudolium species are known 
from the EarK Miocene [e.g., E. aoteanum Beu. 1970 
and another, unnamed species from New Zealand; £. 
biornatum (Tate, 1894) from .Australia] and the Oligo- 
cene {e.g., Cassidaria mutica Michelotti, 1861). .As with 
Tonna species (Morton. 1991 1 the diet consists of holo- 
thurians: the gut of a specimen of E. bairdii from off 
Ma\or Island. New Zealand, contained ossicles of the 
apodid holothurian Protankyra rigida Pawson, 1965 (det. 
D. L. Pawson) Otherwise nothing is known of Eudolium 
biolog). 

Eudolium crosseanum : Monterosato. 1869) 

(figures 1-4. 9. 20. 23-26. 37 1 

Dolium crosseanum Monterosato, 1869:22S. pL 12. fig. 1: Trvon. 

1885:263. pi. 2. fig. 11 (figs. 11 and 12 transptKed. fig. 12 

= D. testardi Montrouzier. 1863). 
Doliopsis crosseana. — Monterosato. 1872:8; Coen. 1930:147. 

figs. 1, 2 [in part — figs. 6-9 = Galeodea echinophora vLin- 

naeus, 1767), P. Bouchet. personal communication]. 
Dolium Eudolium^ croaseanum. — Kobelt. 1908:155. pi. 126. 

fig. 1 I in part — figs. 2. 3. 4 = £. bairdii'. 
Eudolium crosseanum. — Wenz. 1941:1076. fig. 3066: Turner. 

1948:178 un part— pi. 81. figs. 1. 2. te.xt Hg. 5 = £. bairdii). 
Eudolium testardi. — Osima, 1943:132, pi. 4, fig. 3 (not Dolium 

testardi Montrouzier. 1863). 
Eudolium thompsoni McGintv. 1955:80. pi. 1. figs. 5, 6; .\bbott. 

1974:168, fig. 178S: Abbott & Dance. 1986:119. text fig 

(new synonym). 
Tonna {Eudolium) cros.seana. — Kilias. 1962:14, fig. 12 6: Pi- 

ani. 1977:27. 38. figs. 3. 6. 10 [in part — fig. 5 = Galeodea 

echinophora ^ Linnaeus. 1767), P. Bouchet personal com- 
munication, fig. 11 = £. bairdii]. 
Tonna ^Eudolium) thompsoni. — Kilias. 1962:16, fig. 12 5. 
NOT Dolium \EudoIium) crosseanum. — Dall, 1889a.b; Daut- 

zenberg & Fischer, 1906 (= E. bairdiiV 
NOT Dolium crosseanum. — Locard. 1S97 ^Talisman stn. 63 

= £. bairdii, and stn. 139 = a turrid. P. Bouchet. personal 

communication). 



Figure I. Label in Monterosato's handwriting gummed to dorsum of holot\-pe of Eutfo/ium crosseanum. Figure* 2-4. Eudolium 
crosseanum. 2. holot>pe. off SiciK. Hebrew Uni\ersit>. Jerusalem No. 21386 (79.8 x 58 mm). 3. Oregon station 3636. off Belize. 
USNM 751892 (32 8 x 22 m). 4. Oregon station 2021. off French Guiana. USNM 751885 (54 x .39.5 mm). Figures 5-8. Eudolium 
pyriforme. 5. 7. Bohol Straits, Philippines. NMNZ MF. 56381 (5. 51 x 34.5 mm; 7. 40.5 x 27.5 mm). 6. David Starr Jordan 
station TC 40 54. off Oahu. Hawaii. USNM 504464 i79 5 x 47 mm). 8. Southwest of Taiwan. NMNZ MF 56380 S4 5 x 55 mm). 
.\\\ photographs of shells are to scale 

Figure 9. Eudolium crosseanum. Oregon station 3636. off Belize. USNM 751892. detail of feleoconch sculptm-e. 7 x. Figiu-es 
10-19. Eudolium bairdii. 10. detail of teleoconch sculpture, of specimen in figure 16. 7 x. H. off .\ldermen Islands. New 
Zealand. D. Gibbs collection (76 x 49.5 mm). 12. 13. off Tosa-Shimuzu. Japan. NMNZ MF.563S3 ^figure 12. 51 x 37 mm; figure 
13. 59 X 43 4 mm> 14. Combat station C279. Straits of Florida. USNM 715002 (49.7 x 35.5 mml 15. off Scot Reef. Western 
Australia. NMNZ MF.57427 (58 x 38 mm). 16. Oregon 11 station 11133. off Yucatan Peninsula. Mexico. USNM 751930 v.>3 x 
22.7 mm). 17. SMIB 4 station D\V.55. southwest of New Caledonia. MNHN i45 x 28 5 mm' 18. Oregon station 5690. off Colombia. 
USNM 751869 (46 x 31.5 mm:. 19. Albatross station 5590, Sebuku Bay. Borneo. USNM 23911 ^46.5 x 31.7 mm). .Ml photographs 
of whole shells are to scale. 



Page 26 



THE NAUTILUS, Vol. 106, No. 1 




B. A. Marshall, 1992 



Page 27 




Page 28 



THE NAUTILUS, Vol. 106, No. 1 




Figures 20-22. Riglit lateral views of head-foot areas of male 
EudoliuTH animals. 20. E. crosseanum, Oregon station 3636, 
off Ik-lize, USNM 751892 21. E. pyrijormc. David HI an Jor- 
dan station T(;/40/5-l. off Oahu, Hawaii, USNM 804464 22. 
F.. hairdii. M ascareigncs III station 24, off Madagascar, MNHN, 
Scale lines: 20 = 2.5 mm; 21, 22 = 5 mm. ct, cephalic tentacle, 
pe, penis; pr, proboscis; pt, penial tentacle; sg, seminal groove. 



NOT Eudolium crosseanuni. — Tomlin, 1927; Barnard, 1963; 
Abbott, 1974; Okutani, 1983; Kilburn, 1986; Horikoshi, 
1989; Waren & Bouchet, 1990; Poppe & Goto, 1991 ( = 
E hairdii) 

Description: Shell up to 81 mm (est.) high, thin to 
moderately thick, periostracum thin, straw-colored. Pro- 
toconch deep yellowish brown; teleoconch irregularly 
maculated with yellowish brown on a white or buff white 
ground, some major spiral cords with small yellowish 
brown spots, mature outer lip typically with a pinkish 
flush. Protoconch primarily conchiolin, conical, of about 
5 convex worls, 3.46-4.44 mm in diameter, sculptured 
with 3 rows of small periostracal spines. First whorl of 
calcareous mould of inner surface of protoconch (figure 
23 — exterior removed with sodium hypochlorite solu- 
tion) 317 ^m wide, sculptured with very fine irregular 
network of crisp threads that enclose minute, crowded, 
roughly circular pits. Second whorl with fine, crisp axial 
riblets and 2 similar crisp spiral threads, one at shoulder 
angulation, the other on ramp between shoulder spiral 
and suture. Shoulder spiral commencing immediately, 
ramp spiral commencing three quarters of a whorl later. 
Axial ribs and spiral cords becoming obsolete early on 
2nd whorl. Subsequent whorls essentially smooth, round- 
ed. Teleoconch of up to 4 evenly convex whorls, sculp- 
tured with numerous rounded spiral cords that multiply 
by intercalation; and fine, crisp, crowded axial riblets; in 
rare specimens some spiral cords becoming very weakly 
nodular on last adult whorl. Thin specimens with a thin, 
flared, weakly dentate outer lip and smooth columella 
and parietal area. Thickened specimens with strongly 
dentate outer lip, spirally plicate columella, and with 1- 
3 small spirally elongate denticles on parietal area close 
to insertion. 

Animal: (figure 20: subadult male, shell height 22.3 mm, 
Oregon stn. 3636) Everted proboscis large, very wide 
but longer than broad, thin-walled. Cephalic tentacles 
small, slender, rounded in cross section, gently tapered, 
tips roimded. Eyes small, in rounded swellings on outer 
sides of tentacles near bases. Penis base well behind base 
of right cephalic tentacle, penis very large, spathulate, 
laterally compressed, tip broadly rounded, deeply grooved 
along ventral edge to tip, minute tentacle protruding 
from end of groove at distal extremity. Operculum absent 
in adults. Jaw plates (figure 24) ovate, thin, each with 
strong, thickened, hooked anterior projection. 

Raditla: (figures 25, 26) with the formula 2.1.1.1.2, teeth 
curved, sharply pointed, very stout, strongly interlocked. 
Central tooth broad; cutting area acutely angulate, 7-9 
small sharp cusps on each side, terminal cusp large, long, 
narrow; posterolateral projections on face strong, sharp. 
Lateral teeth strongly longitudinally flanged along outer 



Figures 23-26. Eudolitun crosseanuni. 2.'{. protoconch with periostracum removed, Oregon station 5914, I^esser Antilles, USNM 
751903. 24. (jaw plate), 2.^. 26. (radnla) from shell 22 mm high. Oregon station 3636. off Belize, USNM 751892. 25. width of 
radula. 26. detail of central, lateral ;ind marginal teeth Figures 27-.'{0. Eudoliunt pyrijornie. Hadiila from shell 41 mm high, 
MUSORST(JM 6 station I)W39i, Loyalty Islands. 27. width of radula 28-30. details of central and lateral teeth. Scale bars: 27 
= 0.5 mm, others = 1 mm. 



B. A. Marshall, 1992 



Page 29 




Page 30 



THE NAUTILUS, Vol. 106, No. 1 




Figures 3 1 -.'ib. Etidolium hairdii. 3 1 -.'$3. radula from slu'll 24 mm higli, Straits of Florida, Florida State Museum L'F 26666. 
31. detail of central and lateral teeth from anterior end of radular ribbon representing earliest teeth formed. 32. detail of central, 
lateral and marginal teeth at posterior third of radular — note size and morphological differences compared with teeth formed 
earlier (3 1 ). 3.3. width of radula at posterior third. .34. detail of central and lateral radular teeth from shell .57 mm high, Mascareignes 
III station 24, oH' Madagascar, MNHN. 3,'>, 36. radula from shell 76 mm high, off Aldermen Islands, New Zealand, D. Gibbs 
colleclion. 3.'j. width of radula. 36. detail of central tooth Scale bars: 35 = 0.5 mm, others = 0.1 mm. 



edges to interlock with inner marginals, terminal cusp 
very large, 8-11 fine cusps heliiiKl on outer edge. Mar- 
ginal teeth similar, without secondary cusps. 

Type data: Dolium crosseanum. HOLOTYPE Coen 
collection (No. 2521), Hebrew I'niversity, Jeru.salem 
21386, off Palermo, Italy; Eudoliurn tlwinpsani. HO- 
LOTYPE Florida State Museum, Gainsville, Florida UF 



170426, Triton stn. 1206, off Sombrero Ke\ Light, Flor- 
ida Ke\s, 137 m 

Other material examined: WESTERN ATLANTIC— 

off Cape Ilatteras, South C^arolina, south to off French 
(iuiana, 35°06'N-7°18'N (56 specimens in 37 lots USNM). 

Distribution: (figure 37) Mediterranean and Western 
Atlantic, from Cape Hatteras to French Guiana; depth 



B. A. Marshall, 1992 



Page 31 




Figure 37. Map showing ihstrihution of Eudolium crosscanum (•) and £. pyrifornie (A) 



range of material examined 17-914 m, maximum known 
Hving occurrence 457 m. 

Remarks: As discussed by Piani (1977) two specimens 
have been interpreted as the holotype of D. crosseanum, 
one in the Monterosato collection, Museo Zoologico, Roma, 
the other in the Coen collection, Hebrew University of 
Jerusalem. Piani correctly concluded that the Rome spec- 
imen cannot be the holotype because it is extensively 
covered by a brown periostracum and has conspicuous 
nodules, characters not mentioned in the rather detailed 
original description, and because it lacks the maculations 
recorded by Monterosato. From comparison of dimen- 
sions of the Rome and Jerusalem specimens with Mon- 
terosato's original published measurements, Piani (1977: 
37) concluded that the Jerusalem specimen was the clos- 
est match but because there were discrepancies he sug- 
gested that it was perhaps a third example and thus by 
implication not the holotype. Piani (1977: figs. 6, 10) 
reproduced Coen's (1930: figs. 1, 2 "tipo") illustrations 
and a photograph (provided by H. K Mienis) of the 
Jerusalem specimen. Upon receipt of the Jerusalem spec- 
imen (figures 1, 2), kindly lent to me by H. K. Mienis, 
it was immediately obvious that Piani's illustrations are 
of one and the same specimen, though this was neither 
unequivocally stated by Piani nor evident from the il- 
lustrations. 

It transpires that the dimensions of the Jerusalem spec- 
imen were wrongly quoted by Piani (1977:37), as they 
in fact closely accord with Monterosato's (1869) original 
measurements ( "long 81; diam. maj. 56 mill.; apert. 62 
mill, longa, 28 lata "): height 79.8 mm (lacks proto- 
conch — estimated height when intact 81 mm), diameter 
58 mm, height of aperture 62 mm, width of aperture 28 



mm. The general shell facies, color and color pattern of 
this specimen agree closely with the original description, 
while the number of denticles on the outer lip (32) are 
closely accordant: Monterosato stated that there were 33 
denticles, but it is possible that a low swelling at the 
abapical extremity of the lip was also counted. The only 
discrepancy is the thickness of the outer lip, which ranges 
from 3.5 to 4.0 mm over the denticulate area, and not 
3 mm as stated by Monterosato. The smooth zone at the 
adapical extremity, however, is 3 mm thick, suggesting 
that Monterosato may have measured it there to avoid 
including the denticles. Diagnostic features of this par- 
ticular specimen that were realised on the original draw- 
ing include the distinctive pattern of chipping at the rim 
of the anterior siphonal canal, and the growth scar in 
front of the columella. Incontrovertible proof that this 
specimen belonged to Monterosato is the label in Mon- 
terosato's handwriting gummed to the dorsum (figure 1) 
"Genus Doliopsis mihi inedito' . 

The specific identity of the Rome specimen is uncer- 
tain, but judging from Piani's (1977: fig. 11) illustration 
it is much more strongly nodular than any specimen of 
E. crosseanum seen during the present study and seems 
more likely to be E. hairdii (P. Bouchet concurs). Other 
published records of E. crosseanum from the Mediter- 
ranean apart from references to the holotype are based 
on misidentification (see synonymy). 

By direct comparison of the holotypes, the Western 
.Atlantic E. thompsoni McGinty, 1955 is specifically in- 
distinguishable from E. crosseanum. Judging from the 
fact that the species has only been obtained once in the 
Mediterranean yet it is relatively common in the western 
Atlantic, it is likely that the Mediterranean specimen is 
the result of a chance introduction of a teleplanic larva 



Page 32 



THE NAUTILUS, Vol. 106, No. 1 




Figure 38. N4ap showing distribution of Eudolium bairdii. Eastern Atlantic distributions provided by P. Bouchet (personal 
comnuinication). 



transported by the Gulf Stream from Western Atlantic 
populations (see Discussion). 

Eudolium crosseanum is closely similar to the Western 
Pacific and Hawaiian species E. pyriforme (see below). 

Eudolium crosseanum was the first species described 
by Monterosato in an illustrious malacological career, and 
the holotype must have been one of his most prized 
possessions. He would have been impressed to know that 
his specimen would still be the only one known from the 
Mediterranean 122 years after it was described, and that 
it would remain the largest known example of the species. 



Eudolium pyriforme (Sowerby, 1914) 
(figures 5-8, 21, 27-30, 37) 

Doliuni pyriforme Sowerby, 1914:37, pi. 2, fig. 14. 

Eudolium piiriformc. — Osinia, 1943:131, pi. 5, fig. 4; May- 
bloni, 1951:282; Kira, 1959:55, pi, 22, fig, 5; Garrard, 1961: 
17; Kira, 1962:59, pi. 23, fig. 5; Okutani, 1964:400, pi. 1, 
fig. 23, te.xt fig. 2; Habe & Kosuge, 1967:66, pi, 25, fig. 12; 
Kuroda, Habe & Oyama, 1971:135, pi. 37, fig. 4; Wolfe, 
1974:3, text fig.; Powell, 1974:201; Cernohorsky, 1976:1, 
fig. 1; Cernohorsky, 1978:61, pi. 16, fig. 7; Powell, 1979: 
163; Kosuge, 1985:59, pi. 22, fig. 6; Abbott & Dance, 1986: 
116, text fig.; Kilburn, 1986:4, fig. 11; Okutani, 1988:78, 
fig 46. 

Description: Shell up to 85.5 (est.) mm high, thin; peri- 
ostracurn thin, translucent, straw-colored. Protoconch 
yellowish brown; teleoconch sparsely to densely irregu- 
larly maculated with pale to deep yellowish brown, major 
spiral cords alternately spotted yellowish brown and white 
on a buff white ground, outer lip of many specimens 



with a pinkish flush. Protoconch primarily conchiolin, of 
about 5 convex whorls, 2.95-3.45 mm in diameter, es- 
sentially smooth (worn?). Internal calcareous mould not 
seen. Teleoconch of up to 4.6 evenly convex whorls, 
sculptured with numerous rounded spiral cords that mul- 
tiply by intercalation, and fine axial riblets, axials crisp 
on first 3 whorls, less crisply defined on subsequent whorls, 
no nodules. Outer lip of most specimens weakly thick- 
ened in adults, but a few specimens are moderately thick- 
ened and dentate; columella and parietal area simple. 

Animal: (figure 21 — adult male minus radula that had 
been extracted when received from one of 7 shells 64- 
79.5 mm in height, from off Oahu, Hawaii — USNM 
804464) externally similar to that of E. crosseanum (see 
above) but differing in that the penis arises nearer outer 
base of right cephalic tentacle. Jaw similar to that in £. 
crosseanum. 

Radula: (figures 27-30 — juvenile female 22 mm shell 
height, Oregon stn. 3636, off Belize City, USNM 751892) 
similar to that of £. crosseanum except that central and 
lateral teeth have smaller cusps, and sides of central tooth 
subparallel instead of markedly tapered. 

Type data: Dolium pyriforme: HOLOTYPE: The Nat- 
ural History Museum, London 1915.1.6.148, off Kii, Ja- 
pan. 

Other material examined (73 specimens): INDIAN 
OC>EAN — Trawled off Umgababa, Natal, South Africa 
(1 NMP); trawled ofi^ Durban, South Africa, 421 m (1 
NMP); trawled off Bazaruto I., Mozambique (1 NMP); 
12°39.8'S, 48°16.2'E, N Madagascar, 375-385 m (1 
MNHN); I2''42.4'S, 48°14.3'E, N Madagascar, 285-295 



B. A. Marshall, 1992 



Page 33 



m (1 MNHN); 12°42.9'S, 48°12.1'E, N Madagascar, 445- 
455 m (1 MNHN); 12°43'S, 48°15'E, 300-340 m (1 
MNHN); 20°58'S, 55°15'E, off Reunion, 450-580 m (1 
MNHN); 22°14.7'S, 43°04.5'E, S Madagascar, 470-475 
m (1 MNHN); Crevettiere 1986 stn. 80, 22°17'S, 43°04'E, 
SW Madagascar, 530 m (1 MNHN); Crevettiere 1986 
stn. 76, 22°22'S, 43°03'E, SW Madagascar, 530 m (2 
MNHN); Crevettiere 1986 stn. 56, 22°26'S, 43°05'E, SW 
Madagascar, 435 m (2 MNHN); Crevettiere 1986 stn. 
57, 22°26'S, 43°06'E, SW Madagascar, 460 m (1 MNHN); 
Crevettiere 1986 stn. 58, 22°26'S, 43°06'E, SW Mada- 
gascar, 440 m (1 MNHN); SoWa stn. 01/84/74, 14°16.5'S, 
122°54.4'E, NW of Collier Bay, Western Australia, 302 
m (1 WAM); Soela stn. 01/84/54, 15°51.2'S, 120°44.3'E, 
WNW of Lacepede Archipelago, Western Australia, 350- 
348 m (6 WAM) JAPAN— off Tosa Shimuzu, ca. 350- 
400 m (1 NMNZ); off Ashizuri-misaki (2 MCZ); oft Tosa, 
ca. 183 m (1 MCZ); off Kii (1 MCZ). TAIWAN— SW of 
Taiwan, ca. 128-183 m (5 NMNZ). PHILIPPINES— 
Albatross stn. 5289, Batangas Bav, Luzon, 315 in (2 
USNM); Coriolis MUSORSTOM 3 stn. CP103, of Min- 
doro, 193-200 m (1 MNHN); Coriolis MUSORSTOM 2 
stn. CP83, 13°55'N, 120°30'E, off Mindoro, 318-320 m 
(1 MNHN); Bohol Straits, ca. 200 m (5 NMNZ); Albatross 
stn. 5519, off Pt Tagalo, N. Mindanao, 333 m (2 USNM); 
Albatross stn. 5518, off Pt Tagalo, Mindanao, 366 m (2 
USNM); 9°43'S, 130°00'E, Timor Sea, ca. 250 m (2 
NMNZ). HAW Ml— David Starr Jordan stn. TC-40-54, 
21°01.6'N, 156°43.00'W, off Oahu, 223 m (7 USNM). 
QUEENSLAND and NEW SOUTH WALES— Off Cape 
Moreton, 165 m (1 AMS); Kapata stn. 71-08, ca. 32°46'S, 
152°16'E, off Newcastle, 280-549 m (1 AMS); Kapala 
stn. K75-05-03, 33°02'S, 152°31'E, off Newcastle, 475 m 
(1 AMS); between Newcastle and Sydney, 182-549 m (1 
AMS); off Broken Bay, 137 m (1 AMS); between Sydney 
and Norah Head, 412-457 m (1 AMS); Kapala stn' K76- 
07-01, 33°33'S. 151°59'E, off Broken Bay, 384 m (1 AMS); 
Kapala stn. K76-24-05, 33°33'S, 151°59'E, off Broken 
Bay, 373-366 m (1 AMS); off Svdnev, 420-440 m (1 
AMS); 35°50'S, 150°34'E, off Batemans Bay, 366 m (1 
AMS). CHESTERFIELD REEFS— Corio/'is MUSOR- 
STOM 5 stn. 376, 19°51'S, 158°30'E, 280 m (1 MNHN). 
NEW CALEDONIA— Vanfcan MUSORSTOM 4 stn. 
CP193, 18°56'S, 163°23'E, off d'Entrecasteau Reefs, 415 
m (1 MNHN); Vaiiban SMIB 3 stn. DW28, 22°47'S, 
167°12'E, 394 m (1 MNHN); Vaiiban SMIB 1 stn. DW2, 
22°52'S, 167°13'E, 415 m (1 MNHN); Vaiiban SMIB 2 
stn. DW15, 22°53'S, 167°11'E, 375-402 m (1 MNHN); 
Vauban SMIB 2 stn. DW8, 22°54'S, 167°13'E, 435-447 
m (1 MNHN); Vauban SMIB 2 stn. DW17, 22°55'S, 
167°15'E, 428-448 m (1 MNHN). LOYALTY IS- 
LANDS— A/is MUSORSTOM 6 stn. DW391, 20°47'S, 
167°06'E, off Ouvea, 390 m (1 MNHN); Alis MUSOR- 
STOM 6 stn. CP 465, 21°04'S, 167°32'E, off Lifou, 480 
m (1 MNHN). NEW ZEALAND— off Aldermen I., ca. 
366 m (1 Gardner coll'n, Auckland); BS843 (0.589). 
37°14.6'S, 176°51.0'E, Rangatira Knoll, NW of White I., 
407-162 m (fragment NMNZ); off Tokomaru Reef, Gis- 
borne, ca. 220 m (1 Auckland Institute and Museum). 



Distribution (figure 37): Southern Africa, Madagascar, 
Reunion, Japan, Taiwan, Philippine Islands, Indonesia, 
Hawaii, Australia, New Caledonia, northern New Zea- 
land; depth range of material examined 137-580 m, 
deepest known living occurrence 390 m. 

Remarks: Eudolium pyriforme is extremely similar to 
E. crosseanum in shell color, color pattern, protoconch 
morphology, shell thickness, size relative to the number 
of whorls, and in size attained. Although they are also 
similar in sculpture, E. pyriforme differs in having axial 
riblets that are consistently lower, broader and less sharp- 
ly defined than those of E. crosseanum, especially where 
traversing the spiral cords. As described above, a single 
adult male specimen of E. pyriforme differs from that 
of an immature E. crosseanum in having the penis base 
close beside the right cephalic tentacle instead of well 
behind it (figures 20, 21). The external anatomies of 
males and females are otherwise similar. Although the 
position of the penis seems likely to be a major distin- 
guishing feature, with only two different-sized animals 
for comparison it is impossible to ascertain whether or 
not its position is infraspecifically stable or changes dur- 
ing ontogeny. 

As described and illustrated herein (figures 25-30), the 
radula of £. pyriforme differs from that of E. crosseanum 
in the shape of the central tooth and in the size of the 
secondary cusps. With only a single radula from each 
species from dissimilar-sized specimens, however, it is 
impossible to ascertain either the degree of infraspecific 
variability at equivalent size, or the degree of morpho- 
logical change during ontogenetic development (see E. 
bairdii below). 

Specimens of £. pyriforme tend to be a little narrower 
than those of E. crosseanum. although there is a broad 
overlap in shape. Shell height/diameter ratio in E. pyr- 
iforme ranges from 1.41 to 1.72 (mean 1.53, SD 0.095, 
n = 30) as against 1.31-1.66 (mean 1.44, SD 0.085, n = 
18) in £. crosseanum. Specimens from off Hawaii are 
uniformly narrowly ovate (USNM 804464, height/di- 
ameter ratio 1.59-1.72, mean 1.67, SD 0.047, n = 7, 
figure 6). Some specimens from Japan (MCZ 293903), 
the Timor Sea (NMNZ MF.57428), New South Wales 
(AMS C.68870) and Mozambique (NMP H.7800) are as 
narrowly ovate as specimens from Hawaii, however, and 
there is complete integration between narrow and broad 
forms in material from west of Hawaii (height/diameter 
ratio 1.41-1.62, mean 1.48, SD 0.052, n = 22). 

Eudolium bairdii (Verrill & Smith, 1881) 
(figures 10-19, 22, 31-36. 38) 

Doliuni bairdii Verrill & Smitii in Verrill, 1881:299; Verrill, 
1882:.51.5; Verrill, 1884253. pi. 29, figs. 2a,b. 

Dolium hayardi (sic,).~Paetel, 1888:221. 

Dolium (Eudolium) crosseanum. — Dull, 1889a:232 (in part), 
pi. 15. fig, 5. 

Dolium {Eudolium) crosseanum. — Dall, 1889b:134, pi. 15, fig. 
5, pi. 44, fig. 2, pi. 62, figs. 83, 83a; Dautzenberg & Fischer, 
1906:38 (not D. crosseanum Montern,sato, 1869). 



Page 34 



THE NAUTILUS, Vol, 106, No. 1 



Dohum crosseamim. — Locard. 1897293 (in part not D. cros- 
seanurn Monterosato, 1869 — Talisman stn. 139 record is 
based on a turrid: P. Bouchet, personal communication). 

Dolium (Eudolium) cro.sseanum var. solidior Dautzenherg & 
Fischer, 1906:38, pi. 3, fig. 1. {new synonym). 

Dolium (Eudolium) crosseanum. — Kobelt. 1908:1.5.5, pi 126, 
figs. 2, 3, -1 (in part — fig. 1 = £. crosseanum). 

Morio lineata Schepman, 1909:124, pi, 10, fig, 5 (new syn- 
onym). 

Eudolium crosseanum. — Tomlin, 1927:82, fig, 4b; Barnard, 
1963:8; .\bbott, 1974:168, pi, 6, fig, 1787; Okutani, 1983: 
264, text fig,; Kilburn, 1986:4, fig, 12; Horikoshi, 1989:63, 
pi, 15, fig, 11; Waren & Bouchet, 1990:89, figs, 11, 78, 79; 
Poppe & Goto, 1991:128, pi, 22. figs, 1, 2 (not D. cros- 
seanum Monterosato, 1869), 

Eudolium lincatum. — Osima, 1943:133, pi, 5, fig, 1, 

Oocorys lineata.— Turner . 1948:178, 190; .Abbott & Dance, 
1986:116, te,\t fig, 

Eudolium inflatum Kuroda & Habe, 1952:56 (replacement 
name for Eudolium lincatum Osima not Schepman); Ku- 
roda & Habe, 19.57:28, figs, 2, 4, 5; Kira, 1962:.59, pi, 23, 
fig, 4; Okutani, 1988:77, fig. 45; Bieler & Petit, 1990:137 
(new synonym). 

Eudolium lincatum inflatum — Kira, 195444, pi. 22, fig, 4; 
Kira, 19.59:.55, pi, 22. 

Eudolium kuroharai Azunia, 1960:98, pi, 1, fig, 8, text fig, 1 
(new synonym). 

Tonna (Eudolium) crosseana. — Kilias, 1962:14 (in part), 

Dolium (Eudolium) crosseanum solida (sic), — Settepassi, 1971, 
appendix vii, text figs, 

Tonna (Eudolium) crosseana. — Piani, 1977, fig. 11, 

Oocorys solidior. — Piani, 1977:38, figs. 1, 2. 7, 

Description: Shell up to 75,5 mm liigli, thin to rather 
thick, periostracum thin, straw-colored. Protoconch deep 
yellowish to reddish brown; teleoconch white or buff 
white, major spiral cords yellowish to reddish brown, 
outer lip white. Protoconch primarily conchiolin, conical, 
of about 5 convex whorls, 3.00-4.75 mm in diameter, 
sculptured with 3 narrow spiral threads with small spines 
(in well preserved specimens). Calcareous mould of inner 
surface of protoconch (Waren & Bouchet, 1990: fig 79) 
similar to that of E. crosseanum but with 3 instead of 2 
spiral threads. Teleoconch of up to 4.4 evenly convex 
whorls, sculptured with numerous rounded spiral cords 
that iTiultipK by intercalation, and fine axial riblets, axials 
less crisply defined after third whorl, last adult whorl 
with or without weak to strong nodules on major spiral 
cords. Shell ranging through intermediates from thin to 
rather thick. Thin specimens with thin, flared, weakly 
dentate outer lip and simple columella and parietal area. 
Most thickened specimens with thick, strongly dentate 
outer lip, thick spirally plicate callus at base of columella, 
and 1-3 spirally elongate denticles on parietal area below 
insertion. 

Animal: (figure 22, adult male, shell height 57.5 mm, 
Mascareignes III stn. 24, off Madagascar) externally sim- 
ilar to that of E. pyrijorme. 

Radula: (figures 31-36) (from shells 25, 57 and 76 mm 
in height) similar to those of £. crosseanum and E. pyr- 
ijorme. With increasing size central and lateral teeth 



enlarging while size of secondary cusps and area they 
occupy remain rather static. Secondary cusps ultimately 
becoming obsolete on laterals in large adults. 

Type data: Dolium bairdii: HOLOTYPE USNM 51385 
(51 X 25 mm). Fish Hawk station 945, 39°58'N, 71°13'W, 
off Barnegat, New jersev, 379 m; Dolium (Eudolium) 
crosseanum var. solidior. HOLOTYPE Musee Oceano- 
graphique, Monaco, 38°52.50'N, 27°23.05'W, off the 
Azores, 599 m; Morio lineata: HOLOTYPE Zoological 
Museum, Amsterdam 3.09.008, Sihoga stn 173, 3°27'S, 
131°0.5'E, Ceram Sea, 567 m; Eudolium inflatum: TYPE 
MATERIAL (Osima, 1943, pi. 5, fig. 1) possibly in Oshi- 
ma collection, which was destroyed during World War 
II (A. Matsukuma, personal communication), off Wa- 
kayama Prefecture, Japan; Eudolium kuroharai: HO- 
LOTYPE in Mr. M. Azuma's private collection, Takara- 
zuka City, Japan (not seen), off Tosa, Japan. 

Other materia! examined (161 specimens): WEST- 
ERN ATLANTIC OCEAN— Barnegat, New Jersey, south 
to off French Guiana, 39°58'N-7°37'N (12 specimens in 
9 lots MCZ, 91 specimens in 43 lots USNM). WESTERN 
INDIAN OCEAN— 12°01'S, 49°26'E, off Madagascar, 450 
m, A. Crosnier (1 MNHN); 12°42'S, 48°14'E, off N. Mad- 
agascar, 380-375 m, A. Crosnier (1 MNHN); 12°43'S, 
48°12'E, off N. Madagascar, 445-455 m, A. Crosnier (1 
MNHN); Marion-Dujresne crusise 32. stn. CP 144, 
20°50'S, 55°35'E, off Reunion, 605-620 m (1 MNHN); 
Mascareignes III stn. 37, 22°18'S, 43°05'E, off S Mada- 
gascar (1 MNHN); Crevettiere 1986 stn. 80, 22°17'S, 
43°04'E, SW Madagascar, 530 m (1 MNHN); Crevettiere 
1986 stn. 81, 22''23'S, 43°03'E, SW Madagascar, 525 m 
(2 MNHN); Mascareignes III stn. 24, 22°31'S, 43°07'E, 
off S Madagascar, 430-460 m (1 MNHN); trawled off 
Umgababa, Natal, South Africa (1 NMP); trawled off 
Tongaat, Natal, South Africa (1 NMP); off Cape St. Fran- 
cis, South Africa (1 NMP). WESTERN AUSTRALIA— 
off Augustus I. south to off Rowlev Shoals (2 specimens 
in 1 lot NMNZ, 23 specimens in 19 lots W.AM, 1 specimen 
USNM). JAPAN— SW of Ashizuri misaki, Kochi Prefec- 
ture, trawled (1 MCZ); off Tosa Shimuzu, Kochi Prefec- 
ture, 350-400 m (2 NMNZ). PHILIPPINES— Corio/is 
MUSORSTOM 3 stn. CP106, 13°47'N, 120°30'E off Min- 
doro, 640-668 m (1 MNHN). MALA\Sl\— Albatross 
stn. .5592, Sebuku Bay, Borneo, 558 m (1 USNM); Al- 
batross stn. 5590, Sebuku Bav, Borneo, 567 m (1 USNM). 
NEW CALEDONIA— Vai/fcan MUSORSTOM 4 stn. 
DW156, 18°54'S, 163°19'E, Grand Passage, 525 m (1 
MNHN); Vauban MUSORSTOM 4 stn. DWT97, 18°51'S, 
163°21'E, Grand Passage, 550 m (1 MNHN); Vauban 
MUSORSTOM 4 stn. CP 171, 18°58'S, 163°14'E, Grand 
Passage, 425 m (1 MNHN); Jcan-Charcot BIOCAL stn. 
DW36, 23°09'S, 167°11'E, 650-680 m (1 MNHN); Alis 
SMIB 4 stn. DW55, 23°21'S, 168''05'E, 215-260 m (1 
MNHN). LOYALTY ISLANDS— A/is Ml'SORSTOM 6 
stn. CP 467, 2r05'S, 167°32'E, off Lifou, 575 m (1 
MNHN). NEW SOUTH WALES— off Port Stephens 
south to off Brush I. (5 specimens in 5 lots AMS). NEW 
ZEALAND— N of Mavor I., Bav of Plenty, 380-420 m, 



B. A. Marsliall, 1992 



Page 35 



coll. M. Huaki (1 NMNZ); off Aldermen I., New Zealand, 
400 m (1 D. Gibbs coll'n, Auckland). 

Distribution (figure 38): Atlantic, Mediterranean and 
Indo- Western Pacific, 17-823 m, deepest known occur- 
rence of living specimen 560 m. 

Remarks: Eiidoliiim bairdii is rendered high!} distinc- 
tive by the reddish to \ellowish brown spiral bands on 
the major spiral cords, a pattern in marked contrast to 
the irregular mottling of E. crosseanum and E. pyri- 
forme. Endolitim bairdii is indistinguishable from the 
Pacific form that has usualK been known as E. lineatum 
(see below). Compared with E. crosseanum and £. pyr- 
iforme at the same stage of growth, the primary spiral 
cords in E. bairdii tend to be more prominent, while the 
secondary spirals generally enlarge more slowly relative 
to the primaries. E. bairdii differs further in having con- 
siderably stronger a.xial riblets, especially where travers- 
ing the spiral cords. The axial riblets in £. bairdii tend 
also to be less crowded, numbering 3-7 per millimeter 
at the end of the second teleoconch whorl (mean 5.40, 
SD 1.04, n = 20), compared with 6-10 (mean S 33, SD 
1.07, n = 12) in £. crosseanum, and 6-11 (mean 8.69, 
SD 1.55, n = 13) in E. pyriforme. The calcareous internal 
mould of the protoconch of E. bairdii ( Waren & Bouchet, 
1990: fig. 79 — misidentified as E. crosseanum) differs 
from that of E. crosseanum (figure 23) in having 3 instead 
of 2 spiral threads. With only a single protoconch from 
each species for comparison, however, the degree of in- 
fraspecific variability in protoconch morphology is un- 
certain. 

Of the three known living Eudolium species, E. bairdii 
exhibits the greatest variation in shell morphology, un- 
doubted adults ranging from 32.5 mm (est.) to 75.5 mm 
in height, with lightly built or strongly thickened shells, 
and with or without nodules on the last adult whorl. 
There is smooth intergradation between all of the ex- 
tremes both within and between samples from the At- 
lantic and the Indo-Pacific. B\ direct comparison of ho- 
loty pes and topotypes, I am unable to detect any constant 
differences between D. bairdii, D. crosseanum var. so- 
lidior. M. lineatum, E. inflatum and E. kuroharai, and 
so it is concluded that they are all conspecific. It is sig- 
nificant, however, that thick-shelled forms with heavily 
thickened outer lips and strongly dentate apertures (ex- 
treme solidior form) (figures 14, 16) have no strict par- 
allel (figure 17) outside the Atlantic, while specimens 
from off eastern Australia are thick-shelled and often 
particularly heavily sculptured, yet they do not exhibit 
the apertural features of the solidior form. These differ- 
ences suggest that there may have been some genetic 
drift between isolated, probably largely self-recruiting 
populations, but these may also be non-genetic responses 
to local environmental conditions. The latter interpre- 
tation is suggested by the fact that a solidior-Vike form 
is also rarely exhibited by the Atlantic E. crosseanum 
(figure 4) yet not by E. pyriforme, its Indo-Pacific sister 
species. Heavy shelled forms (estimated height 34.5-61.0 
mm) are clearly adults and perhaps senescent, and it is 



likely that many lightly built forms (32.5-75.5 mm) are 
also mature. Although type specimens of D. bairdii and 
the synonymized taxa are not illustrated here, illustra- 
tions of strongly similar specimens are provided for ori- 
entation: D. bairdii, E. inflatum and E. kuroharai (figures 
12, 13, 15), D. crosseanum var. solidior (figures 14, 16), 
Morio lineata (figures 18, 19). 

E. bairdii and E. crosseanum have fully overlapping 
geographic and bathymetric ranges in the Atlantic (fig- 
ures 37, 38), and the two species have been taken together 
at three stations in the Western Atlantic {Oregon stn. 
2391, USNM 878126 and 751929 respectivelv; Oregon 
stn. 4911, USNM 878128 and 751872; Oregon II stn. 
11253, USNM 766104 and 878129). E. bairdii and £. 
pyriforme also have overlapping, geographic and bathy- 
metric ranges and are thus probably locally sympatric. 

DISCUSSION 

Although larvae of Eudolium species have not been ob- 
served in the field or reared in the laboratory, the pro- 
toconch morphologv and species distributions are char- 
acteristic of tonnoideans with teleplanic larvae 
(Scheltema, 1966; Laursen, 1981). Teleplanic larvae re- 
main planktonic for many months, in some species for 
a year or more, and can be transported great distances 
in ocean currents (Scheltema, 1966, 1971; Pechenik et 
ai, 1984). Species with teleplanic larvae often have ex- 
tremely wide amphioceanic, Indo-Pacific, or Indo-Pa- 
cific-Atlantic distributions in their benthic stage, es- 
pecially those with wide tolerances (Scheltema, 1986 and 
references therein). Although some localities for individ- 
uals in the benthic stage do not necessarily support self- 
recruiting populations, it is clear that widely separated 
self-recruiting populations are able to maintain their spe- 
cific integrity through periodic influx of larvae from up- 
current populations. 

Whether or not Eastern Atlantic and Mediterranean 
specimens respectively of £. bairdii and £. crosseanum 
arose from local self-recruiting populations is unknown, 
but judging from the fact that they are indistinguishable 
from Western Atlantic specimens and that both species 
are evidently far more common in the Western Atlantic 
than in the Mediterranean, it is probable that some if 
not all originated from Western Atlantic larvae that were 
transported by the Gulf Stream (Scheltema, 1986). While 
E. bairdii may well be at least partly self-recruiting in 
the Eastern Atlantic, this is unlikely to be true for £. 
crosseanum in the Mediterranean with a single con- 
firmed specimen. Specific integrity of the strongly iso- 
lated Hawaiian population of £. pyriforme is probably 
maintained through intermittent recruitment of larvae 
from the Western Pacific via the Kuroshio Current (Zins- 
meister & Emerson, 1979). 

It is likely that the two most closely related species, 
£. crosseanum and £. pyriforme originated from dis- 
junct populations of an ancestor that formerly had a 
continuous distribution in the Tethyian Atlantic-Indo- 
Pacific. Since Eudolium species are unknown living ei- 



Page 36 



THE NAUTILUS, Vol. 106, No. 1 



ther in the Eastern Pacific or from the Western Atlantic 
south of Columbia, it seems probable that distribution 
was continuous via Africa rather than America. Unfor- 
tunately, the southwestern coast of Africa has been rel- 
atively poorly sampled at depths most likely to yield 
Eudolium species (300-600 m) and it is unknown wheth- 
er or not they occur there. The apparent absence of £. 
crosseanum off South Africa suggests that larvae are 
incapable of reaching there from northeastern South 
America in the great counterclockwise South Atlantic 
gyral. Similarly, larvae of E. pyrifornie in the warm 
Aguihas Current are perhaps unal)le to survive injection 
into the Atlantic Ocean via the cold Bengueia CJurrent. 
.\tlantic and Indian Ocean populations of £. bairdii may 
be tulK isoialed 1)\ the same means. Unless hydroiogicai 
conditions ofl southwest .Africa were formerly more fa- 
vorable, perhaps during Pleistocene interglacials, £. 
bairdii and the ancestor of E. crosseanum and £. pyri- 
fornie may have ranged through the circumtropical Te- 
thys Ocean, in which case isolation of eastern and western 
populations would date from the late Miocene following 
establishment of the Middle Eastern Landbridge (Por, 
1986 and references therein). The second alternative is 
favored here. 

ACKNOWLEDGMENTS 

For the loan of material I am grateful to the following: 
A. G. Beu (Geology and Geophysics Division, Lower 
Hutt); P. Bouchet (Museum National d'Histoire Natu- 
relle, Paris); D. Gibbs (Auckland); C. Carpine (Musee 
Oceanographique, Monaco); M. G. Harasewych, R. S. 
Houbrick and S. Greenhouse (National Museum of Nat- 
ural History, Washington, D.C.); D. G. Herbert and R. 
N. Kilburn (Natal Museum, Pietermaritzburg); S. Kool 
and R. D. Turner (Museum of Comparative Zoology, 
Harvard); A. Matsukuma (National Science Museum, To- 
kyo); W. F. Ponder and I. Loch (Australian Museum, 
Sydney); T. Schiotte (Zoological Museum, Copenhagen); 
Y. Shibata and R. Yamanishi (Osaka Museum of Natural 
History); F. G. Thompson (Florida State Museum, Gains- 
ville); K. M. Way (The Natural History Museum, Lon- 
don); F. E. Wells (Western Australian Museum, Perth). 
For constructive comments on the manuscript I am grate- 
ful to A. G. Beu, P. Bouchet and R. N. Kilburn. Special 
thanks to R. Thomson (Victoria University, Wellington) 
for access to the scanning electron microscope; and to 
my staff colleagues J. Lord and J. Nauta for photography, 
A. Marchant for printing, K. Ryan for word processing, 
and R. Webber for the excellent illustrations of Eudolium 
animals. 

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THE NAUTILUS 106(1 ):39-42. 1992 



Page 39 



Buridrillia deroyortim. New Species from the Galapagos Islands, 
a Living Record of a Neogene Turrid Genus 



William K. Emerson 

American Museiiiii ol Natural Histor\ 
Central Park West at 79tl) Street 
New York, NY 1002-1, USA 



James H. McLean 

Natural Histors Museum of Los 
Angeles C^ountv 
900 Exposition Boulevard 
Los Angeles, C.\ 90007, USA 



ABSTRACT 

Buridrillia (U'riiyiiruni new species is described from 3 10 meters 
oft Isla Floreana (l>pe localit> I and 365 meters off Isla Duncan, 
Galapagos Islanils It is the first li\ing species of a turrid genus 
otherwise known from Neogene fossils from the New World 
tropics. The genus is assigned to Crassispirinae on shell and 
radular characters. It has a columellar plication, evidentK con- 
vergent with that of Borsoniinae, 

Key M'ords: Prosobranch gastropod; Turridae; Clrassisiiirinae; 
Buridrillia new species; Galapagos Islands, Ecuador. 



INTRODUCTION 

Andre and Jacqueline DeRoy were for many years avid 
collectors of Galapagan marine mollusks and are knowl- 
edgeable students of the molluscan fauna of the archi- 
pelago. Mr. DeRoy was a commercial fisherman who 
undertook numerous dredging operations, and Mrs. 
DeRoy spent much of her time collecting in shallow 
waters. For the past 30 years, they have contributed 
specimens to the American Museum of Natural History 
and the Natural History Museum of Los Angeles County 
for study and report. Among these specimens is a new 
species of turrid gastropod collected in the 1970s. We 
take great pleasure in describing this new turrid in honor 
of the DeRoys. 

The species described here is of more than usual in- 
terest for two reasons: it represents the first living record 
of a genus previously believed to have become extinct 
in the late Neogene, and study of its radular characters 
results in its assignment to a subfamiK other than that 
which comes to mind upon first examination of shell 
morphology. 

Abbreviations for institutions used in the text: AMNH — 
American Museimi of Natural History, New York; 
LACM — Los Angeles County Museum of Natural His- 
tory, Los Angeles. 



SYSTEMATICS 

Familv TURRIDAE Swainson, 1840 



Subfamily CRASSISPIRINAE Morrison, 1966 
Genus Buridrillia Olsson, 1942 

T\pe species (original designation): Clathrodrilla (Bur- 
idrillia) panarica Olsson, 1942:51. Pliocene of the Burica 
Peninsula, Panama and Costa Rica. 

Olsson (1964:98) extended the distribution of the type 
species to the Esmeraldas formation of Ecuador, from 
which sediments he also described five additional species 
of Buridrillia. He allocated the genus to "Drillinae" as 
then understood, which predated the proposal of Cras- 
sispirinae. 

In his revision of turrid classification, Powell (1966: 
62) assigned Buridrillia to the Borsoniinae, evidently based 
on the prominent columellar plication of the type species. 
However, the radular tooth of the new species B. deroij- 
orum is clearly of the duplex (terminology of Morrison, 
1966) or modified wishbone type, comparable to that 
illustrated by Kantor and Taylor (1991, fig. 2B) for Cli- 
onclla sinuata (Born, 1778). This tooth type is charac- 
teristic of (but not limited to) the Crassispirinae (McLean, 
1971; McLean in Keen, 1971). It is substantially different 
from the long hollow tooth of the Borsoniinae. 

Shell characters other than the columellar plication of 
Buridrillia are remiiiLscent of the crassispirine genera 
Ciassispira (Crassispira) Swainson (1840:152, 313), and 
Hindsiclava Hertlein & Strong (1955:227), in which there 
is a subsutural cord on the smooth shoulder, a deep sinus 
on the concave shoulder, and axial and spiral sculpture 
on the body whorl. The columellar plication of Buri- 
drillia is evidently convergent with that of Borsoniinae. 
Assignment of the genus to Crassispirinae is supported 
both on radular characters and the balance of shell char- 
acters. 

Buridrillia deroijonim new species 
(figures 1-14) 

Description: Shell sturdy, fusiform, attaining 57-1- mm 
in length. Spire acuminate, whorls shouldered, shoulder 
concave, with prominent subsutural cords; axial sculpture 
of nodose ribs; spiral sculpture of depressed lirae that 
decussate the axial ribs to form three rows of nodules; 



Page 40 



THE NAUTILUS, Vol. 106, No. 1 




Figures 1-14. BuridriUia dcroijorum new .specie.s. 1-3. Holotype, .AMNH 2321ti3; 4-6. I'aratype #8, LACM 2461; 7,8. Paratype 
#5, AMNH 232165 (Kigure.s l-«, slightly reduced, see Table 1 for measurements). 9, 10. Paratype #8, LACM 2461, spire enlarged 
about X 2.5. 1 1, 12. Paratype #10, Operculum, 1 1. View of interior side, 12. View of exterior side (Figures 11, 12, about X 2.5). 
13, ll-. Paratype #12, Raciuiar (icnlitinn (Scale bar for 1:3 = lOO^m, for 14 = .W^jin). 



body whorl with numerous spiral lirae that interrupt the 
axial sculpture on the anterior half. Protoconch lost on 
14 of the 15 specimens, protoconch poorly preserved on 
paratype #8 (Table 1), of 2 1/2 apparently smooth w horls 



(Figures 9, 10); bod\ whorl sculptured with 12 to 13 axial 
ribs. Aperture large, outer lip thin, inflated, explaned 
with a deep, L'-shaped anterior notch, posterior notch 
deep, w idely open; columella with a prominent plication 



W K. Emerson and J. H McLean. 1992 



Page 41 



Table 1. Buridrillia dcroijorum new species. Shell measure- 
ments in mm. Spires not preser\'ed, except for parat\pe #8. 
Number of specimens examined = 15 









# 




Length 


Widtli 


Whorls 


AMNH 232163 








Holotype 


50.1 


17.8 


7',/2 


AMNH 232164 








Paratvpe #1 


34.4 


13.7 


5'/2 


Paratvpe #2 


34.2 


13.0 


5'/2 


Paratvpe #3 


25.7 


10.5 


5'/2 


AMNH 232165 








Paratvpe #4 


52.4 


18.7 


7'/2 


Paratvpe #5 


45.5 


17.1 


6'/2 


Paratvpe #6 


31,0 


16.1 


61/2 


Parat\'pe #7 


24,3 


10,7 


6 


LA(.:M 2461 








ParatN'pe #8 


57.3 


194 


9'/2 


Paratvpe #9 


56.1 


18.8 


7'/2 


♦Paratvpe #10 








(operculum) 


48.9 


17.3 


7'/2 


Parat\'pe #1 1 


35 8 


13 1 


IVi 


*Paratvpe #12 








(radula) 


22.4 


9.8 


6'/2 


Paratvpe #13 


15 2 


6 4 


5 ',''2 


Paratxpe #14 


12,3 


5.1 


5 


Range 


12 3-57 3 


5.1-19.4 




Mean 


:> 1 1 


13 7 





■ Specimens providmg radula and operculinn tor photograpin 



on the proximal part of the pillar. Periostracum dense, 
flak\, greenish-brown. Shell color buff with light tan 
bands in the subsutural area and across base; aperture 
glossy white, tinged with tan. Operculum, small, thin, 
light brown, long (H=7.9 mm) and narrow (W=2.6 mm) 
with a marginal ridge and a terminal nucleus (Figures 
II, 12). Radula of both rows of marginal teeth only, of 
the duplex or modified wishbone type (Figures 13, 14). 

Type locality: North of Isla Floreana [also known as Santa 
Maria Island or Charles Island] I°I4'S, 90°26'VV, Gala- 
pagos Islands, Ecuador in 310 m, dredged bv the DeRoys, 
April 15, 1979. 

Specimens examined: (All dredged by A. and J. DeRoy, 
in the Galapagos Islands, see Table I for measurements): 
Holotype AMNH 232163 (Figures 1-3) and paratypes 
1-3, AMNH 232164, from the type locality; paratypes 
4-7, AMNH 232165, from the type locality, May 15, 
1978, in 310 m; paratypes 8-14; LACM 2461, off Isla 
Duncan [0°25'S, 90°43'W] in 365 m, March. 1979. 

Distribution: Known only from the Galapagos Islands 
from the t\pe localitv and off Isla Duncan, in 310 and 
365 m. 

Remarks: Of the five known Neogene species of Bur- 
idrillia from the Esmeraldas formation described bv Ols- 



son (1964), B. deroyonim most resembles the type spe- 
cies, B. panarica. from the Pliocene of Pacific Panama 
(type locality) and Costa Rica (Charco Azul formation) 
as well as from the Pliocene of Ecuador (Esmeraldas 
formation). The two species are of similar size and have 
a similar, well-developed columellar plication. From B. 
panarica it differs in having a raised subsutural cord 
rather than a constricted subsutural band, and in having 
much more strongK nodose axial ribs. 

The columellar plication of B. deroyorum is present 
on all fifteen specimens regardless of size. However, the 
strength of its continuation on the internal pillar may 
differ. A strong plication is visible through a hole on the 
back side of paratspe 10 made to extract the radula. 
However, the pillar in Paratype 8 (Figures 5, 6), which 
is visible through a naticid drill hole, has a very taint 
plication. Similar variation has been noted for fossil spe- 
cies of Buridrillia. Olssoii (1964:98) pointed out that some 
specimens of the type species have a well-marked col- 
umellar fold, but that others lack it or have slight swell- 
ings or raised lines. Other species described by Olsson 
have poorly marked folds or lack folds 

The present distribution of Buridrillia is that of a relict 
genus, a survivor in the eastern Pacific at the Galapagos 
refugium of a turrid group that was once more widely 
distributed in the New World tropics during the Neo- 
gene. 



ACKNOWLEDGMENTS 

We are grateful to Andre and Jacqueline DeRoy of Isla 
Santa Cruz, Galapagos Islands, for donating the type 
specimens to our respective institutions. We gratefully 
acknow ledge the assistance of our colleagues: Walter E. 
Sage, III, Stephanie Crooms and Andrew S. Modell of 
the AMNH provided technical assistance, word-process- 
ing and photographic services, respectively; C, Clifton 
Coney of the LACM operated the electron microscope 
at the I'niversit) of Southern California. We thank Yuri 
I. Kantor of the A. N. Severtzov Institute of Animal 
Evolutionary Morphology and Ecology, Moscow for 
commenting on a draft ol the manuscript. 

LITERATURE CITED 

Born. 1 1778 Index Rerum Naturalium Musei Caesarei V'in- 
dobonensis. Part 1; Testacea. Vienna, i-xlii + 1-458 p 

Hertlein, L. G. and A. M. Strong. 1955. Marine mollusks 
collected during the Askoy Expedition to Panama, Colom- 
bia, and Ecuador in 1941. Bulletin of the American Mu- 
seum of Natural History 107(2):159-318. 

Kantor, Y. I. and J. D. Taylor. 1991. Evolution of the toxo- 
glossan feeding mechanism: new information on the use 
of the radula. Journal of Molluscan Studies 57(11:129-134. 

Keen, .\. M 1971. Sea Shells of Tropical West .\merica ed 
2, Stanford University Press, xiv -I- 1064 p. 

McLean, J. H. 1971. A revised classification of the family 
Turridae, with the proposal of new subfamilies, genera, 
and subgenera from the eastern Pacific. The V'eliger 14(1): 
114- 130. 



Page 42 



THE NAUTILUS, Vol. 106, No. 1 



Morrison, J, P E. 1966, On the hiMiilii-sol Turridae, American 

Malacological L'nioii, Annual Reports, tor 1965: 1-2, 
Olsson, .'\. .\. 1942, Tertiars atitl (Quaternary fossils from the 

Buriea Peninsula of Panama and Costa Hiea Bulletins oi 

Amerieau Paleoiitolog) 27( 106): 1-106, 
Olsson, A. ,\, 1964, Neogene moiiusks Irom northwestern 

Ecuador, Paleoiitological Research Institution, Ithaca. New 

York, 256 p,, 38 pis. 



Powell, A, W B 1966, The moliuscan families Speightiidae 
and Turridae Bulletin of the .Aucklanil Institute and Mu- 
seum. 5: 184 p., 23 pis, 

Swainsou, W, 1840, .\ Treatise on Malacology or the Natural 
C;lassification of Shells and Shell-fish, London, 419 p. 



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rHE NAUTILUS 



Volume 106, Number 2 
June 5, 1992 
ISSN 0028-1344 

A quarterly devoted 
to malacology. 



Marine Biological Laboratory 
LIBRARY I 

JUN 1 5 1992 




EDITOR-IN-CHIEF 
Dr. M. G. Harase-wyeh 
Division oi Mollusks 
National Museum of 
Natural Histor\ 
Smithsonian Institution 
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CONSULTING EDITORS 

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Field Museum of 
Natural History 
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Dr. Robert T. Dillon, Jr. 
Department of Biology 
College of (;harleston 
Charleston, SC: 29424 



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Department of Living Invertebrates 

The American Museum of Natural 

History 

New York, NY 10024 



Dr. Robert Hershler 
Division of Mollusks 
National Museum of 
Natural Histors' 
Smithsonian Institution 
Washington, DC 20560 

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Division ot Mollusks 
National Museum of 
Natural History 
Smithsonian Institution 
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Mr. Richard I. Johnson 
Department of Mollusks 
Museum of ("omparative Zoology 
Harvard University 
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THEt^NAUTILUS 



CONTENTS 



Volume 106, Number 2 
June 5, 1992 

ISSN 0028-1344 

Mari ne Biological lahnr^tf;;^'] 
LIBRARY 

Two new species of Cancellariidae (Gastropocfc: JUN 1 S IQQ? 
Neogastropoda) from Brazil 1 43 

New species of Solariella (Gastropoda: Trochidae) from the 

western Atlantic Ocean 50 

A new fossil land snail of the genus Hemitrochus from 

Bowden, Jamaica 55 



M. G. Harasewych 
Richard E. Pelil 
Andre V erhecken 



James F. Quinn, Jr. 



Glenn A. Goodfriend 



R. Douglas Hunter 
John F. Bailey 


Dreissena pohjmorpha (Zebra mussel): Colonization of soft 
substrata and some effects on unionid bivalves 


60 


Edward J. Petueh 


New Ecphoras (Gastropoda: Thaididae: Ecphorinae) from 

the Calvert Formation of Maryland (Langhian Miocene) 


68 


Elizabeth Potter 
Michael A. Rex 


Parallel development-depth trends in deep-sea torrid snails 

from the eastern and western north Atlantic 


72 


John K. Tucker 


Drillia maclcani, new name for Drillia sintiosa McLean 

and Poorman, 1971 (Gastropoda: Turridae) 


76 



Notices 



58th ANNUAL MEETING OF THE AMERICAN 
MALACOLOGICAL UNION 

The 58th annual meeting of the American Malacolog- 
ical Union will be held August 2-7, 1992, at the Hyatt 
Sarasota, Sarasota, Florida. Local assistance is being pro- 
vided by the Sarasota Shell Club and the Mote Marine 
Laboratory. The meeting will feature contributed pa- 
pers, bourse and exhibits, a collectors' evening, auction, 
banquet, and fieldtrips to marine, freshwater, and ter- 
restrial habitats and a nearby fossil pit. Two international 
symposia are planned: Biology of Caribbean Mollusks, 
organized by Dr. Riidiger Bieler [Department of Zool- 
ogy, Field Museum of Natural History, Roosevelt Road 
at Lake Shore Drive, Chicago, IL 60605, Tel. (312) 922- 
9410, ext. 270, FAX (312) 663-5397], and Advances in 
Gastropod Phylogeny, organized by Dr. Terrence M. 
Gosliner [Department of Invertebrate Biology and Pa- 
leontology, California Academy of Sciences, Golden Gate 
Park, San Francisco, CA 94118, Tel. (415) 750-7277, FAX 
(415) 750-7090]. Information, registration forms, and a 
call for papers will be included in the Spring 1992 issue 
of the AMU Newsletter. Additional information about 
the meeting and membership applications can be ob- 
tained by contacting: 

Dr. Robert C. Bullock, President 
American Malacological Union 
Department of Zoology 
Biological Sciences Building 
University of Rhode Island 
Kingston, RI 02881 
Tel. (401) 792-2372 
FAX (401)792-4256 



WESTERN SOCIETY OF MALACOLOGISTS 

The Twenty-fifth Annual Meeting of the Western So- 
ciety of Malacologists will be held at the Asilomar Con- 
ference Grounds on Monterey Bay, Pacific Grove, Cal- 
ifornia, from June 30 to July 3, 1992. In addition to 
contributed papers on marine, freshwater and terrestrial 
mollusks, both living and fossil, the agenda includes two 
special symposia, on Opisthobranchs, and on Cocos Is- 
land, Costa Rica. A shell auction, reprint sale, and ban- 
quet round out the program. For further information 
contact WSM President David K. Mulliner (5283 Vickie 
Drive, San Diego, California 92109; (619) 488-2701) or 
WSM Treasurer Henry W. Chane\ (Museum of Natural 
History, 2559 Puesta del Sol Road, Santa Barbara, Cal- 
ifornia 93105; (805) 682-4711, ext. 334; fax (805) 569- 
3170). Deadline for receipt of abstracts is May 15, 1992. 



October 11-14, 1992 — Symposium on Freshwater Mus- 
sel Conservation and Management. Embassy Suites Ho- 
tel, St. Louis, MO. Sessions devoted to regulations, com- 
mercial harvest, conservation, sampling methodology, 
data management, and environmental awareness. For 
info: Kurt Welkc, Wisconsin Dept. of Natural Resources, 
11 1 W. Dunn St., Prairie du Chien, Wl 53821, 608/326- 
0233. 



THE NAUTILUS 106(2):43-49, 1992 



Page 43 



Two New Species of Cancellariidae 
(Gastropoda: Neogastropoda) from Brazil 



M. G. Harasewych 
Richard E. Petit 

Department of Invertebrate Zoology 
National Museum of Natural History 
Smithsonian Institution 
Washington, DC 20560 USA 



Andre Verhecken 

Scientific Associate 
Malacology Section 
Koninklijk Belgisch Instituut 
voor Natuurwetenschappen 
B- 1040 Brussels, BELGIUM 



ABSTRACT 

Two new species of cancellariids are described from shallow 
waters of the Brazilian Province. Each has a sister species in 
the Caribbean Province. Tritonoharpa leali, n. sp. differs from 
T. lanceolala in having a more inflated and thinner shell, in 
which the axial costae are more pronounced than the spiral 
cords. Cancellaria petuchi ii. sp differs from C. reticulata in 
having a thick, w hite callus on the columellar side of aperture, 
a broad, rounded shoulder, and a proportionalK longer aperture 
and larger protoconch than C. reticulata. CaJicclUiria petuclii 
lacks strongU cancellated sculpture on the bod\ whorl and the 
bifid posteriormost columellar fold of C. reticulata 

Key words: Cancellariidae; Tritonoharpa; Cancellaria. new 
species; Brazil. 



INTRODUCTION 

The nioiluscan fauna (if Brazil has been regarded bv 
mo.st authors (e.g., Abbott, 1974; Rios, 1970, 1975, 1985) 
as comprising the southernmost component of the Ca- 
ribbean Province. Several workers, among them Ekman 
(1953), Work (1969) and Rios (1970), had commented 
on an endemic component of the Brazilian molluscan 
fauna, while others (Briggs, 1974; Coelho & Koening, 
1972; Coelho & Ramos, 1972) had suggested the existence 
of a Brazilian Subprovince or Province. More recentK , 
Petuch (1988:166) delimited the region extending from 
the mouth of the Amazon River southward into the Mar 
de Plata as the Brazilian Province and listed a number 
of endemic molluscan indicator species. Other authors 
(e.g., Yokes, 1990) have begun to distinguish between 
Brazilian species and their often closely related Carib- 
bean counterparts. 

The cancellariid fauna of Brazil is poorK known. Lange 
de Morretes (1949, 1953) did not include any member 
of this family in his catalogs of Brazilian mollusks, while 
Abbott (1974:246) and Rios (1970:111, 1975:127, 1985: 
126) listed only Cancellaria reticulata Linne, 1767 and 
Tritonoharpa lanceolata (Menke, 1828) (as Cohibraria 
lanceolata) as occurring in the Brazilian fauna. Ver- 
hecken (1991) underscored the paucity of records for this 



family in the fauna of the subequatorial western Atlantic, 
and described two bathyal cancellariids from off south- 
eastern Brazil. 

A large series of specimens of an undescribed Can- 
cellaria as well as two specimens of a new species of 
Tritonoharpa were recently made available to us from 
commercial and institutional sources. We take pleasure 
in naming these new species after the persons who first 
brought them to our attention. 



ABBREVIATIONS USED IN TEXT 

KBIN — Koninklijk Belgisch Instituut voor Natuurweten- 
schappen, Brussels 

MNHN — Museum national d'Histoire naturelle, Paris 

MORG — Museu Oceanografico da Funda^ao Universi- 
dade do Rio Grande, Rio Grande, Brazil. 

UFJF — Universidade Federal de Juiz de Fora, Minas 
Gerais 

USNM — National Museum of Natural History, Smith- 
sonian Institution, Washington, DC 

SYSTEMATICS 

Family Cancellariidae Forbes & Hanley, 1851 
Subfamily Plesiotritoninae Beu & Maxwell, 1987 

Genus Tritonoharpa Dall, 1908 

Beu and Maxwell (1987) reviewed the genus Tritono- 
harpa, enumerating the following diagnostic characters: 
elongate-ovate to biconic shells with spire half or more 
the shell length; weakly to moderately convex teleoconch 
whorls; prominent non-collabral varices on early as well 
as later whorls; well developed columellar collar; colu- 
mellar plait either single and weak or absent; radula 
absent. 

Recent western Atlantic species of Tritonoharpa in- 
clude the Carolinian and Caribbean T. lanceolata 
(Menke, 1828); T. cubapatriae (Sarasua, 1975), based on 
a unique holotype from off Havana, Cuba; T. baijeri 
(Petuch, 1987) from the Caribbean coast of Colombia; 



Page 44 



THE NAUTILUS, Vol. 106, No. 2 




M. G. Harasewvch et al., 1992 



Page 45 




Figures 5-6. Tritonoharpa leali new species. 5. Protoconch of holotype, scale bar = 500 ^m. 6. Surface sculpture of holotype, 
scale bar = 200 ^m. Figures 7-8. Tritonoharpa lanceolata (Menke, 1828). 7. Protoconch of specimen in figure 3, scale bar = 
500 ^m. 8. Surface sculpture of specimen in figure 3, scale bar = 200 nm. 



and the new species described below. Minihraria mon- 
roei (McGinty, 1962) has been referred to the Muricidae 
on the basis of the morphology of its varices and pro- 
toconch (Beu & Maxwell, 1987:56). 

Tritonoharpa leali new species 
(figures 1, 2, 5, 6) 

Description: Shell (figures 1-2) to 18.4 mm, thin, tall, 
narrow. Spire high (spire angle 31-35°), stepped, com- 
prising about % of total shell length. Protoconch paucis- 



piral, of 2y4 inflated, weakly shouldered whorls (figure 
5). First l'/2 whorls nearly planispiral, descending rapidly 
thereafter. Protoconch coaxial with, or deviated from, 
teleoconch axis by up to 5° Transition to teleoconch 
abrupt, marked by flared protoconch lip and onset of 
pronounced spiral sculpture. Teleoconch with up to 5% 
slightly inflated, weakly shouldered whorls. Suture deep- 
ly impressed, obscured by axial costae of succeeding 
whorl. Spiral sculpture of strong, low, equally spaced 
cords, 20-26 on bod\' whorl, 12-15 on penultimate whorl; 
cords, 0-3, weak or absent on siphonal canal. Axial sculp- 



Figures 1-2. Tritonoharpa leali new species. 1. Holotype, MORG 286.59, Davis Bank, southeastern Brazil, 20°40'S,34°41'W, in 
60 m, Marion-Dufresne MD55 sta DC40, 7.5 x. 2. Paratype 1, USNM 860521, off Itaparica Island. Bahia, Brazil, trawled in 20 
m, 5.0x. Figure 3. Tritonoharpa lanceolata (Menke, 1828). USNM 798073. Off St. Thomas, Virgin Islands, 5.0x. Figure 4. 
Tritonoharpa haijeri (Petuch. 1987), Holotype, USNM 859853, Cabo La Vela. Goajira Peninsula, Colombia, in 35 m, 5.0 x. 



Page 46 



THE NAUTILUS, Vol. 106, No. 2 




Figures 9-10. Camcllaria pcluchi new species. *>. Ilolotvpe, MORG 286(i0, 2.0 x 10. Paralvpe 4. USNM 859409, Off Vitoria, 
Espirito Santo, Brazil, trawled on sand bottom in 40 ni, 2 Ox Figure 11. Caiicellaria adclac Pilsbrs, 1940, USNM 508663, Little 
Duck Key, Florida, 1 5x. Figure 12. CanccUaria rclirulata (Linne, 1767), LISNM 811479, 30 mi. ENE of Eau Gallie, Florida 
[28''17'N,'80°01'W], in 34-41 fms (62-75 m), K/V Silver Bav Sta 2010, 2.0x. 



lure of pronounced varices and narrow, opistliocline cos- 
tae. Varices high, non-collabral, narrow, rounded aba- 
pcrturally, undercut adaperturally, 180-200° apart on 
early whorls, up to 238° apart on later whorls. Costae (7- 
22 between varices, 15-33 per whorl) offset from varices 
by 5-10°, producing cancellate sculpture ar their inter- 
section with spiral cords. Aperture narrow, elliptical, with 



well defined posterior notch. Inner lip smooth, \\ith col- 
uiuellar collar reflected over, but not adherent to, fascicle 
and p.seudouinbilicus. Outer lip with up to 8 pairs of 
denticles confined to ilaretl region beyond varix. Siphonal 
canal short, dorsally reflected, partially covered abaxi- 
ally. Rase color khaki to yellowish tan, with darker brown 
maculations anterior to suture and along varices. 



M. G. Harasewych et ai, 1992 



Page 47 



Table 1. Shell measurements of Canccllaria reticulata and C. petuchi. All measurements in mm (N = 10). For C. reticulata, the 
10 specimens were selected from throughout the species range, while for C. petuchi. data is based on specimens from a single lot 
(paratypes 4-13). 



Shell length (SL) 

Aperture length (AL) 

(AL/SL) 

No. whorls, teleoconch 

No. whorls, protoconch 

Diameter, protoconch 

Diameter no. whorls, protoconch 



c 


re tic 


ulat 


a 


C. 


petuchi 




X 






(J 


X 




a 


39.67 






6.21 


33.68 




1-98 


25.64 






3.99 


23.64 




1.33 


0.65 






0.03 


0.70 




0.01 


6.34 






1.76 


5.51 




0.20 


2.67 






0.10 


2.74 




0.06 


1.23 






0.06 


1.60 




0.05 


0.46 






0.02 


0.59 




02 



Type locality: Davis Bank, southeastern Brazil, 
26°40'S,34°41'W, in 60 m, Marion-Dufresne MD.55 sta. 
DC40. May 1987. 

Material examined: Holotype, MORG 286.59, 9.22 mm; 
Paratype 1, USN'M 860521, off Itaparica Island, Bahia, 
Brazil, trawled in 20 m, 18.44 mm. 

Distribution: This species is known from two records, 
both off the state of Bahia, Brazil, in depths of from 20 
to 60 m. 

Etymology: This species honors Dr. Jose H. Leal, in 
recognition of his contributions to the study of Brazilian 
mollusks. 

Comparative remarks: Tritonoharpa leali most closely 
resembles T. lanceolata, from which it differs in having 
a somewhat more inflated and thinner shell, in which 
the axial costae are more pronounced than the spiral 
cords (compare figures 5 and 7). Tritonoharpa haijeri is 
even broader and thinner than T. leali, and is the only 
western Atlantic Tritonoharpa in which the aperture 
length exceeds half the shell length. 



Subfamily Canceilariinae Forbes & Hanley, 
Genus Cancellaria Lamarck, 1799 



1851 



The genus Cancellaria is characterized by the presence 
of a thick, ovate to biconic shell with strongly cancellate 
sculpture, prosocline outer lip, a weak stromboid notch, 
short but distinct siphonal canal, prominent siphonal fas- 
cicle, lirate outer lip, and columella with two to three 
strong folds, the posteriormost strongest and usually bifid. 
Jung and Petit (1990:100) reduce Pijniclia Olsson, 19.32 
to subgeneric status within Cancellaria, and distinguish 
it from the nominotypical subgenus on the basis of its 
pyriform shape with reduced or absent sculpture on body 
whorl and a columella with two strong, non-bifid folds. 

Cancellaria petuchi new species 
(figures 9, 10, 14-16) 

Description: Shell (figures 9-10) to 37 mm, heavy bi- 
conic, pseudoumbilicate. Spire high (spire angle 62-70°), 
conical, comprising about % of total shell length. Pro- 
toconch paucispiral, of 2'/2 low, inflated, glassy whorls 



(figure 14). Protoconch may be co-axial with, or deviated 
from teleoconch axis by up to 15°. Teleoconch with up 
to 6% strongly convex whorls. Suture deeply impressed. 
Spiral sculpture of 18-24 major cords on body whorl, 6- 
10 on siphonal canal, 6-8 on penultimate whorl. Axial 
sculpture of 20-38 prosocline ribs that form strongly 
cancellate sculpture at intersection with spiral cords, es- 
pecially on first four teleoconch whorls. Thereafter, axial 
ribs become broader, less pronounced, may disappear 
entirely, especialK' below the shoulder. Aperture large, 
broad, hemi-elliptical, deflected from coiling axis by 12- 
18° Outer lip with shallow indentation posterior to junc- 
tion with siphonal canal and 10-12 strong lirae, slightly 
recessed, either diminishing Vi whorl into aperture, or 
more usually, disappearing and reappearing V4 whorl 
within the aperture. Inner lip with 2 columellar and 1 
siphonal folds. Posteriormost columellar fold largest, 
overlying siphonal fasciole. All folds with single, sharp 
keel. Siphonal canal short, deflected dorsally and abax- 
ially. Base color white to light tan, with spiral bands 
ginger to dark brown interrupted by narrow axial bands 
of base color, especially on early whorls. Some specimens 
pure white. Aperture with white overglaze forming thick 
callus along columella. 

Type locality: N.E. of Vitoria, Espirito Santo, Brazil, 
19°35'S, 39°42'W, in 15 m. South Eastern Brazil N.O. 
Marion-Dufresne MD55 sta. DC87, May 1987. 

Material examined: Holotype, MORG 28660, 28. 16 mm; 
Paratypes 1-3, MNHN, N.E. of Vitoria, Espirito Santo, 
Brazil, 19°34'S,39°34'W, in 34 m, N.O. Marion-Dufresne 
MD55 sta. CB90, May 1987, 32.52 mm, 11.01 mm, 8.06 
mm; Paratypes 4-23, USNM 859409, Paratypes 24-36, 
Petit collection, Paratypes 37-38, KBIN, Paratypes 39- 
52, Verhecken collection, all from Off Vitoria, Espirito 
Santo, Brazil, trawled on sand bottom in 40 m, 29.26- 
36.56 mm. Paratypes 53-.55, Petit collection, Mangue da 
Olaria, Guarapari, Espirito Santo, Brazil, in sand, minus 
tide. Paratypes 56-57, UFJF, Niteroi, Rio de Janeiro, 
Brazil. Paratype 58, MORG 19472, off Salin6polis, Pard, 
Brazil, in 36 m. Paratypes 59-62, MORG 23307, Coroa 
Vermeiha, Abrolhos Islands, Bahia, Brazil, in 1 m. 

Distribution: This species ranges from off Salinopolis, 
Para, to off Niteroi, Rio de Janeiro, Brazil. It occurs from 
intertidal depths to 40 m. 



Page 48 



THE NAUTILUS, Vol. 106, No. 2 




Figure 13. Cancellaria reticulata (Linne, 1767). Protoconch, scale bar = 500 ^m. Figure 14-16. Cancellaria petuchi new- 
species. 14. Protoconch of paratype, same data as fig. 10. Scale bar = 500 iim. 15. Frontal and 16. Lateral view of distal ends of 
rachidian teeth. Scale bars = 5.0 nm (fig. 15), 3.0 ^m (fig. 16). 



Etymology: This species honors Dr. Edward J. Petuch, 
as a tribute to his contributions to our knowledge of the 
inoUuscan fauna of Brazil, and the Neogene zoogeog- 
raphy of the western Atlantic faunas. 

Comparative remarks: Cancellaria pettichi differs from 
its geographically pro.xiniate congener C reticulata in 
having reduced or absent cancellate sculpture on the 
body whorl and lacking a bifid posteriormost columellar 
fold. Cancellaria petuchi has a thick, white callus on the 
columellar side of the a|)erture and a broad, rounded 
shoulder, features lacking in C. reticulata. Cancellaria 
petuchi also has a proportionally longer aperture and 
larger protoconch than C. reticulata (table 1). Radular 
morphology of Cancellaria petuchi (figures 15, 16) agrees 
in all essential features with that of C. reticulata (Har- 
asewych & Petit, 1982:figures 14, 15 — these figures show 
the ventral surfaces of the distal tips of two radular teeth). 
Of the western Atlantic species of Cancellaria, C. pe- 
tuchi most strongly resembles C. adelae Pilsbry, 1940 



(figure 11), a species endemic to the Florida Keys, with 
which it shares its predominantly smooth body whorl 
sculpture and thick parietal overglaze. Cancellaria ade- 
lae however, has a strongK bifid columellar fold, and 
indeed, may have an additional fold along the posterior 
portion of columella. 

ACKNOWLEDGMENTS 

We thank Dr. Philippe Bouchet (MNHN), Dr. Edmund 
Gittenberger, (Nationaal Natuurhistorisch Museum, Lei- 
den), Robert Moolenbeek (Instituut voor Taxonomische 
Zoologie, Amsterdam), Prof. Maury Pinto de Oliveira 
(UFJF), Prof. Elesar C. Rios (MORG), and Dr. Edward 
J. Petuch, (Florida .Atlantic I'niversity, USA) for making 
specimens available for study. Drs. Edward J. Petuch 
and Jose H. Leal critically read drafts of the manuscript 
and provided some of the references on the biogeography 
of the Brazilian fauna. Ms. Susanne Braden provided the 
SEM micrographs. 



M. G. Harasewych et a/., 1992 



Page 49 



LITERATURE CITED 

Abbott, R T. 1974, American seashells, second edition. Van 
Nostrand Reinhold, Co., New York. 663 p. 

Beu, A. G. and P A Ma.wveli. 1987 A revision of tossij and 
living Gastr<ipods related to Plesiotritun Fisciier, 1884 
(Family Cancellariidae, SubfamiK Plesiotritoninae n. 
subfam.) with an appendix: Genera of Buccinidaee Pisa- 
niinae related to Colubraria Schumacher, 1817. New Zea- 
land Geological Survey Paleontological Bulletin 54.1-140, 
pis. 1-30. 

Briggs, J. C. 1974. Marine Zoogeography. McGrav\-Hill, New 
York. 475 p. 

Coelho, P. A. and M L Koening. 1972. .\ distribui9ao dos 
crustaceos pertencentes as ordens Stoniatopoda, Tanai- 
dacea e Isopoda no Norte e Nordeste do Brasil, Trabalhos 
Oceanograficos da Universidade Federal de Pernambiico 
13:245-260. 

Coelho, P. A. and M A. Ramos. 1972. A constitu9ao e dis- 
tribui(jao da fauna de decapodos crustaceos do litoral leste 
da America do Sul entre as latitudes de 5°N e 39°S. Tra- 
balhos Oceanograficos da Universidade Federal de Per- 
nambuco 13:133-236, 

Ekman, S. 1953. Zoogeography of the Sea. Sidgwick and 
Jackson, London .\iv -I- 417 p. 

Harasewych, M. G. and R. E. Petit. 1982 Notes on the mor- 
phology of Cancellaria reticulata (Gastropoda: Cancel- 
lariidae). The Nautilus 96(3):104-113. 

Jung, P. and R. E. Petit. 1990. Neogene Paleontology in the 
northern Dominican Republic, 10 The family Cancellar- 
iidae (Mollusca: Gastropoda), Bulletins of American Pa- 
leontology 98(334):87-144, 



Lange de Morretes, F. 1949. Ensaio de catalogo dos moluscos 

do Brasil Arquivos do museu Paranaense 7(1):5-216. 
Lange de Morretes, F, 1953. Adenda e corrigenda au ensaio 

de catalogo dos moluscos do Brasil, .Arquivos do Museu 

Paranaense 10(l):37-76, 
McGinty, T, L, 1962. Caribbean marine shells. The Nautilus 

76(2):39-44, pl.3. 
Petuch, E. J. 1987. New Caribbean Molluscan Faunas. The 

Coastal Education & Research Foundation, Charlottes- 
ville. 1-154, A1-A4. 
Petuch, E. J 1988. Neogene history of tropical American 

mollusks. The Coastal Education & Research Foundation, 

Charlottesville. 217 p. 
Pilsbrv, H. A. 1940. A new race of Cancellaria from Florida. 

The Nautilus 54(2):54. 
Rios, E. C. 1970. Coastal Brazilian Seashells. Museu Ocean- 

ografico, Rio Grande. 255 p., 60 pis. 
Rios, E. C. 1975. Brazilian Marine Mollusks Iconography. 

Museu Oceanografico, Rio Grande. 331 p., 91 pis 
Rios, E. C. 1985. Seashells of Brazil. Museu Oceanografico, 

Rio Grande. 328 p., 102 pis. 
X'erhecken, A 1991. Description of two new speciesof bathy- 

al Cancellariidae (Mollusca, Gastropoda) from off Brazil. 

Bulletin Museum national d Histoire naturelle, Paris. Se- 
ries 4, 12, Section A (3-4):547-553. 
\'okes, E. H. 1990. Two new species of Chicoreus subgenus 

Stratus (Gastropoda: Muricidae) from northeastern Brazil. 

The Nautilus 103(4):124-130. 
Work, R. C, 1969, Systematics, ecology, and distribution of 

the mollusks of Los Roques, Venezuela. Bulletin of Marine 

Science 19(3):614-711. 



THE NAUTILUS 106(2):50-54, 1992 



Page 50 



New Species of Solariella (Gastropoda: Trochidae) from the 
Western Atlantic Ocean 



James F. Quinn, Jr. 

Florida Marine Research Institute 
Department of Natural Uesiiurces 
100 Eighth Avenue, SE 
St. Petersburg, FL 33701-5095, USA 



ABSTRACT 

Three new species of the genus Sulariclhi Wood, 1842, are 
described. Solariella qtiadricincta is known only from the con- 
tinental shelf off northeastern Venezuela in deptlis of 26-8(i m, 
and S. staminea is known only from the Davis Seamount off 
southeastern Brazil in 60 m; shells of both species are very 
similar to those of S. carvalhoi Lopes and Sa Cardoso, 1958. 
Solariella cristata is known from the upper continental slope 
of Isla Cancun, Yucatan, Mexico, and off Key Largo, Florida 
Keys, in depths of 155-256 m, and off St. Vincent, Lesser 
Antilles in 165-201 m. 

Key words: Trochidae; Solariellinae; Solariella; systematics; 
new species. 



INTRODUCTION 

Two recent monographs of western Atlantic Trochidae 
present accounts of the faunas of restricted geographical 
areas: the Straits of Florida (Quinn, 1979) and tlie Gulf 
of Mexico (Quinn, in press). In the process of examining 
specimens for those reports, several new species from 
other areas of the western Atlantic Ocean were discov- 
ered. This paper presents descriptions of three new spe- 
cies of Solariella Wood, 1842. These species are included 
in Solariella because of the similarities of their shells to 
tho.se of species such as S. lacunella (Dall, 1881); how- 
ever, because radular characters are important (Herbert, 
1987) and animals of the three species were unavailable 
for study, such a.ssignment of these species is tentative 
at present. 

Institutional abbreviations used in this paper are as 
follows: MNIIN (Museum National d'llistoire Naturelle, 
Paris, France); MORG (Museu Oceanographico da Fun- 
da(,'ao Universidade do Rio Grande, Rio Grande, RS, 
Brazil); UMML (Rosenstiel School of Marine and At- 
mospheric Science, University of Miami, Miami, Flori- 
da); USNM (National Museum of Natural History, Smith- 
sonian Institution, Washington, DC). 



SYSTEMATICS 

Genus Solariella Wood, 1842 

Type species (monotvpv): Solariella maculata Wood, 
1842. 

Solariella quadricincta new species 
(figures 1-4) 

Material examined: 1 fragment, UMML 30.6695 (para- 
type); JOHN ELLIOTT >ILLSBURY Station P-727, 
10°20'N, 65°02' W, 64 m; 10-ft otter trawl; 21 July 1968.— 
1 specimen, L'MML uncatalogued [examined and pho- 
tographed in 1975 (figs. 3, 4), but a recent attempt to 
relocate this specimen was unsuccessful, and it is pre- 
sumed lost]; JOHN ELLIOTT PILLSBURY Station P- 
721, ir06.5'N, 64''22.5'W, 26-27 m; 10-ft otter trawl; 
21 Julv 1968. — 1 specimen, USNM 859437 (holotype); 1 
specimen, UMML 30.6528 (paratype); JOHN ELLIOTT 
PILLSBURY Station P-718, 1 1°22.5'N, 64°08.6'W, 60 m; 
10-ft otter trawl; 20 Julv 1968.— 1 specimen, UMML 
30.6376 (paratype); JOHN ELLIOTT PILLSBURY Sta- 
tion P-705, 10°45'N, 62°00'W, 77-86 m; 10-ft otter trawl; 
18 July 1968. 

Description: Shell of moderate size for genus, attaining 
6.55 mm height, 7.15 mm width, umbilicate, ivory with 
light orange-brown spots and flammules, nacreous under 
thin outer porcelaneous layer. Protoconch 300-325 /um 
maximum diameter, of about 1 whorl. Teleoconch whorls 
5.5, tubular, shouldered; first 3 whorls with 3 strong, 
subequal spiral cords forming whorl periphery, adapical 
cord forming whorl shoulder; suKsequent whorls with 
subsutural spiral cord forming narrow channel with su- 
ture; last whorl with fourth primary spiral cord, subequal 
to other peripheral spiral cords and coincident with su- 
ture on previous whorls; 1-2 weak spiral cords sometimes 
occurring between primary spiral cords; fine spiral threads 
overK ing interspaces and primary spiral cords on last 3 
whorls. Axial sculpture consisting of rather strong riblets 
on whorls 2 and 3, fading in strength to numerous col- 
labral threads in interspaces of spiral cords on subsequent 



J. F. Quinn, Jr., 1992 



Page 51 




r- 



K 






p^ 



^ 



r/ 



V, 



^ 






%^^^JI 



Figures 1-t. SolaricUa quadricincta new species. 1, 2. .^pertural and basal views of holotype, USNM 8.594.37, height 6.40 mm, 
widtli 6.40 mm, from oft Isia cle Margarita, Venezuela, 11°22..5'N, 64°08.6'W, 60 m. 3, 4. Apertural and basal views of specimen 
(lost; measurements imknowii), UMML .30.66.'59, from off Isla de Margarita, Venezuela, 11°06..5'N, 64°22..5'W, 26-27 m. Figures 
5, 6. SolaricUa carvalhoi L<ipes and Sa C^ardoso, 19.58. FSBC 1 39514, height 7.9 mm, width 8.2 mm, from Cagarras Island, Rio 
de Janeiro, Brazil, 32 m. 



whorls; axial threads form lamelliform beads on subsu- 
tiiral spiral cord, w eakK beading shoulder spiral cord on 
whorls 3-5. Subsutural shelf rather wide, flat, sloping 
abapically from subsutural spiral cord to shoulder spiral 
cord; fine spiral threads appearing on third whorl near 
shoulder, progressively covering shelf surface on subse- 
quent whorls; 1-2 stronger spiral threads appearing on 
last whorl. Base convex, with 6-7 strong spiral cords; 
cords and interspaces with fine spiral threads; interspaces 
with fine collabral threads. Umbilicus wide, about 40- 
45% maximum shell widtli, funnel-shaped; v\ alls convex, 
with 6 strongly beaded spiral cords. Shell ground color 
ivory, with regularly spaced, spirally elongate light or- 
ange-brown spots on major spiral cords, with occasional 
flammules of same color on subsutural shelf. Operculum 
thin, corneous, multispiral 

Type locality: Off Isla de Margarita, Venezuela, 
11°22.5'N, 64°08.6'W, 60 m. 

Remarks: Shells ot SolaricUa quadricincta closely re- 
semble those of S. staminea. new species, and S. car- 
valhoi Lopes and Sa Cardoso, 1958, from Brazil (figs. 5, 
6). Shells of all three species have four strong, subequal 



primary spiral cords on the last whorl, of which the 
shoulder spiral cord is the most distinctly beaded; have 
fine spiral threads on suprabasal whorl surface; and have 
flat subsutural shelves. Shells of S. quadricincta differ 
from those of S. starnirica by being slightly narrower 
(height;width ratios = 0.92-1.01 and 0.79-0.90, respec- 
tively); by having a channeled suture; by having crisp 
rather than flattened axial threads; by having fewer, nar- 
rower, rounded basal spiral cords; by having a narrow 
and more strongly beaded circumumbilical cord; by hav- 
ing more numerous, more strongly beaded intra-mnbil- 
ical spiral cords; and by having a strong, distinct color 
pattern. Shells of S. quadricincta differ from those of S. 
carvalhoi by being smaller at similar whorl number; by 
ha\ ing canaliculate sutures; by having more steeply slop- 
ing subsutural shelves; by having finer, crisper, more 
closely spaced axial threads that are present in the in- 
terspaces of all spiral cords; by having a more finely 
beaded shoulder cord; by having wider interspaces be- 
tween the basal spiral cords; and by having more strongly 
beaded intraumbilical spiral cords. SolaricUa quadri- 
cincta is only known from the continental shelf off north- 
eastern Venezuela in depths of 26-86 m. 



Page 52 



THE NAUTILUS, Vol. 106, No. 2 




"\ 







t 



10 



Figures 7-10. Solariclla cristata new species. 7, 8. Holotype, IISNM 859421, height 9 1 mm. width 9 4.5 mm, from SE of St. 
Vincent, Lesser Antilles, 13°11.2'N, 6r05.1'W, 16.5-201 m, 9, 10. Paratype, UMML 30.5669, height SI mm, width 8.0 mm, from 
off Isla Mujeres, Yucatan, Mexico, 21°07'N, 86°21'W, 155-205 m. 



Solariella cristata new species 
(figures 7-10) 

Material examined: 1 specimen, UMML 30.5669 (para- 
type); JOHN ELLIOTT PILLSBURY Station P-598, 
21°07'N, 86°21.0'W, 155-205 m; 10-ft otter trawl; 15 
March 1968.— 1 specimen, UMML 30.7433 (paratype); 
GERDA Station G-6I, 25°14'N. 80°02'W, 256 m;' 6-ft 
otter trawl; 29 August 1962.— 1 specimen, L'SNM 859421 
(holotype); JOHN ELLIOTT PILLSBURY Station P- 
874, 13''11.2'N, 61°05.1'W, 165-201 ni; 5-ft Blake trawl; 
6 July 1969. 

Description: Shell rather large for genus, attaining 9.1 
mm height, 9.45 mm width, umbilicate, ivory vvith light 
brown spots and llannnulcs, nacreous under thin outer 
porcelaneous layer. Protoconch 300-320 ^lm maximum 
diameter, of about 1 whorl. Teleoconch whorls 6.1, tu- 
bular, shouldered; first 3 whorls with 4-5 spiral cords, 
increasing to 14 on body whorl; abapical cord on whorls 
3-5 strongest, forming peripheral carina; cords on last 
whorl becoming subequal to peripheral cord, giving whorl 



more evenly rounded appearance. Axial sculpture ap- 
pearing on whorl 2, consisting of rather strong, low, flat- 
tened folds on adapical half to two-thirds of whorls 2- 
4, becoming narrower, roimded, more crowded, and ex- 
tending to level of suture on subsequent whorls; axial 
folds forming strong, rounded beads on spiral cords, beads 
strongest on adapical part ol whorl. Subsutural shelf nar- 
row, sloping adapically from suture to shoulder spiral 
cord; single strong spiral cord bisecting shelf. Base weakly 
convex, with 7-10 strong, smooth spiral cords. Umbilicus 
wide, about 30% maximum shell width, funnel-shaped; 
walls convex, with 4-6 strong, strongly beaded spiral 
cords. Shell ground color ivory, with scattered light brown 
spots appearing on fourth whorl, becoming more nu- 
merous and forming irregular axial flammules on last 
whorl. 

Type locality: SE of St. Vincent, Lesser Antilles, 
I3°11.2'N, 6r°05.1'W, 165-201 m. 

Remarks: The peripherally carinate, rather strongly ax- 
ially sculpted shells of Solariella cristata most closely 



J. F. Quinn, Jr., 1992 



Page 53 





Figure 1 1-12. Soldriclla staminea new species Apertural and basal views of holotype, MORG 26530, height 3.8 mm, width 4 8 
mm, from Davis Seamount, Brazil, 20°40'S, 34°41'W, 60 m. 



resemble shells of S. cincta (Pliilippi, 1836) (see Fretter 
& Graham, 1977:46-48, figs. 31, 32) from the north- 
eastern Atlantic Ocean. However, shells of S. cristata are 
larger, have a narrower subsutural shelf, and have a 
distinct color pattern, whereas those of S cincta are 
uniformly ivor\ . The only other western Atlantic species 
having shells with a strongly carinate periphery is S. 
patriae Carcelles, 1953, but shells of that species lack 
other spiral cords except the shoulder and circumbasal 
spiral cords, and have only three basal cords (see Rios, 
1985:23, pi. 10, fig. 97). The three widely separated lo- 
calities from which S. cristata has been collected (the 
Florida Keys, Yucatan, and the Lesser Antilles) suggest 
that the species is widespread, but rare, in the Caribbean 
Sea. 

Solariella staminea new species 
(figures 11, 12) 

Material examined: 1 specimen, MORG 26530 (holo- 
type); 3 specimens, MNHN uncatalogued (paratypes); 
MARION-DUFRESNE Cruise MD-55, Station DC-40, 
20°40'S, 34°41'W, 60 m; dredge; May 1987; P. Bouchet, 
J. H. Leal, and B. Metivier collectors. 

Description: Shell of moderate size for genus, attaining 
6.7 mm height, 7.45 mm width, umbilicate, white with 
few light yellow to orange-brown spots and streaks, na- 
creous under thin outer porcelaneous layer. Protoconch 
about 300 ^m ma.ximum diameter, of about one whorl. 
Teleoconch whorls 5.6, tubular, shouldered; spire whorls 
with 3 strong spiral cords, adapical one forming whorl 
shoulder; last whorl with fourth strong spiral cord, sub- 
equal to other spiral cords and coincident with suture on 
previous whorls; 1-2 additional, weaker, intercallary spi- 
ral threads sometimes present; fine spiral threads over- 
ly ing interspaces and lower 2-3 spiral cords on last 3 
whorls. Axial sculpture of rather strong riblets on whorl 
2, fading in strength to numerous, crowded, rather flat- 
tened collabral threads in interspaces of spiral cords on 
subsequent whorls; axial threads forming weak, rounded 



beads on shoulder spiral cord, finely beading other spiral 
cords. Subsutural shelf moderately wide, flat, sloping 
abapically from suture to shoulder spiral cord; fine spiral 
threads appearing on third whorl; one spiral cord ap- 
pearing near beginning of first whorl, but fading to ob- 
scure angulation or disappearing on second whorl; 1-2 
spiral cords appearing on third whorl, one near middle 
of shelf strongest. Base weakly convex, with 9-11 strong, 
flattened spiral cords; innermost cord strap-like and 
weakly beaded; interspaces with fine collabral threads, 
forming weak rugae on spiral cords on adaxial half of 
base. Umbilicus wide, about 35-45% maximum shell 
width, funnel-shaped; walls convex, with 4-5 beaded 
spiral cords. Shell ground color white, with few, widely 
spaced light yellow to orange-brown spots, principally 
on shoulder cord, but occasionally extending abaxially 
as short streaks or flammules. 

Type locality: Davis Seaniount, off southeastern Brazil, 
20°40'S, 34°41'W, 60 m. 

Remarks: Shells of Solariella staminea are very similar 
to those of S. quadricincta and S. carvalhoi. Differences 
that distinguish shells of S. staminea from those of S. 
quadricincta are discussed in the Remarks section of the 
latter species. Shells of S. staminea differ from those of 
S. carvalhoi by being broader (height:width ratios = 
0.79-0.90 and' 0.93-0.98, respectively); by having a 
broader subsutural shelf with stronger spiral cords and 
axial threads; by having a shoulder spiral cord bearing 
rounded rather than spirally elongate beads; by having 
more numerous, flatter basal spiral cords; by having a 
circumumbilical spiral cord that is broad, flat, and weak- 
ly beaded rather than narrow, sharp, and strongly bead- 
ed; by having stronger, more weakly beaded intraum- 
bilical cords; and by having a very weak color pattern. 

ACKNOWLEDGMENTS 

I thank the late Gilbert L. Voss (Rosenstiel School of 
Marine and Atmospheric Science, University of Miami) 



Page 54 



THE NAUTILUS, Vol. 106, No. 2 



for access to the collections under liis care and for per- 
mission to d(>.scribe the new species in that material. The 
late Joseph Ilo.sewater and Richard S. Houbrick (both 
USNM) kindly allowed me to examine types and other 
material. Jose H. Leal brought the specimens of Solariella 
staminea to my attention and kindK provided the SEM 
micrographs of that species. Eliezer de C. Rios (MORG) 
generous!) provided comparative material of Solariella 
carvalhoi. Marjorie Myers, Llyn French, and Judy Leiby 
provided valuable word-processing and editorial assis- 
tance. Thomas H. Perkins and William G. Lyons (both 
Department of Natural Resources, Florida Marine Re- 
search Institute), and two anonymous reviewers provided 
valuable critiques of previous drafts of this paper. 

LITERATURE CITED 

C^arcelies, A. R. 19.53. Nuevasespeciesdegastropoclosmarinos 
de las republicas oriental del I'ruguay y Argentina Co- 
municaciones Zooiogicas del Museo de Historia Natural 
de Montevideo 4(70): 1-16. 

Dall, W. H. 1881. Reports on the results of dredging, under 
the supervision of Alexander Agassiz, in the Gulf of Mex- 
ico, and in the Caribbean Sea, 1877-79, by the United 
States Coast Survey steamer "Blake", Lieutenant-Com- 
mander C. D. Sigsbee, U.S.N., and Commander J l\. Bart- 



lett, U.S.N., commanding. W. Preliminary report on the 
Mollusca. Bulletin of the Museum of Comparative Zool- 
ogy, Har\ard L'niversity 9(2)::3-3-144, 

PVetter. \' and .\. (Jraham 1977. The prosobrancli molluscs 
of Britain and Denmark. Part 2 — Trochacea Journal of 
Molluscan Studies, Supplement 3:39-100. 

Herbert, D. G. 1987. Revision of the Solariellinae (Mollusca: 
Prosobranchia: Trochidae) in southern Africa. Annals of 
the Natal Museum 28(2):283-382. 

Lopes, H de S. and P de Sa Cardoso. 1958. Sobre un novo 
gastropodo brasileiro do genero "Solariella" Wood, 1842 
(Trochidae). Revista Brasileira de Biologia 18(l):59-64. 

Philippi, R .\ 18.3(1 Eniuneratio molluscorum Siciliae cum 
viventium tuni tellure fossilium, quae in itinere suo ob- 
servavit. Vol. 1. Berolini, Sinionis Schroppii et Soc, xiv -I- 
267 pp. 

Quinn, J. F., Jr. 1979. Biological results of the University of 
Miami Deep- Sea Expeditions. 130. The systematics and 
zoogeography of the gastropod family Trochidae collected 
in the Straits of Florida and its approaches. Malacologia 
19(l):l-62. 

Quinn, J. F.,Jr. In press. The Trochidae t)f the Ckilf of Mexico 
(Prosobranchia: Archaeogastropoda). Memoirs of the 
Hourglass Cruises. 

Rios, E. de C. 1985. Seashells of Brazil. Funda^ao Cidade do 
Rio Grande, Rio Grande, RS, Brazil, 328 -t- [103] p. 

Wood, S. V. 1842. A catalogue of shells from the Crag, .\nnals 
and Magazine of Natural History, Series 1, 9:527-544. 



THE NAUTILUS 106(2):55-59, 1992 



Page 55 



A New Fossil Land Snail of the Genus 
Hemitrochus from Bowden, Jamaica^ 



Glenn A. Goodfriend 

Geophysical Laboratory 
Carnegie Institution of Washington 
5251 Broad Branch Road, NW 
Washington, DC 20015 LISA, and 
Department of Environmental 

Sciences and Energy Research 
Weizmann Institute of Science 
76100 Rehovot, ISRAEL 



ABSTRACT 

A new species of Hemitrochus, H. bowdenensis, is described 
from a collection made in the Bowden Beds at Bowden, Ja- 
maica. The species differs from its nearest relative H. gramin- 
icola (the only modern Hemitrochus found in Jamaica) on the 
basis of shell sculpture, the siiape of the whorls, and umbilical 
width. Analysis of amino acid D/L ratios from one specimen 
indicates a late Holocene age for the sample and that therefore 
the material does not belong stratigraphically to the Bowden 
Beds (of early Pliocene age). The species is apparently extinct, 
probably as a result of either habitat destruction following 
European settlement or late Holocene climatic changes. 

Key words: land snail, fossil, new species, Jamaica, Hemitro- 
chus. Bowden. 



INTRODUCTION 

The Bowden Beds, located near Bowden, St. Thomas 
Parish, in southeastern Jamaica, have yielded an ex- 
tremely rich marine mollusk fauna (Woodring, 1925, 
1928) as well as a number of land snails, including a 
species of helicinid of the genus Lticidella (Simpson, 
1895), three species of poteriids (Simpson, 1895; Bartsch, 
1942; Morrison, 1955), and two species of camaenids of 
the genus Pleurodonte (Simpson, 1895; Kimball, 1947). 
Various other land snail species have been mentioned as 
having been collected from the Bowden Beds ("Thijsa- 
nophora", Opeas striata. Succinea latior, Simpson, 1895; 
"Stenogyra", "Melaniella", "Truncatella", Woodring, 
1928), but because of the fresh appearance of some of 
these shells, it has been suggested (Woodring, 1928) that 
these mav be recent material washed into the Bowden 



' Contribution No. 24, Department of Environmental Sci- 
ences and Energy Research, Weizmann Institute of Science 



sediments. The Bowden Beds are considered to be of 
early Pliocene age based on analysis of the foraminiferal 
fauna (assigned to the Globorotalia margaritae zone by 
Bolli and Bermudez (1965), which was placed in the early 
Pliocene by Bolli and Premoli Silva (1973)). Recent work 
on marine mollusks from Bowden has accepted this age 
assignment (Jung, 1989). 

E.xamination of the collections at the Academy of Nat- 
ural Sciences of Philadelphia (ANSP) has turned up a 
new species of land snail collected from the Bowden Beds 
and belonging to the family Helminthoglyptidae (= Xan- 
thonycidae sf nst/ Baker (1943) and Nordsieck (1987); = 
Fruticicolidae sensu Turner (1958)). This new species is 
described below and its relationships are discussed. 

Hemitrochus boivdenensis new species 

Description: Shell of average size for the genus {ca. 11- 
12 mm diameter), low-trochoidal (height % of diameter), 
moderately thin; spire weakly convex; shell periphery 
subangular, becoming nearly rounded at the lip; suture 
deeply impressed; base weakly convex, with the apex of 
the convexity shifting from a position in the middle of 
the base in the younger part of the shell to a position 
nearer to the umbilicus (than to the periphery) as the 
adult lip is approached; descent of the suture behind the 
lip unknown, since the upper part of the lip is not pre- 
served in the specimens; lip unreDected at periphery but 
gradually becomes reflected on the base in the direction 
of the umbilicus; bordering the umbilicus, the lip is 
strongly reflected, but only for a short distance, whereas 
in the middle of the base of the shell, the lip reflection 
starts earlier but is weaker; the basal insertion of the lip 
is drawn out, forming a rim on one side of the umbilicus 
which is reflected over a small part of the umbilicus; 
aperture round to weakly elliptical, with the width usu- 
ally slightly exceeding the height; protoconch 1.6-1.7 
whorls, smooth; the sculpture of later whorls consists of 



Page 56 



THE NAUTILUS, Vol. 106, No. 2 



Table 1. 


Measurements of the 


type material 


of Hemitrochus 


boudenensis. 










Specimen 


Life history 
stage 


Diameter 

(mm) 


Height 

(mm) 


Height/ 
diameter 


Whorl 
number 


Aperture 
width 

(mm) 


Aperture 
height 

(mm) 


.Aperture 

height/ 

width 


Holotype 
Paratype 
Paratype 


adult 

subadult 

?juveniie 


12.4' 

10.5' 

8.6 


8.3 
6.9 
5.5 


0.67 
0.66 
0.64 


4.5 
4.4 
4.0 


5.2- 
4.6 
3.9 


49^ 
4.6 

3.8 


0.942 
1 00 
0.97 



' Diameter is the appro.ximate adult diameter; a precise measurement is not possible since the lip is broken. 
^ Measured 0.1 whorl behind aperture, due to broken lip. 



low, broad ribs {ca. 12-14 on the penultimate V4 whorl) 
that are parallel to the growth lines and irregular, with 
each rib varying in both height and width along its length; 
the length of the ribs also varies, with some disappearing 
on the dorsum of the shell, and others continuing across 
the periphery and terminating just below the periphery; 
on the rest of base, sculpture consists only of irregular 
growth lines, which get stronger nearer the lip; color 
opaque white. Measurements of specimens are presented 
in Table 1. 

Material examined: The holotype (ANSP 75798) and 
two paratypes (ANSP 75799) are the only known spec- 
imens of this species. The holotype (figures 1-3) is an 
adult shell missing the upper lip and the end of the lip 
where it becomes the umbilical wall. The first paratype 
(figures 4, 5) is apparently a subadult specimen. The lip 
shows the reflection which characterizes the adult form 
but the lip is a little thinner than that of the holotype, 
suggesting that the shell is not fully mature. The dimen- 
sions of the shell should be representative of the adult 
dimensions, since further growth would result only in 
thickening of the shell. The upper lip of this specimen 
is broken as in the holotype but the umbilical end of the 
lip is intact. The second paratype has the last ca. 1 whorl 
missing, the remains of which can be seen only as a rim 
around the umbilicus. An additional ca. 'A of the upper 
part of the last whorl was originally present but was 
removed for amino acid analysis. 

Type locality, Mratigraphy, and collection information: 

The locality data provided with the material indicates 
that it was collected "Among oligocene fossils, Bowden, 
Jamaica", by Uselma C. Smith and S. L. Schumo. Bowden 
is a small village located on the east side of Port Morant 
Bay in southeastern St. Thomas Parish, Jamaica. No 
stratigraphic information is provided with the material. 
However, concerning the well-known Bowden fossil beds 
(later considered to be Miocene in age; Woodring, 1928), 
Woodring (1925:7) stated that "At the type locality [i.e., 
at Bowden] a bed of imj)erfectly consolidated gravel 
consisting of small pebbles and grains of igneous rocks, 
limestone, and other sorts of rock in a marly matrix crops 
out in the road-cut for a distance of several hundred feet 
along the bay. This bed, which is not more than 2 or 3 
feet thick, contains perfectly preserved fossils, and, so 
far as known, has furnished all the fossils collected at 
Bowden." Chubb (1958:28) indicates that at Bowden 



there is "one bed, some 2-3 feet thick lying a few feet 
above road-level, of unconsolidated gravel consisting of 
small pebbles and grains of igneous rocks in a marly 
matri.x. The bed is lenticular and runs for only about a 
couple of hundred yards, and it is extremely rich in well- 
preserved fossils. ..." The land snail material was among 
an extensive collection of marine mollusks made b\ Smith 
and Schumo and it seems almost certain that all this 
material was collected from the locality described by 
Woodring and by Chubb. In a visit to Bowden by the 
author in April, 1991, it was found that the beds con- 
taining marine mollusks run along the roadcut on the 
east side of the Bowden road, northward from the junc- 
tion of the road climbing up the hill from Bowden. This 
should be taken as the type locality for Hemitrochus 
bowdenensis. 

No date of collection is associated with the Schumo 
and Smith collection from Bowden, but the catalogue at 
the Academy of Natural Sciences of Philadelphia indi- 
cates that this material was presented to the museum on 
January 7, 1902 (G. Rosenberg, personal communica- 
tion), so it was collected some time before this date. 

Comparative remarks: The only other helminthoglyp- 
tid genus found in Jamaica is Dialeuca. The shells in this 
genus differ from Hemitrochus shells in several respects. 
The umbilicus (covered in adults) is very narrow and the 
basal lip inserts at a high angle to the columella (in 
Hemitrochus this insertion is at a low, near vertical angle); 
these features, together with a higher expansion rate of 
the last whorl, result in the aperture being considerably 
wider than high. The basal lip expands gradually and 
evenly across the base (this also occurs in some Bahamian 
Hemitrochus species) and the dorsal lip is reflected. The 
sculpture consists only of weak growth lines (this occurs 
also in some Bahamian Hemitrochus species). The ribbed 
sculpture characteristic of most species of Hemitrochus 
occurs also in Plagioptycha, a genus found in Hispaniola, 
the Bahamas, and the Lesser Antilles. But Plagioptijcha 
differs from Hemitrochus in having a high-angle inser- 
tion of the basal lip onto the columella (as in Dialeuca) 
associated with a relatively wide aperture; the shell is 
more depressed and a ridge is usually present on the 
inside of the lower lip on the left (umbilical) side. Thus 
the new species clearly belongs in Hemitrochus rather 
than in a related helminthogU ptid genus. 

Hemitrochus bowdenensis most closeH resembles the 
only other Hemitrochus species inhabiting Jamaica, H. 



G. A. Goodfriend, 1992 



Page 57 




Figures 1-3. Heniitrochus huwdencnsis new species, dorsal, apertural, and ventral views of holotype. 4, 5. Hernitrochus bow- 
denensis new species, apertural and ventral viev\s of paratype. 6-8. Hernitrochus graminicola (from Happy News, SE of Alexandria, 
St. Ann, Jamaica), dorsal, apertural, and ventral views. Scale line (in mm) in figure 2 refers to figures 1-5 and scale line in figure 
7 refers to figures 6-8. 



graminicola (C. B. Adams). H. graminicola (figures 6- 
8) differs in having a more weakly developed sculpture 
consisting of fine, regular ribs {ca. 30 on the penultimate 
'/4 whorl) which cross the base of the shell, disappearing 
only near the umbilicus. It also differs in having a round- 
ed periphery on the whole of the last whorl, a more 
inflated base, less impressed sutures, and a generally larg- 
er size (12-16 mm diameter). However, in the form of 
the lip reflection and in the shift of the apex of the basal 



convexity toward the umbilicus in the direction of the 
lip, H. boivdenensis resembles H. graminicola precisely. 
H. pseudogtjra (Torre) from Cuba closely resembles H. 
graminicola but is less close to H. boivdenensis in that 
its sculpture is both weaker and finer than that of H. 
graminicola. It should be emphasized that, although these 
are the species nearest phenotypically to H. boivdenensis, 
they are not very closely related to it — the character of 
the sculpture of H. boivdenensis is altogether different 



Page 58 



THE NAUTILUS, Vol. 106, No. 2 



Table 2. D/L amino acid ratios in paratype specimen of 
Hemitrochus howdcncnsis 



Amino acid 



D/L 



Alanine 

Alloisoleucine, isoleucine 

Proline 

Aspartic acid 

Methionine 

Glutamic acid 

Phen\ialanine 



0.12 

0.053 

0.33 

0.27 

0.24 

0.062 

014 



from that of any modern species. H. bowdenensis is not 
banded as other Hemitrochus species are. The lack of 
banding on the shells could possibly be the result of 
fading, but this seems unlikely for such young material. 
Hemitrochus has been considered a subgenus of Ce- 
polis (Pilsbry, 1939; Baker, 1943), but Turner (1958) 
raised it to generic status on the basis of anatomical 
characteristics. Altliough the earlier inclusion of Hemi- 
trochus within Cepolis was retained by Nordsieck (1987) 
and Vaught (1989), the taxonomy of Turner (1958) is 
followed here. 



AGE OF THE SAMPLE 

Although the Bowden Beds are considered to be of early 
Pliocene age, the possibility of inclusion of some modern 
material in the collections still exists, for example if ma- 
terial were collected from slumped sediments which could 
have incorporated modern terrestrial shells or from ex- 
posed fissure infills. Woodring (1928) considered that 
some of the land snail material that had been collected 
from the Bowden Beds may have been modern, stating 
that some of the material may represent "the remains 
of living snails that fell into openings in the ground and 
thus were collected with the fossil material" (p. 109). 
Because of this possibility, amino acid enantiomer/epi- 
mer analyses were carried out on a fragment of one of 
the H. bowdenensis shells as a check on its age (see 
Goodfriend, 1991, for analytical methods). Material of 
early Pliocene age would be expected to give D/L amino 
acid ratios near equilibrium (1.3 for D-alloisoleucine/L- 
isoleucine and 1.0 for other D/L amino acid ratios) and 
may be highly depleted in amino acid content. 

The analytical results (Table 2) indicate that the Hemi- 
trochus shell is relatively young. As would be expected, 
faster-raccmizing amino acids such as aspartic acid, pro- 
line, methionine, and phenylalanine (Goodfriend, 1991) 
show higher D/L ratios than the slower racemizing/ 
epimcrizing amino acids glutamic acid and isoleucine. 
A calibration of the rate of isoleucine epimerization in 
Pleurodonte during the Holocene at a site on the north 
coast of Jamaica (Goodfriend and Mittercr, 1988) leads 
to an age estimate based on the alloisoleucine/isoleucine 
(A/I) ratio of the Hemitrochus of 1700 year B.P. (as- 
suming an initial A/I value of 0.013, as is typical of 
modern land snail shells). This estimate is very approx- 



imate since there are a number of errors involved which 
compound to produce the total error: the error of mea- 
surement of the A/I ratio of the Hemitrochus sample 
(5-10%), possible differences in the epimerization rate 
between different genera (usually on the order of 10%), 
and, most importantly, the uncertainty of the rate dif- 
ference between Bowden and the north coast calibration 
site. For example, a 2° difference between the sites would 
lead to a 40% difference in the epimerization rates (equa- 
tion 3 in Goodfriend and Mitterer, 1988). Allowing a 
total uncertainty of ±50% would indicate a probable age 
between 800-2400 year B.P. Thus it is clear that the 
sample is of late Holocene age, and therefore represents 
material which does not belong to the Bowden Beds. A 
visit to the Bowden Bed type locality revealed that the 
moUusk-containing unit was in many cases covered by 
colluvial material, slumped down from the steep slope 
above. It seems likely that the H. bowdenensis material 
was collected from such a slumped deposit, perhaps at 
the same level as the mollusk beds. 

DISCUSSION 

This new species of Hemitrochus is not represented in 
modern collections from Jamaica and is therefore pre- 
sumably extinct. Whether this presumed extinction was 
the result of forest clearance subsequent to European 
settlement of Jamaica or occurred before this time, as a 
result of natural processes such as climatic change, cannot 
be ascertained without additional dated records of this 
species. Evidence of human induced local extirpations 
of land snail species exists for the north coast of Jamaica 
(Goodfriend and Mitterer, 1988). But climatic changes 
in the late Holocene have also been documented in Ja- 
maica (Goodfriend, 1987). 

The occurrence of this apparently extinct species in 
the recent fossil record of Jamaica is somewhat surprising 
in view of the fact that other Holocene and late Pleis- 
tocene deposits on the island contain only extant species 
(Goodfriend and Mitterer, 1988; Goodfriend, 1989). Re- 
cent extirpations of species in north-central Jamaica 
(Goodfriend, 1987) and the central north coast of Jamaica 
(Goodfriend and Mitterer, 1988) have been noted, but 
these species have survived elsewhere on the island. Local 
endemics, as Hemitrochus bowdenensis may have been, 
will be more sensitive to environmental changes; when 
more widespread species undergo local extirpations, local 
endemics may undergo extinction. The forests of south- 
eastern St. Thomas have been almost completely cut 
down and replaced by agriculture — degraded forest re- 
mains in only a very few areas. One vNonders whether 
other extinct species may turn up in the recent fossil 
record in this area and other areas that have been sim- 
ilarly degraded. 

ACKNOWLEDGMENTS 

I am indebted to Dr. G. Rosenberg for bringing this 
material to my attention and providing information on 



G. A. Goodfriend, 1992 



Page 59 



the Schumo and Smith Bowden collection, and to Dr. G, 
M. Davis for use of the collections of the Academy of 
Natural Sciences of Philadelphia, loan of the samples, 
and permission to carry out amino acid analysis on a 
shell fragment. Dr. K. Emberton and an anonymous re- 
viewer provided useful comments that were incorporat- 
ed into the manuscript. Dr. P. E. Hare provided pho- 
tographic equipment. 



LITERATURE CITED 

Baker, H, B, 1943. Some Antillean helicids. Nautilus 56(3): 
81-91. 

Bartsch, P. 1942. Thecyclopiiorid niollusksof the West Indies, 
exclusive of Cuba. Bulletin of the United States National 
Museum 181:43-141, -l-pl. 8-18 and 41 

Belli, H. M. and P. J. Bermudez 196.5 Zonation based on 
planktonic foraminifera of Middle Miocene to Pliocene 
warm-water sediments. Boletino de Informacion de la As- 
sociacion Venezueiana de Geologia, Mineria, v Petrologia 
8:121-149. 

Belli, H. M. and I Premoii Silva. 1973 Oiigocene to Recent 
planktonic foraminifera and stratigraphy of the Leg 15 
sites in the Caribbean Sea. /;i.- Edgar, N. T, et al. (eds.). 
Initial reports of the Deep Sea Drilling Project, Vol. 15. 
U.S. Government Printing Office, Washington, DC, p. 475- 
497. 

Chubb, L. J. 1958. Higher Miocene rocks of southeast Ja- 
maica. Geonotes l(l/2):25-31. 

Goodfriend, G. .\. 1987. Late Holocene morphological changes 
in a Jamaican land snail: evidence for changes in rainfall, 
/n- Berger, W. H. and L. D. Labeyrie (eds.). Abrupt cli- 
matic change. D. Reidel Publishing Co., Dordrecht, p. 
123-126. 

Goodfriend, G. A. 1989. Quaternary biogeographical history 
of land snails in Jamaica. In: Woods, C. A. (ed). Bioge- 
ography of the West Indies: past, present, and future. 
Sandhill Crane Press, Gainesville, FL, p 201-216, 



Goodfriend, G A. 1991 Patterns of racemization and epi- 
merization of amino acids in land snail shells over the 
course of the Holocene. Geochimica et Cosmochimica Acta 
55:293-302. 

Goodfriend, G .\. and R. M. Mitterer. 1988. LateQuaternar> 
land snails from the north coast of Jamaica: local extinc- 
tions and climatic change. Palaeogeography, Palaeocli- 
matology, Palaeoecology 63:293-311. 

Jung, P. 1989 Revision of the Strombina-group (Gastropoda: 
Columbeiiidae), fossil and living. Schweizerische Palaon- 
tologische Abhandlungen 111. 

Kimball, D. 1947. A new Pleurodonte from the Miocene, 
Bowden, Jamaica. Nautilus 61(2):37-39. 

Morrison, J. P. E. 1955. Notes on American cyclophorid land 
snails, with two new names, eight new species, three new 
genera, and the family Amphicyclotidae, separated on 
animal characters. Journal of the Washington Academy 
of Sciences 45(5): 149-162. 

Nordsieck, H. 1987. Revision des Systems der Helicoidea. 
Archiv fiir Molluskenkunde 118(l'/3):9-50. 

Pilsbry, H. A. 1939. Land Mollusca of North America (north 
of Mexico), Vol. I, part I Monographs of the Academy of 
Natural Sciences of Philadelphia, 3:I-XVII, 1-.573, i-ix. 

Simpson, C. T. 1895. Distribution of the land and fresh-water 
mollusks of the West Indian region, and their evidence 
with regard to past changes of land and sea. Proceedings 
of the I'nited States National Museum 17:423-451. 

Turner, R D 1958. The genus Hemitrochus in Puerto Rico. 
Occasional Papers on Mollusks, Museum of Comparative 
Zoology, Harvard University 2(22): 153-178. 

Vaught, K. C 1989. A classification of the living Mollusca. 
Abbott, R. T. and K, J. Boss (eds.) American Malacologists, 
Inc., Melbourne, Florida 

Woodring, W. P. 1925. Miocene mollusks from Bowden, Ja- 
maica. Pelecypods and scaphopods. Carnegie Institution 
of Washington Publications 366:I-IV, 1-222, pi. 1-28. 

Woodring, W. P. 1928 Miocene mollusks from Bowden, Ja- 
maica Part II Gastropods and discussion of results. Car- 
negie Institution of Washington Publication 385:I-VII, 1- 
564. 



THE NAUTILUS 106(2):60-67, 1992 



Page 60 



Dreissena polymorpha (Zebra Mussel): Colonization of Soft 
Substrata and Some Effects on Unionid Bivalves 



R. Douglas Hunter 
John F. Bailey 

Department of Biological Sciences 

Oakland University 

Rochester, MI 48309-4401 USA 



ABSTRACT 

Zebra mussel {Dreissena polymorpha) and native bivalve 
(Unionidae) densities were determined at three sites in southern 
Lake St. Clair (Ontario) using SCUBA Lake St. Clair bottom 
in this area is mostly soft (silty clay) and unionids were virtually 
the only hard surfaces present. Dreissena abundance and bio- 
mass increased from west to east: 152 individuals/m- and 26.1 
g/m- live mass (west site), to 11,655/m- and 845.8 g/m- (east 
site). Density of live unionids showed a reverse trend: 2.3/m- 
and 129.5 g/m- (west site), to 0.1/m- and 16.6 g/m- (east site). 
At the central and east sites all that was visible of any unionid 
(live or empty shell) was a mound of Dreissena. In a few of 
those samples, recruitment was so intense that the colony had 
grown laterally from the original settled surface, resulting in a 
loose mat of Dreissena over the bottom. Among the three sites 
there was a strong negative correlation between Dreissena bio- 
mass and abundance of live unionids (r= — 1.0, P<0.01, N=3). 
Most of the more heavily colonized unionids showed damage 
to the posterior valve edges. These data demonstrate that the 
zebra mussel can not only colonize lakes consisting almost en- 
tirely of soft substrata but can reach densities of 10-20 thou- 
sand/m- partially by lateral extension of colonies from an orig- 
inal "seed" surface. At such densities they will have a major 
impact on the benthos, especially indigenous unionid bivalves, 
which are likely to face virtual elimination. 

Key words: Dreissena polymorpha, unionids, soft substrata. 
Lake St. Clair. 



1991), rapid dispersal by passive means either involving 
drift by veliger larvae or boat /ship dispersal, growth to 
se.xual maturity in one season, and others (Mackie, 1991). 
In addition, it is ecologically unique, i.e. it has a niche 
which is not represented in North American fresh waters. 
Specifically, there is no other attached macrofouling in- 
vertebrate of any ecological significance in the Great 
Lakes. In some areas such as Lakes St. Clair and Erie, 
relatively high primary productivity and otherwise op- 
timal trophic conditions have further contributed to its 
success. Information on tlie distribution and abundance 
of Dreissena in Lake St. Clair has been reported by 
Hebert et al., 1991, and its biology reviewed by Mackie, 
1991. Hebert et al. (1989) and Hunter and Bailey (1991) 
made the observation that Dreissena settled on and at- 
tached readily to a variety of hard surfaces including the 
shells of native clams (Bivalvia; Unionidae), of which 
there are at least 18 species in Lake St. Clair (Nalepa 
and Gauvin, 1988). Since settled individuals attach by 
means of a byssus, they require a hard substratum, hence 
it is often assumed that a soft-bottom lake or river will 
not support significant zebra mussel colonization. 

The purpose of this study was to document the success 
of Dreissena in colonizing one region of a soft-bottom 
lake. Lake St. Clair, and to provide preliminary evidence 
of its impact on the communit\ of unionids in that area. 



INTRODUCTION 

Lake St. Clair is part of the Great Lakes drainage system 
between Lake Huron and Lake Erie. It is a relatively 
shallow lake, with a mean depth of 3.0m, a length of 
43km, and a width of 40km (Leach, 1991). Most au- 
thorities believe that the zebra mussel (Driessena poly- 
morpha Pallas, 1771) was first introduced into North 
America in Lake St. Clair in 1985 or 1986 (Hebert et 
al.. 1989, 1991; Mackie, 1991). Since then, it has spread 
rapidly and is now found in all five of the (;reat Lakes. 
Reasons for its remarkable success include relatively high 
fecundity (ca. 10'' eggs/female spawning event; Sprung, 



MATERIALS AND METHODS 

Three sites in southern Lake St. Clair were sampled for 
zebra mussels and unionids on 29 and 30 September, 
1990 (Fig. 1). All three sites were located on 42°22'3" N 
latitude which is roughly parallel to and about 5 miles 
north of the southern shoreline of Lake St. Clair in Ca- 
nadian waters (Ontario). The site farthest west (hence- 
forth the "west site") was at 82°47'30" W; the central 
and east sites were at 82°40'00" W and 82°32'30" W, 
respectively. Total distance between adjacent sites was 
about 10.3 km (6.4 mi; F'ig. 1). These sites were the same 
as locations 15, 18, and 21 of Pugsley et al., 1985, and 



R. D. Hunter and J. F. Bailey, 1992 



Page 61 



two of them, the w est and east sites, were also sampled 
b\ Nalepa and Gauvin (1988). The depth at each site 
was 6.0 ± 0.61 m. 

The bottom of Lake St. Clair is nearly all soft sediment, 
the main component of which is muddy sand, especially 
in the central part, with areas of gravel sand closer to 
shore (Leach, 1991). In the areas sampled here, the sur- 
ficial sediment was virtually the same from site to site; 
silty clay overlain by a thin layer of detritus. The lake 
bed was relatively flat with few low undulations. Mac- 
rophytes were almost non-e.\istent and nearly the only 
visible solid substrata consisted of mounds of zebra mus- 
sels. 

Zebra mussel and unionid sampling were done using 
SCUBA in two dive teams working independenth , with 
two members per team. An aluminum scjuare frame of 
0.25 m- area was used to obtain a series of samples by 
casting (throwing forward) the frame, then by touch, 
collecting all hard objects to a depth of approximately 2 
cm into the sediment. All objects within each quarter m- 
were separately bagged, brought to the surface, labeled, 
and returned to the laboratory for further analysis. Sam- 
pling was not statistically random, however due to the 
poor visibility (30-50 cm), bottom features, including 
mounds of zebra mussels, were not visible to the divers 
when the quarter m- frames were cast. Therefore this 
procedure was relatively free of sampler bias. Each dive 
team moved about 1-2 m further over the bottom be- 
tween successive samples. Sampling protocol requireti 
that a total of ten "hits" were needed for each dive team 
in order to finish a site. A hit was a sample containing 
at least one zebra mussel. Underwater records were kept 
of total casts so that misses could later be used for density 
calculations. Hence the data reported here are based on 
a different number of casts (=samples) at each site, with 
each site having 20 hits. 

In the laboratory, each quarter m- sample was indi- 
vidually e.xamined. All hard surfaces (mostly unionid 
shells) were scraped free of zebra mussels b) cutting the 
byssal threads. Removed mussels were rinsed free of sed- 
iment and the rinse water collected and passed through 
a 0.5 mm sieve in order to recover small individuals. The 
smallest zebra mussels, mostly 0.2-0.5 mm shell width, 
were not retained by this process, because the large vol- 
umes of animals and sediment made working with small- 
er meshes impractical. Care was taken to separately re- 
cord clusters of zebra mussels that were not attached to 
any other hard surface. After draining, the zebra mussels 
from each quarter m- sample were pooled for weighing. 
Live unionids were also cleaned, weighed, and prepared 
for identification. All weights reported herein are total 
(shell -I- soft tissues) live weight. A random subsample 
of zebra mussels from each site was preserved in 10% 
formalin (neutralized with CaCO,) and used for shell 
length measurements. Length measurements were to the 
nearest 0.1 mm. From these data, density vs. size class 
plots could be done by site. 

Dreissena shell to tissue mass was determined by dry- 
ing to a constant mass at 70°C, animals that had been 



St Clair/ 



MICHIGAN 




42° 30 



ONTARIO 82 30 

Figure I. Location of the three sample sites in Lake St. Clair. 
The shipping channel is shown as a dashed line. 



sorted by shell length . Whole dry animals were decal- 
cified using 5% HCl for 2 hours or until CO2 evolution 
stopped. This method gives identical results to "ashing" 
methods (Hunter and Lull, 1976). The remaining non- 
shell material (internal tissues + periostracum) was dried 
and weighed giving "tissue" dry mass. Computed from 
these values was the shelhtissue mass ratio w hich is simply 
shell mass divided by tissue mass, a number which is a 
useful inde.x of the general condition of the animal. 

RESULTS 

A total of 104 quarter m- samples were taken. At the 
west site, out of 43 quarter m- samples (=casts), 20 had 
Dreissena (=hits) and 23 did not (=misses), i.e., 53.5% 
of the 43 casts contained no zebra mussels. Numbers for 
the other two sites were: 33 casts, 13 misses for the central 
site (39.4%); and 28 casts, 8 misses for the east site (28.6%). 
Table 1 gives the abundance and biomass for Dreissena 
and for unionids at the three sites. Dreissena density was 
greatest at the east site averaging 11,655 individuals/m- 
and 845.8 g/m-; and lowest at the west site, averaging 
152 individuals/m- and 26.1 g/m-. Each of the sites 
differed significantly in biomass from the other two (one- 
factor ANOVA, P=0.0001; Scheffe's F-test, P<0.01). In 
terms of abundance, the west and central sites were not 
significantly different but the other comparisons (west 
and east; central and east) were significantlv different 
(one-factor ANOVA, P<0.000L Scheffe's F-test, P<0.01). 
Both biomass and abundance increased from west to 
central to east site. The maximum biomass of Dreissena 
recorded for any sample was at the east site at 1409 g/m^ 
which corresponds to 19,417 individuals/m^. 



Page 62 



THE NAUTILUS, Vol. 106, No, 2 



15 



I 10 

u 
O 
Si 

E 

Z 5 



■ Wood & other 

D Druses 

E Dead unionids 

S Live unionids 



S\S-.V;\.A. 



■'•^■■'•^■^■^ 



•yrrrjrjrji 



^.>.^.^.^.i». 



•./.^.^.^.^. 



S''.«\-%'\*^ 



••^•^'f'^•^• 



'■■.'■■iX'i-i-i-: 



'•'■•i-i-iX-i-iX-i-^ 



'•■.■ '.••■.■ ••.• •'.••'.*'''.■ 'if ^ 



•• J- . ^i^Z^'f' f3 



• '.' '.•^: ^.- ^.•^.•'.- '.•^.* 



■ . ^ . ^•^•^- .»«^ ■ 



West 



Central 
Site 



East 



Figure 2. Mean density of hard substrata in Lake St. Clair by 
type of material and by site. Druses are discreet clusters of 
zebra mussels. 



In contrast to Dreissena density trends, density of live 
unionids was highest at the west site and decreased mov- 
ing to the centra! and east sites (Table 1). Unionid density 
was greatest at the west site averaging 2.3 individuals/ 
m- and 129.5 g/m-. Out of 28 quarter m- samples at the 
east site, only one live unionid was found. At the west 
site there were 24 live individuals found over 43 quarter 
m- samples. The data for the east site were non-normal 
due to the presence of zeros for all but one sample. For 
purposes of statistical analysis, a small number (0.01) was 
added to all values which were then log (In) transformed. 
This transformed data set was then statistically analyzed 
for differences between sites. For both biomass and abun- 
dance, there were significant differences among sites (one- 
factor ANOVA, P<0.002), and the east site differed sig- 
nificantly from the other two (Scheffe's F-test, P<0.01). 

It was readily apparent b\' inspection of the lake bot- 
tom earlier in the season, when visibility was 2.5-3.5 m, 



that hard substrata in this general area of the lake con- 
sisted of small, discrete objects imbedded in the soft 
bottom. Later observations indicated they were fairly 
common, ranging from about 2.4 objects/m- at the west 
site to 14 objects/m- at the east site, and they were 
sufficiently small so as not to cover a very large proportion 
of a square meter except in a few of the east samples. 
With the exception of 3 pieces of wood, and two snail 
shells (Elimia livescens Menke 1830), all of these hard 
substrata were either unionids, unionid shells, or inde- 
pendent clusters of zebra mussels (Fig. 2). The densit\ 
of live unionids at the three sites has been presented in 
Table 1 and is shown in Fig. 2 relative to other hard 
substrata. The number of dead unionids (shells only with 
both valves usually connected), shows a reverse trend to 
that of the living unionids. There was, on average, 0.1 
shell at the west site, 1.4 at the central, and 7.4 at the 
east site. Likewise the number of Dreissena clusters 
(druses) that were independent from other hard substrata 
also increased from west to east (0 west, 2.6 central, and 
6.1 east; Fig. 2). 

Figure 3 illustrates the negative correlation between 
Dreissena biomass and unionid abundance. This corre- 
lation is significant P<0.01, r=-1.0, N=3. The plot of 
unionid density also shows the ratio of live to dead (L: 
D) animals by site. At the west site there were 25 live 
individuals for ever\ dead unionid. This ratio was 0.9:1 
at the central site, whereas at the east site there was only 
a single live unionid, giving an L:D ratio of 0.02:1. 

Based on subsamples of fixed material, size distribu- 
tions of Dreissena were expressed on a per m- basis (Fig. 
4). It is evident that at both the central and east sites 
there was substantial recent recruitment based on the 
large group of juveniles from about 1 to 5 mm shell 
length. Fifty-two percent of the central sample and 62% 
of the east sample were made up of juveniles 5 mm or 
less. 

In contrast, at the west site there was a pronounced 
absence of Dreissena in these smaller juvenile size classes 
(Fig. 4). The smallest animal in the \\est subsample was 
2.6 mm and only 3% of the subsample was less than 5 
mm shell length. Larger size classes were well-repre- 
sented especially in the central and east subsamples. For 
example at the central site, 34% of the subsample was 
comprised of mussels measuring 12 mm or greater, com- 



Table 1. Mean abundance and biomass (±SE) for Dreissciui and lor unionids at three sites in southern Lake St. Clair. Only a 
single live unionid was found in the east samples. 



Site in Lake St. Clair 



West 



Central 



East 



Dreissena density 
Number/m^ 
Biomass (g/m^) 

Unionid density 
Nuinber/in- 
Hiomass (g/m-) 



152 ± 36,9 
26,1 ± 6,34 

2,3 ± 0.49 
129.5 ± 27,79 



2,847 ± 620,9 

402.1 ± 87.69 

L3 ± 25 

73 4 ± 14,69 



1L655 


± 2,000.4 


845.8 


± 145.17 




0.1 




16 6 



R. D. Hunter and J. F. Bailey, 1992 



Page 63 



8 
7 
6 

CM 

Is 

n 4H 

2 2- 

1 





D Live 
■ Dead 
L:D Ratio 



Unionids , 0-02:i 



25:1 




West Central East 



1000 



800- 



^- 600 



O) 



400- 



200- 



Dreissena 

Mean ± SE 



k\\\\\\\\\N 




West 



Central 
Site 



East 



Figure 3. N4ean abundance of live and dead (empty siiells) 
unionids by site (upper plot) and mean biomass of Dreissena 
(lower plot) by site in Lake St. Clair. 



pared to 12% at the east and 9% at the west sites. Although 
ages were not determined from this material, it is likely 
that the majority of Dreissena measuring 12 or more 
mm were at least one-year-old. 

In addition to the above differences, the condition of 
the animals differed among the sites. There was a ten- 
dency for the central and especially the east animals to 
have areas of eroded shell, ie. patches where the peri- 
ostracum was clearly missing revealing the underlying 
whitish CaCOj. Although no data were recorded, there 
was also a higher proportion of empty Dreissena shells 
at the east site suggesting that mortality rates were sub- 
stantially higher there. Not only were west site densities 
far lower (there were major areas of live unionid shell 



200 



150- 



100- 




5 10 15 20 25 30 35 



200 




5 10 15 20 25 30 35 



800 



600 



400 



200 




5 10 15 20 25 30 

Shell Length (mm) 



35 



Figure 4. Density of Dreissena by shell length size class at 
three sites in Lake St. Clair. 



surface unoccupied b\ zebra mussels) but the Dreissena 
there had shells with fully intact periostraca. 

Figure 5 shows the relationship between shell length 
and shelLtissue mass ratio for each site. The general ten- 
dency is that as the mussels grow (increase in length) 
their shelhtissue ratio also increases, with a fairly linear 
relationship between the variables. Smaller size animals 
(<10 mm shell length) do not differ much between sites 
and their shelhtissue ratios are mostly within the range 
of 7-10. In contrast, larger animals (>20 mm) show- 
widely differing shelhtissue ratios at different sites (Fig. 
5). For example, a 25 mm animal from the east site would 
have a dry tissue mass averaging 34 mg and a dry shell 



Page 64 



THE NAUTILUS, Vol. 106, No. 2 



22 
20 - 

18 - 

16 
14 
r. 12 



I 10 
8 



- ^ - West 







---a- Centra 






East 




^, 

D 


- 






D 


a - - 

1 




1 1 



10 



15 20 25 30 



Shell length (mm) 

Figure 5. Shell to tissue mass ratio vs shell length for Dreissena 
at three sites in Lake St. Clair 



mass of 552 mg giving a shell:tissue ratio of 16.2. Animals 
at the central and west sites of the same 25 mm length 
would have 54 and 86 mg dry tissue and 812 and 821 
mg dry shell for a shelbtissue ratio of 15.0 and 9.5 re- 
spectively. When groups of mussels over 20 mm shell 
length were compared across the three sites, the east and 
central sites were not significantK' different in shelhtis- 
sues mass ratio, however the west site differed from the 
other two (one-factor ANOVA, P<0.0001; Sheffes F-test, 
P<0.01). In other words, larger sized Dreissena at the 
east and central sites are "tissue-deficient" compared to 
west site animals. 

It was evident from close inspection of the unionid 
shells whether of live or dead material, that many of 
them showed damage to their posterior salve edges which 
was a direct result of dense aggregations of Dreissena. 
An Anodonta grandis that is heavily infested with zebra 
mussels is shown in Fig. 6. The posterior one third to one 
half of this and other species is typically covered with a 
thick and solid mass of Dreissena. often more than 2 cm 
thick. Dreissena attachment of this magnitude renders 
normal valve movement virtually impossible and activ- 
ities normally occurring through the gape (feeding, re- 
production, respiration, locomotion, etc.) are severly cur- 
tailed. The resulting damage, illustrated in Figs. 7 and 
8 with Lampsilis siliquoidea, involved substantial de- 
formation of the shell so that the normally gradual curve 
of the posterior valve border was jagged or indented. 
The majority of specimens collected had levels of damage 
that fell between the extremes shown in Figs. 7 and 8. 
In some specimens, this damage extended internally so 
that the nacre was rough, irregular, and/or discolored. 
In a few individuals the damage was so extensive that a 
gap of 2-5 mm remained at the posterior edge even 
through the valves were fully closed (Figs. 7 and 8). For 




Figure 6. Specimen of Anodonta grandis from Lake St. Clair 
that has been heavily colonized b\ Dreissena polyrr^orpha. Such 
colonized indi\ iduals ranged from about 5 5-9.5 cm in length. 
Anodonta is shown posterior end up. 



Lampsilis siliquoidea, the only species for which there 
was sufficient material, 22.5% of the shells had no dam- 
age, 55/c had moderate damage, and 22.5% had rela- 
tively heavy damage. 

Nine species of uilionids were found among the three 
sites (Table 2). For six of the species, at least one live 
individual was obtained, but three species, Lampsilis 
ventricosa, Truncilla truncata. and Elliptiodilatata. were 
represented onK by empty shells. The unionid fauna 
from these sites was dominated by Lampsilis siliquoidea. 
such that this species comprised 79.4% of all the live 
unionids collected. When empty shells alone are consid- 
ered, then L. siliquoidea comprised 67.8% of the unionid 
communit\-, suggesting that its dominance has recently 
increased. For all of the species found, the number of 
dead individuals was greater than or equal to that of the 
live indi\iduals. 

DISCUSSION 

.\ number of European investigators have reported on 
unionids as a substratum for Dreissena. Sebestyen (1938: 
178) concluded that there was "no doubt that Dreissena 
has a decided ill effect on unionidae. " In that same study 
it was observed that the native unionids (6 species) w ere 



R. D. Hunter and ]. F. Bailey, 1992 



Page 65 



7 • 


4 


Ik 




mi 


Rki 




I 


i m 


I^^mI 


PVmI 


■'*^ 


\ ■ H 


K, ^1 


r l^^fo 


I 


\ ifl 


H^'iirl-.^afla 


^^■L ^n ^^^1 


■ 


kJ^ 


k..Imi 


^^B V A^l 


f_ 




Figures 7-8. Dorsal (7.) and posterior (8. ) views of two Lamp- 
silis siliqtioidea shells from Lake St. Clair from which all at- 
tached Dreissena have been removed. The shells on the left 
are nearly normal, whereas the shells on the right show con- 
siderable damage. One (8.) appears fuzzy due to remaining 
byssal threads. Shells are between 6 and 7 cm in length. 



in danger of disappearing entirely from Lake Balaton, 
Hungary. Sebestyen (1938) suggested that the interaction 
resulted in interference with locomotion and reduction 
in food suppK for the unionids. In contrast, Lewandowski 
(1976) concluded that Dreissena had no unfavorable ef- 
fects. In spite of this finding, he cited other European 
studies that identified the interaction as one of a com- 
petitive nature. There was evidence that Dreissena caused 
a slight negative effect on growth in shells of \ounger 
individuals and that in older unionids, the shells of Dreis- 
sena-infested individuals were thicker than shells of 
Drei.ss^na-free indixiduals. The body (soft-tissue) weight 



Nun- 


iber/m- 


Live 


Dead 


1.04 


1.73 


0.12 


0.23 


0.04 


0.15 


0.04 


0.04 


0.04 


0.04 


0.04 


0.04 





0.23 





04 


(.1 


04 



Table 2. Mean abundance of unionids by species for all three 
Lake St. Clair sites combined. Dead = empty shells. Total area 
sampled = 26 m-, so that a density of 0.04/m^ corresponds to 
one individual 



Lanipsilis siliquoidea (Barnes, 1823) 
Leptodea jragihs (Rafinesque, 1820) 
Proptera alata (Say, 1817) 
Anodonta grandis (Say, 1829) 
Ligttmia nasitta (Say, 1817) 
Fusconaia flava (Rafinesque, 1820) 
Lampsilis vcntricosa (Barnes. 1823) 
Truncilla truncata Rafinesque, 1820 
Elliptio dilalaln Rafinesque, 1820 



was not significantly altered by infestation. Lewandowski 
(1976) also observed posterior shell deformations in some 
of the Anodonta piscinalis and speculated that the si- 
phonal region was particularly attractive to post-veliger 
Dreissena. 

It is likely that the intensitv of the infestations with 
which Lewandowski was working, (20 Dreissena on each 
unionid, specificalK Anodonta piscinalis) was below the 
level required to cause an observable impact. The above 
densities are comparable to those reported here for the 
west site but are much lower than occur in the central 
or east sites in southern Lake St. Clair. 

Hebert et al. (1989) observed that Dreissena is often 
located on unionid shells in Lake St. Clair. They consid- 
ered Dreissena to be an "interference competitor" of 
native mussels (Hebert et al.. 1989; 1991). Field exper- 
iments in Lake Erie have shown that Dreissena has both 
species and sex-specific effects on unionids (Haag et al., 
1991). 

Two bits of data presented in this studv' are strongly 
suggestive that Dreissena is having a negative impact on 
the community of unionids in those areas of Lake St. 
Clair where Dreissena densities are relatively high. This 
evidence includes the negative correlation between 
Dreissena biomass and unionid densitv , as well as dam- 
age to unionid shells beneath Dreissena accumulations 
(Hunter and Bailey, 1991 ). Negative impact of Dreissena 
on unionids was also shown by Hebert et al., (1991) who 
reported that heaviK- infested unionids had half the lipid 
reserves as unaffected individuals. Mackie (1991) con- 
cluded that heavy infestations of Dreissena cause several 
kinds of negative effects on unionids, some of which 
interfere with normal locomotion. Lhiequivocally dem- 
onstrating that zebra mussels cause death of unionids is 
difficult since direct observation of death is impractical 
and the cause is likely to be multifactorial. Unionids also 
die of a varietv of other causes resulting in emptv shells 
on which Dreissena can settle. Hence, observation of 
unionid shells colonized by Dreissena is not necessarily 
evidence that they were the cause of death. Observation 
of empty unionid shells bearing deformation to the pos- 
terior valve edges of the sort illustrated in Figs. 7 and 8, 



Page 66 



THE NAUTILUS, Vol. 106, No. 2 



is strongly suggestive that Dreissena has played a role in 
the death since there is little else that will cause this kind 
of damage (R. Hoeh, pers. comm.). 

This study also suggests that the general health or 
condition of zebra mussels is inversely related to their 
densit\'. Specifically, where the population density is high, 
as is the case at the central and east sites, the larger 
animals are emaciated. The result is a shell to tissue mass 
ratio that is high compared to that of animals from low 
density situations, i.e., under such conditions adult-sized 
shells contain an undersized mass of tissue. As larger 
(older) individuals are settled on and overgrown by more 
recent recruits, it is likely they experience increasingly 
suboptimal conditions, resulting in an emaciated animal. 
Tissue degrowth under poor trophic conditions is known 
to occur in gastropods (see Russell-Hunter and Eversole, 
1976) and it is likely to be occurring in Drcisscna-in- 
fested unionids as well. 

Stanczykovvska (1975) concluded that as the Dreissena 
population density increased, their condition decreased. 
Her findings were based largely on size and weight data, 
i.e., animals of specific age groups in dense populations 
(e.g., 700/m-) tended to be smaller in both length and 
weight than those from low densitv conditions (e.g., 30/ 

Size-distribution data suggest that there are certainly 
two and probably three year-classes present at each site. 
This contention agrees with other studies w hich indicate 
that Dreissena in both Lakes Erie and St. Clair can grow 
to well over 10 mm shell length in one year (Nichols et 
al., 1991; Mackie, 1991). Thus the largest individuals 
sampled (i.e., 20 mm shell length) are likely to be in 
their second year w ith a few larger individuals possibly 
three years old. The timing of recruitment of the most 
recent cohort is distinctly different at the west site com- 
pared to the other two (Fig. 4). At the west site there is 
an absence of small juveniles, whereas they are well 
represented at both the central and east sites. The modal 
size for juveniles (defined here as individuals <10 mm 
shell length) at both the central and east sites was 2 mm. 
Forty-three percent of the central sample and 35% of 
the east sample consisted of juveniles less than 3 mm 
shell length. It would appear that a relatively intense but 
brief recruitment of post-veligers occurred at the west 
site perhaps 4-8 weeks before these samples were taken, 
resulting in a relatively well-defined cohort of about 5- 
10 mm in late September (as seen in Fig. 4). In contrast, 
a more recent recruitment is in evidence at the other 
two sites beginning perhaps four weeks previously and 
possibly still continuing or just ending as of late Septem- 
ber. It appears that there was little recruitment activity 
at the central site during the time when recruitment was 
intense at the west site (Fig. 4). 

These contrasting recruitment patterns are likeK to be 
due to differences in veliger density in different major 
water masses and to wind-induced irregularities influ- 
encing the extent of these water masses. Water on the 
western and northwestern side of the lake ("Huron wa- 
ter) is largely from the Lake Huron/St. Clair River flow 



system (Leach, 1980, 1991). Huron water is relatively 
veliger-poor due to very low Dreissena densit\ in that 
area in 1990, which would result in low recruitment 
around the west site. Water in eastern and southeastern 
Lake St. Clair ("St. Clair" water) is more stable and 
productive due to nutrient enrichment from Ontario riv- 
ers (Leach, 1980, 1991). In this water mass Dreissena 
reached much higher densities (eg. central and east sites) 
than elsewhere in the lake. Changing wind speed and 
direction move the boundries of these water masses to 
some degree. If the St. Clair water mass extended over 
the west site for a few da>s in summer, then it could 
produce the recruitment pulse that is seen in September 
as a distinct cohort of settled juveniles. 

Two of the sites sampled in this study (west and east) 
were approximately the same as sampled by Nalepa and 
Gauvin (1988) in their study of unionid distribution in 
Lake St. Clair. Since their samples were taken in Sep- 
tember, 1986, they contained no zebra mussels. The av- 
erage unionid densitv thev reported for the southern half 
of Lake St. Clair was 2^6 ± 1.91/m- (.x ± s, N=15) 
compared to the average for the present study of 1.2 ± 
1.10/m2 (N=3). Nalepa and Gauvin (1988)' found 18 
species of unionids of which nine were the same as those 
in Table 2. Of the nine species listed in Table 2, four were 
represented by empty shells only. Although they col- 
lected material from a much larger area it is interesting 
to note that seven out of nine species common to the two 
studies, declined as a percentage of the total numbers 
collected. Only Lampsilis siliquoidea (=L. radiata sili- 
qiioidea of Nalepa and Gauvin, 1988) increased substan- 
tially as a proportion of the total unionid community. 
That species comprised 45% of Nalepa and Gauvin's 
unionid abundance whereas it was 79% of our samples. 
Although the goals of these two studies differ, it is possible 
that L. siliquoidea may be less affected by Dreissena 
colonization than other species hence has become an even 
more dominant member of the unionid community than 
it was before. In their study, Nalepa and Gauvin con- 
cluded that the diversity and composition of unionids in 
Lake St Clair appeared to be relati\ely unchanged since 
the turn of the century. With the introduction of the 
zebra mussel at very nearly the same time and location 
as their stud)-, the unionid communit\ will now un- 
doubtedK change very substantialK . Because no indig- 
enous species appears to be able to tolerate or escape 
Dreissena colonization, it is likeK that the unionid com- 
munit) in Lake St. Clair will be virtually eliminated in 
a few \ear's time. 

ACKNOWLEDGEMENTS 

This study was supported in part by the Michigan Sea 
Grant College Program (Project No. M /PM-3H). The 
cheerful enthusiasm and careful sampling of SCUBA 
divers Luke Clyburn, Phyllis Higman, and Mary Black- 
burn is gratefully acknowledged. Joe Leach (Ontario 
Ministr\ of Natural Resourses) and Bob Haas (Michigan 
DNR) provided thoughtful discussion on Lake St. Clair 



R, D. Hunter and J. F. Bailey, 1992 



Page 67 



water masses. Special thanks are due to LCDR Luke 
CK'burn (NSCC) for providing the boat transportation 
and dive support. Randy Hoeh (Univ. of Michigan, Mu- 
seum of Zoology) kindly identified or confirmed our iden- 
tification of the unionids. 

LITERATURE CITED 

Garton, D., and VV. Haag. 1990. Reproduction and recruit- 
ment of Drcissena during the first invasion year in western 
Lake Erie. Abstract. In. Zebra mussels: the Great Lakes 
E.xperience, Feb., 19, Univ. of Guelph, Guelph, Ontario, 
p4. 

Haag, W R . D J Berg, and D W Garton 1991 Drcissena 
polynunplia colonies encrusting native unionid bivalves 
produce species-specific and .sex-specific effects. Journal of 
Shellfish Research 10:2.59. 

Hebert, P.D.N. , B.W. Muncaster, and G.L. Mackie. 1989. 
Ecological and genetic studies on Drcissena polijrnorpha 
(Pallas): A new mollusc in the Great Lakes, Canadian 
Journal of Fisheries and Aquatic Science 46:1587-1591. 

Hebert, P.D N,, C.C. Wilson, M.H. Murdoch, and R, Lazar. 
1991, Demography and ecological impacts of the invad- 
ing mollusc Drcissena poli/nwrpha. Canadian Journal of 
Zoology 69:405-409, 

Hunter, R,D. and VV.W. Lull. 1976. A comparison of two 
methods for estimating the weight of inorganic materials 
in molluscs. Malacological Review 9:118-120. 

Hunter, R.D, and J.F. Bailey. 1991, Colonization of soft sub- 
strata by zebra mussels: role of and cost to native bivalves. 
Journal of Shellfish Research 10249-250 

Leach, J H. 1980. Limnological sampling intensity in Lake 
St. Clair in relation to distribution of water masses. Journal 
of Great Lakes Research 6:141-145. 

Leach. J.H. 1991. Biota of Lake St. Clair: habitat evaluation 
and environmental assessment. Hydrobiologia 219:187- 
202. 



Lewandowski, K 1976. Unionidae as a substratum for Dreis- 
sena polymorpha Pall. Polish Archives of Hydrobiology 
23:409-420. 

Mackie, G.L. 1991. Biology of the exotic zebra mussel, Dreis- 
sena pohjmorpha, relative to native bivalves and its po- 
tential impact in Lake St. Clair H\drobiologia 219:251- 
268. 

Nalepa, T.F. and J.M. Gauvin. 1988. Distribution, abundance, 
and biomass of freshwater mussels (Bivalvia: Unionidae) 
in Lake St. Clair. Journal of Great Lakes Research 14:41 1- 
419. 

Nichols, S.J., A.M. Bitterman, and F, Ely, 1991, Growth rates 
of Drcissena polymorpha in the St. Clair River and Lakes 
St. Clair, Erie, and Ontario from June to November, 1990. 
Journal of Shellfish Research 10: 250. 

Pugsley, C.W., P.D.N, Hebert, G,W. Wood, G.W. Brotea, and 
T.W. Obal. 1985. Distribution of contaminants in clams 
and sediments from the Huron-Erie corridor. I - PCBs and 
octachlorostv rene Journal of Great Lakes Research 11: 
275-289. 

Russell-Hunter. WD, and AG. Eversole. 1976. Evidence for 
tissue degrowth in starved freshwater pulmonale snails 
(Helisoma trivolvis) from tissue, carbon, and nitrogen 
analyses. Comparative Biochemistry and Phvsiologv 54A: 
447-453. 

Sebestyen, O. 1938. Colonization of two new fauna-elements 
of Pontus-origin (Drcissena polymorpha Pall, and Coro- 
phium curvispinum GO, Sars forma devium Wundsch) 
in Lake Balaton, Verhandhmgen Internationale Vereini- 
gung Limnologie 8:169-181, 

Sprung, M, 1991, Costs of reproduction: a study on metabolic 
requirements of the gonads and fecundity of the bivalve 
Drcissena polymorpha Malacologia 33: 63-70. 

Stanczykowska, A. 1975. Ecosystem of the Mikolajskie Lake. 
Regularities of the Drcissena polymorpha Pall (Bivalvia) 
occurrence and its function in the lake, Polish Archives of 
Hydrobiology 22:73-78. 



THE NAUTILUS 106(2):68-71, 1992 



Page 68 



New Ecphoras (Gastropoda: Thaididae: Ecphorinae) from the 
Calvert Formation of Maryland (Langhian Miocene) 



Edward J. Petuch 

Department of Geology 
Florida Atlantic University 
Boca Raton, FL 33431 



ABSTRACT 

Three new ecphoras are described from the relatively unstudied 
Shattuck Zones 12 and 14 of the Plum Point Member of the 
Calvert Formation (Langhian Miocene) of Maryland. The new 
species, Ecphora {Ecphora) chesapeakensis n.sp., Ecphora (Ec- 
phora) turneri n.sp., and Ecphora (Trisecphora) scientistensis 
n.sp., fill in gaps in the evolutionary lineages of the Ecphora 
ganlnerae species complex, the Ecphora choptankensis species 
complex, and the Ecphora (Trisecphora) tricostata species 
complex, respectively. 

Key words: Ecphora, Gastropoda; Calvert Formation; Mio- 
cene; Marvland. 



INTRODUCTION 

Since the publication of the "Field Guide to the Ec- 
phoras" (Petuch, 1989), three new species of Ecphora 
Conrad, 1843 have been brought to my attention. These 
were collected in the relatively unstudied Beds 12 and 
14 (Zones 12 and 14 of Shattuck, 1904:xxvii) of the Cal- 
vert Formation (Plum Point Member), along the Calvert 
Cliffs of western Chesapeake Bay, in Calvert County, 
Maryland. As the preservation of the moUuscan fossils of 
Beds 12 and 14 is not good, the result of extensive leach- 
ing by groundwater, most specimens of upper Calvert 
ecphoras are collected in a fragmentary state. In spite of 
this difficult collecting of extremely fragile and shattered 
specimens, Mr. Joseph Turner of Baltimore, Maryland, 
through great diligence and patience, has managed to 
extract a number of unusual forms from these virtually 
unworkable beds. Through his generosity in the donation 
of numerous study specimens, I was able to determine 
that three ecphoras were new to science, and these are 
described here. This paper is considered an addendum 
to "Field Guide to the Ecphoras." 

The three new species fill gaps in the evolutionary 
lineages of three main groups of ecphoras. One new 
species, here named Ecphora (Ecphora) turneri n.sp., is 
morphologically intermediate between E. [Ecphora) 
wardi I'etuch, 1989 (figure 5) from Bed 10 of the Calvert 
Formation (Langhian Stage, Miocene) and £. [Ecphora) 
choptankensis vokesi Petuch, 1989 (figure 6) from Bed 



16 of the Choptank Formation (Serravallian Stage, Mio- 
cene). Likewise, a new three-ribbed ecphora, here named 
Ecphora (Trisecphora) scientistensis n.sp., is interme- 
diate between E. [Trisecphora) eccentrica Petuch, 1989 
(figure 9) of Bed 10 of the Calvert Formation and E. 
(Trisecphora) smithae Petuch, 1988 (figure 11) from Bed 
16 of the Choptank Formation. The third new species, 
here named Ecphora (Ecphora) chesapeakensis n.sp., is 
the oldest known member of the E. (Ecphora) gardnerae 
Wilson, 1987 species complex. This new ecphora is an- 
cestral to £. (Ecphora) williamsi Ward and Gilinsky, 
1988 (figure 2) from Bed 19 of the Choptank Formation, 
which previously had been thought (Petuch, 1989) to 
have been the original progenitor of the wide-ribbed E. 
gardnerae species group. 

In this paper, I use the morphological criteria for ec- 
phora species-level determinations that are outlined in 
my earlier works (Petuch, 1988, 1989). The holotypes of 
the new taxa are deposited in the invertebrate paleon- 
tology collection of the Florida Museum of Natural His- 
tory, University of Florida, Gainesville, Florida, and bear 
UF numbers. 

SYSTEMATICS 

Class Gastropoda 

Subclass Prosobranchia 

Order Caenogastropoda 

Superfamily Muricacea 

Family Thaididae 

Subfamily Ecphorinae Petuch, 1988 

Genus Ecphora Conrad, 1843 



chesapeakensis new species 



Ecphora [Ecphora 
(figures 3, 4) 

Materials examined: Holotijpe: Length (incomplete 
and reconstructed) 28 mm, in Shattuck Zone 14, Plum 
Point Member, Calvert Formation, at Scientists C'liffs, 
(Calvert County, Maryland, Chesapeake Bay, UF 23798; 
Paratype: length (incomplete and reconstructed) 35 mm, 
.same locality as holotype, Petuch collection. 

Description: Shell cylindrical in shape, slightly inflated; 
shoulder rountled; body whorl ornamented with 4 large, 



E. J. Petuch, 1992 



Page 69 




Figures 1-11. Ecphoras from the Miocene of Maryland. 1. Ecphora (Ecphora) calvertensis Petuch, 1988, dorsal view of 43 mm 
specimen, Shattuck Zone 12, Calvert Formation. 2. Ecphora (Ecphora) willianui Ward and Gilinsky, 1988, dorsal view of 57 mm 
specimen, Shattuck Zone 19, Choptank Formation. 3. Ecphora (Ecphora) chesapeahensu n.sp., ventral view of holotype, length 
(incomplete) 28 mm, UF 23798, Shattuck Zone 14, Calvert Formation. 4. Ecphora (Ecphora) chesapeakensis n.sp., ventral view 
of paratype, length (incomplete) 35 mm, Shattuck Zone 14, Calvert Formation 5. Ecphora (Ecphora) wardi Petuch, 1989, ventral 



Page 70 



THE NAUTILUS. Vol 106, No. 2 



wide, rounded, adherent cordlike ribs that are sHghtK 
"T"-shaped in cross section; ribs sculptured with 1-4 
thin, impressed spiral threads; shoulder rib largest and 
widest, curving upward ( posteriorward ) to produce can- 
aliculate spire whorls; wide shoulder rib slightK in- 
curved, producing distinctly rounded appearance; areas 
between ribs relatively smooth, with only few very fine 
spiral threads; siphonal canal well developed, orna- 
mented with numerous large spiral threads; umbilicus 
narrow. 

Etymology : Named for the Chesapeake Bay, which bor- 
ders the type locality. 

Discussion: Ecphora chesapeakensis is closest to E. wil- 
liarwii Ward and Gilinsky, 1988 (figure 2) from the youn- 
ger Choptank Formation, and appears to be its direct 
ancestor. The new species differs from its Choptank de- 
scendant in being a smaller, less inflated shell with thin- 
ner, less rounded ribs. The ribs of E. williamsi are lower 
and more adherent, while those of E. chesapeakensis 
project farther from the body whorl. Both species have 
similar wide, rounded, incurved shoulder ribs. Ecphora 
cheaapeakenais is ai.so similar to £. calvertenaiH Petuch, 
1988 (figure Ij from Shattuck Zone 12, but differs in 
having distinctly rounded, cordlike ribs instead of the 
sharply-flanged, "T"-shaped ribs of the older Calvert 
species. This new ecphora constitutes a morphfjlogical 
link between the generalized E. calvertemis and the E. 
gardnerae species complex of the later Miocene. Ecphora 
cheaapeakennia is confined to Shattuck Zone 14 (Bed 14, 
Calvertj. 

Ecphf/ra (Ecphora) turneri new species 
(figures 7, H) 

Material examirH'<l: llololype: Length (incomplete) 38 
rnrn, in Shattuck Zone 14, Plum Point Member, Calvert 
FoTinayioii, approximately I km rjortli of (iovernor Bun, 
Calvert fJiifs, Calvert Oiurrty, Maryland, ( ;(iesa[jcakc 
Bay, UF 2)465; ParaUjpe: length 44 rrun, sann- locality 
as holotype, Peluch collection. 

DeHcriplion: Shf;ll cylindrical in shape, with sharply 
angled shoulder; body whorl ornamented with 4 thin, 
narrow, hladelike ribs; edge of ribs rounded; subsutural 
areas flattened, (jroducing stepjjed, s<alariiorin spire; ar- 
eas between rib.s smooth, without spiral s( iilptnre; si- 
phonal canal (elongated; utnbilic us n;irrow Uiit well rie 
veloped. 



Etymolog>': The ta.xon honors Mr. Joseph Turner of 
Baltimore, Maryland, who generously donated a large 
suite of research material from Shattuck Zones 12 and 
14 of the Calvert Formation. 

Discussion: Ecphora turneri is closest to £. choptanken- 
sis vokesi Petuch, 1989 (figure 6) from Shattuck Zone 16 
of the Choptank Formation, but differs in being a much 
smaller, much more cylindrical and slender shell, with 
lower, less projecting ribs. The umbilicus of E. turneri 
is also much narrower than that of E. choptankensis 
vokesi, and the new species has a proportionally longer 
siphonal canal. Ecphora turneri is confined to Shattuck 
Zone 14 (Bed 14, Calvert). 

Subgenus Trisecphora Petuch, 1988 

Ecphora (Trisecphora) scientistensis 
new species 
(figure 10) 

Material examined: Holotype: Length (incomplete, 
fragmentary) 29 mm, in Shattuck Zone 12, Plum Point 
Member, Calvert Formation, at Scientists Cliffs, Calvert 
County, Maryland, Chesapeake Bay, UF 23799. 

Description: Shell cylindrical, slightly inflated in shape; 
shoulder sharply angled; subsutural area flattened, pla- 
nar; spire whorls .scalariform, ste[)ped; body whorl or- 
namented with 3 large cordlike ribs; ribs rounded on 
edges, sculptured with 1 or 2 faint, shallow impressed 
spiral threads; areas between ribs smooth, without spiral 
sculpture. 

Etymology: Named for the Scientists (>liffs, ("alvert 
County, Maryland, the type locality. 

Discussion: Ecphora (Trisecphora) scientistensis is 
(•los<-st to /•;. (Trisecphora) eccentrica Petuch, 1989 (fig- 
ure 9) from Shattuck Zone 10 of the Calvert I'ornKilion, 
but differs in being a more cylindrical shell, by lacking 
fine spiral threaded .sculpturing between the ribs, and by 
having a lower, ;idherent spire that does not become 
detaclied and uncoiled. 'I"he ribs of /•,'. (Trisecphora) 
scientistensis are also thicker than those of E. (Trisec- 
phora) eccentrica. The new si)ecies is also somewhat 
similar to E. (Trisecphora) tricostata Martin, 1904, also 
from Zone 10, but that well known species luis w ider ribs 
that are "T"-sliai)ecl in cross section and al.so has a more 
inflated, globo.se shell. Ecphora (Trisecphora) tricostata, 
like /','. (Trisecphora) eccentrica, has detached, uncoiled 
whorls, wliile the whorls of E. (Trisecphora) scientislen- 



vicw of lioffjtypc, len(;lli 70 iiiiii, Slialliick Zone 10, Calvert I'oriniitioii (tiikcn from Ward and (illiiisky, 1988: plate I. figure 4) 
h. Erphora (Ecphora) clioptankciisis ixikcsl I'i^Iik li \'IH'), vcnlral view of paratype, fenglli ()2 nun, Siialluck Zone !(>, Ofioplank 
(''orriiiition 7, K. Ecphora (Ecphora) Inrncrt w .p doisal .imi ventral views of liololype, Icngtfi (iiudinpiete) 38 nnn, I'l'' 214(i5, 
SlialtiKJc /one M, Calvert T'or/iialioii '). Etjihorii {'t'rb.c(pli()r(i) ccccnlrlra retiicli, lOKO, cforsaf view ol paiatvpe. fcnglli 5(i mm, 
Sfiiitliiek /.one 10. (lufvcrt I'drnialion 10. /'>7)/i«r« C/V/.srr/i/ifyff/) vi /cii^/.v/c/i.vi.v nsp. dorsal view ol ' 
(raxnienlary) 2\) mm. Ill'' 23700, Sliallmk /one 12, < advert lormalioir I 



viitw ol paratype, leiigtli 75 mm, Stiallii<k /one 10, ( liioplinjk I'onnatioii 



lololvpe, leiiglfi (incomplete, 
l'.( liliiiKi ( '/'rrvcc/i/iora) siiilllidf I'dniii, 10S8. dorsal 



E. J. Petuch. 199: 



«s are tightly coiled and attached. The new species is Petuch, E. J 19S9 Field guide to the Ecphoras. CoastsI Ed- 

confined to Shattuck Zone 12 of the Calvert Formation ueation and Research Foundati.:- " - -:es%ille. \"A_ 

(Bed 12. Calvert). 140 p. 

Shattuck. G B, 1904. Geological and pai^nJuccaC'^iical relatiotis. 

with a review of eariier investigations. In. Clark. W R. 

G B. ShattucL and W H. DalT eds. . The Miocece de- 

LITER.\TIRE CITED poats of Mar> land Mar> land Geological Sun.e> . Miocene. 

p. xiii-cniL 
Petuch. E. J. 19SS. New species of Ecphora and ecphorine \\'ard. I_ \V. and X. L Gilinsk> . 19SS. Ecphonz Gastropoda; 

thaidids from the Miocene of Chesapeake Ba>. Mainland. Muricidae' from the Chesapeake group of Maryland and 

U.S-.\. Bulletin of Paleomalacolog> lvl':l-16. Virginia. Notuke Naturae 469(15 Marcfa':l-il. 



THE NAUTILUS 106(2):72-75, 1992 



Page 72 



Parallel Development-Depth Trends in Deep-Sea 
Turrid Snails from the Eastern and Western 
North Atlantic 



Elizabeth Potter 

Massatliusetts Water Resources 

Authority 
100 First Avenue 
Charlestown, MA 02129 



Michael A. Rex' 

Department of Biology 
University of Massachusetts 
Boston, MA 02125 



Bouchet (1976a) first pointed out that a surprisingly high 
proportion of deep-sea prosobranchs bear larval shells 
that indicate planktotrophic development, a finding since 
supported by numerous surveys of deep-sea snail faunas 
(e.g. Bouchet and Waren, 1979, 1980, 1985; Rex and 
Waren, 1982; Colman et al., 1986a). Studies using oo- 
cyte-size distributions (Rex et al., 1979; Colman et al., 
1986b), stable isotope composition of larval and adult 
shells (Bouchet and Pontes, 1981; Killingley and Rex, 
1985) and direct recovery of larvae from surface plank- 
ton and benthic egg capsules (Bouchet, 1976b; Bouchet 
and Waren, 1980; Gustafson et al., 1991) have corrob- 
orated the use of larval shell morphology to infer mode 
of development, and revealed that planktotrophic larvae 
of some species undergo vertical migration in the water 
column. Both the frequency of planktotrophic devel- 
opment in deep-sea snails and ontogenetic vertical mi- 
gration violate theoretical predictions about larval de- 
velopment in this cold, remote and nutrient-poor 
environment (Thorson, 1950; Vance, 1973). Rex and 
Waren (1982) showed that the incidence of plankto- 
trophic development in prosobranchs actually increases 
with depth in the western North Atlantic. Bathymetric 
trends in developmental mode have not been analyzed 
in other regions of the deep sea to establish the generality 
of this pattern. Here, we compare development-depth 
trends between the eastern and western North Atlantic 
for the Turridae, the largest family of deep-sea proso- 
branchs. 

The morphological criteria for determining the mode 
of larval development are particularly clear in turrids 
(Bouchet, 1990) and have been confirmed by direct ob- 
servation in coastal representatives (e.g. Labour, 1934; 
Thorson, 1946; Thiriot-Quievreux, 1972). Planktotro[ihic 



' Correspond witfi M A Rex 



larval shells have high spires, 2-5 whorls, brown color 
and fine reticulate or ribbed sculpture. The shell consists 
of two distinct parts: a minute Protoconch I at the apex 
which is deposited before hatching, and a larger Proto- 
conch II grown during the planktotrophic phase of de- 
velopment (Robertson, 1974). Commonly, these larval 
shells are of the sinusigera type (Robertson, 1974) which 
have a distinctive beak on the aperture that projects 
between the velar lobes during the free-swimming stage. 
In contrast, shells of nonplanktotrophic larvae have a 
single bulbous whorl, are of the same white or gray color 
as the adult shell, and either lack visible sculpture or 
have very simple robust sculpture. Nonplanktotrophic 
larvae are provisioned with sufficient food to undergo 
development and hatch from benthic egg capsules. 
Though nonplanktotrophic larvae generally seem to have 
less dispersal potential than planktotrophic forms, the 
nonplanktotrophic larva of at least one turrid has a brief 
swimiTiing phase after hatching and then lives demersally 
before metamorphosis (Shiniek, 1986). 

The two data .sets compared are from Rex and Waren 
(1982) and Bouchet and Waren (1980) for the western 
and eastern North Atlantic respectively. We used only 
those species for which larval shells are known and which 
belong to genera included in the Turridae by Bouchet 
and Waren's taxonomic revision. This included 39 west- 
ern species (80% with planktotrophic development) and 
90 eastern species (74% with planktotrophic develop- 
ment). Rex and Waren (1982: Table 1) provide relative 
abundance data for species in 20 epibenthic sled samples 
collected from south of New England, whereas Bouchet 
and Waren report depth records for material from many 
sources includeil in their taxonomic revision. To make 
the two data bases more compatible, we determined the 
known depth range for each species in the separate 
regions, and calculated the percentage of species with 
planktotrophic development in each 500 m depth inter- 
val from the continental shelf to the aby.s.sal plain. We 
then regressed the frequencies of planktotrophic devel- 



E. Potter and M. A. Rex, 1992 



Page 73 



.2 B 

S o 
•r Q 



•S £ 
§2 

o a 



10 - 



0.8 - 



0,6 - 



0.4 



0.2 



' 












y 














D D ^/^mr 


■ 










^^y^yl^r. 


■ 






• 


V 


^y^ 






• 


A 


<y 




i:?^ 


r 


^ 




D 










n Weslem North Atlanlic 




- • 










• Eastern North AUanUc 






-1 • 1 ' 1 









1000 



2000 



3000 



4000 



5(XX) 



6000 



Depth (m) 

Figure I . Relationships between the proportion of species with 
planktotrophic development in deep-sea turrid snails and depth, 
for the western (WNA) and eastern (ENA) North Atlantic. 
Regression equations and their statistics for WNA and ENA 
respectively are; Y = 0.-161 + 0.t)0012X; N=ll, R^ = 0.801. 
F = 36.155. P<0.001; Y = 486 + O.OOOIO.X; N = I1. R- = 
0.798. F = 35.476. P<0.001. 



opment against depth, and performed an analysis of co- 
variance (ANCOVA) to test for differences in slope and 
elevation of the development-depth relationships be- 
tween the two regions. 

Turrids from both regions show highly significant in- 
creases in the incidence of planktotrophic development 
with depth, and the trends are very similar (Figure 1; 
see caption for regression ecinations and their statistics). 
The ANCOVA reveals that the regressions for the eastern 
and western North Atlantic faunas are not statistically 
different in either slope (F = 0.855; d.f. = 1,18; n.s.) or 
elevation (F = 0.952; d.f. = 1,19; n.s.). When variables 
used in a regression are percentage values, such as here 
with the frequency of planktotrophy, arcsine transfor- 
mations can be used to correct for departures from nor- 
mality (Sokal and Rohlf, 1981). When this is done the 
regressions remain highly significant (R- = 0.736, F = 
25.040. N = 11. P < 0.001; a'nd R- = 0.797, F = 35.406, 
N = 11, P < 0.001 for western and eastern faunas re- 
spectively), and are still indistinguishable in slope (F = 
2.246; d.f. = 1,18; n.s.) and elevation (F = 3.688; d.f. = 
1,19; n.s.) by the ANCOVA. There are some idiosyncra- 
sies in the trends. For example, there is a marked increase 
of about 40% in planktotrophy at 2500 m in the western 
fauna, and a large increment of about 20% at 1000 m 
in the eastern group (Figure 1). But overall, the two 
faunas show convincing increases in planktotrophic de- 
velopment with depth, and are remarkably alike in both 
the level of planktotrophy and its rate of increase with 
depth. 

The analysis above suggests that a depth-related in- 
crease in the proportion of turrid species with plankto- 



trophic development is widespread in the North Atlantic. 
It would be interesting to expand this research to include 
the proportion of endemic and shared species in eastern 
and western basins. This would enable us to explore, for 
example, whether horizontal geographic range is asso- 
ciated with mode of development (Jablonski and Lutz, 
1983). We could also determine what proportion of the 
similarity in response show n in Figure 1 is attributable 
to shared and endemic components of the two faunas. 
At present, however, it is not possible to critically eval- 
uate similarity in species composition on such large spa- 
tial scales. There has been no taxonomic synthesis of 
deep-sea turrids for the North Atlantic as a whole. Except 
for abundant and well-known species (see e.g. Etter and 
Rex, 1990), it is still unclear which species are actually 
common to both regions. Also, the large difference in the 
number of species represented in the two data bases 
places severe limitations on any calculated faunal simi- 
larity. The degree to which the two available species lists 
reflect a real shift in the regional diversity of turrids or 
merely the very substantial inter-regional difference in 
sampling intensity remains to be established. 

It is intriguing that deep-sea turrids should have such 
a high proportion of species with planktotrophic devel- 
opment, and even show an increase in planktotrophy 
with depth, in an environment where nonplanktotrophic 
development appears to predominate in most taxa. 
Bouchet (1988) proposed that modes of larval develop- 
ment in deep-sea taxa, including snails, represent phy- 
logenetic constraints, and are of no immediate adaptive 
significance. His arguments are especially compelling for 
groups like the archaeogastropods, which are found 
throughout the deep sea and have only nonplanktotroph- 
ic development. In these groups, larval development may 
be incidental to other adaptive features directly related 
to geographic distribution. In turrids, developmental 
mode is a fixed species-wide characteristic (Bouchet 1989, 
1990). Planktotrophy is an ancestral condition that has 
been lost in many lineages, presumably during the course 
of speciation and adaptive radiation. The loss of feeding 
larvae in snails is potentially reversible (Strathmann, 
1978), but there is no evidence that species have reac- 
quired planktotrophic development during the evolu- 
tionary histor\ of turrids (Bouchet, 1990). 

However, while historical evolutionary events are re- 
sponsible for establishing the overall developmental 
makeup of deep-sea snail assemblages, it is difficult to 
accoimt for the clear and consistent depth trends shown 
in Figure 1 in purely nonadaptive terms. Environmental 
gradients associated with depth have caused marked 
bathymetric changes in the density (Rex et ai, 1990), 
species diversity (Rex, 1983) and species composition 
(Rex, 1977) of deep-sea snail faunas Individual species 
also exhibit clinal effects with depth (Etter and Rex, 
1990), particularly across the upper bathyal region where 
nonplanktotrophic development is more common (Fig- 
ure 1). Depth-correlated selective gradients might be 
responsible for both driving the processes of population 
differentiation and speciation that ultimately generate 



Page 74 



THE NAUTILUS, Vol. 106, No. 2 



new species, and for actively maintaining their geo- 
graphic distributions. Rex and Waren (1982) suggested 
two ways in which modes of larval development in tur- 
rids might be adaptively related to this selective gradient. 
Nonplanktotrophic development might confer an ad- 
vantage at upper bathyal depths because species' bathy- 
metric and horizontal ranges are much more restricted 
there. Large-scale dispersal in ocean currents would car- 
ry planktotrophic larvae away from adult habitats and 
limit successful recruitment. Conversely, selection may 
favor increased dispersal ability at greater depths to en- 
able turrids to track progressively more rare and patchy 
pre> resources. 

Clearly, adaptation cannot provide a complete expla- 
nation for the trends shown in Figure 1, because species 
with planktotrophic and nonplanktotrophic develop- 
ment are found at all depths. Latitudinal gradients in 
the proportion of development types in prosobranchs 
(Thorson, 1950) present a similar problem of interpre- 
tation. Adaptation and phyletic constraint are both plau- 
sible hypotheses to account for part of the bathymetric 
variation in developmental modes of deep-sea turrids. 
Presently, it is not feasible to test these hypotheses ex- 
perimentally b\ showing whether modes of development 
are adaptive to the ecological conditions at different 
depths in the sense that they actually improve fitness. 
However, a phylogenetic study of deep-sea turrids could 
demonstrate whether or not the same developmental 
modes have arisen independentK in similar selective re- 
gimes. 

We thank Philippe Bouchet and Ron Etter for reading 
the manuscript. Ron Etter also provided assistance in 
performing the ANCOVA. This research was supported 
by the Department of Biology, University of Massachu- 
setts at Boston. 



LITERATURE CITED 

Bouchet, P. 1976a. Mise en evidence destades larvairesplaiic- 

loniques chez des Gasteropodes Prosobranches des etages 

hathyal et abyssal. Bulletin du N4useum national d'Histoire 

naturelle 277; 947-972. 
Bouchet, P. 19761) Mi.se en evidence d uiie migration de 

larves veligeres entre I'etage abyssal et la surface, t^omptes 

Kendus des Seances de rAcademie des Sciences, Paris, 283: 

821-824. 
Bouchet, P. 1988. Adaptive and non-adaptive strategies in 

larval biology of deep-sea snails. Abstracts of the Fifth 

Deep-Sea Biologv Svin[)osiuni, Centre de Brest, France, 

IFREMEK, p. 46 
Bouchet, P. 1989. .'\ review ol ijoci ilogoin in gastropods. 

Journal of Molluscan Studies .55; 67-78. 
Bouchet, P. 1990. 'I'urrid genera and mode ol iic\flopmciit; 

the use and ahu.sc ot protocdmli moipliologN Malaroloiiia 

32; 69-77. 
Bouchet, P. and J.-C. Pontes. 1981. Migrations verticales des 

larves de Gasteropodes ahyssaux; arRumcnIs nouveanx dus 

a I analy.se isolopi(|ue de la cocpiillc larvaire et posllarvaire. 

(>omptes Rendus des Seances de I Academic des Sciences, 

Paris (3) 292; 1005- 1008 
Bouchet, P. and .■\. Waren. 1979 Theahvssal mollusraii lauii;i 



of the Norwegian Sea and its relation toother faunas. Sarsia 
64; 211-243. 

Bouchet. P. and A. Waren 1980. Revision of the north-east 
Atlantic batliyal and abyssal Turridae (Mollusca, Gastrop- 
oda), Journal of Molluscan Studies, Supplement 8; 1-119. 

Bouchet, P and A Waren. 1985. Revision of the northeast 
Atlantic bathyal arid abyssal Neogastropoda, excluding 
Turridae (Mollusca, Gastropoda). Bollettino Malacologico, 
Supplemento 1; 123-296. 

Cx)lman, J.G., PA. Tyler and J.D. Gage. 1986a. Larval de- 
velopment of deep-sea gastropods (Prosobrancliia; Neo- 
gastropoda) from the Rockall Trough Journal of the Ma- 
rine Biological Association of the Lnited Kingdom 66; 951- 
965, 

Colnian, J.G., PA. T\ler and J D Gage 1986b The repro- 
ductive biology of Colus jcjjreysianus (Gastropoda; Pros- 
obranchia) from 2200 m in the N.E. Atlantic. Journal of 
Molluscan Studies 52; 45-54 

Etter, R.J. and M.A. Rex. 1990, Population differentiation 
decreases with depth in deep-sea gastropods, Deep-Sea 
Research .37; 1251-1261. 

Gustahson. R.G., D T J Littlewood and R,A, Lutz 1991, Gas- 
tropod egg capsules ant! their contents from deep-sea hy- 
drothermal vent environments. Biological Bulletin 180; 
34-.55, 

Jahlonski, D and R,A, Lutz 1983. Larval ecology of marine 
benthic invertebrates; paleobiological implications. Bio- 
logical Reviews 58; 21-89. 

Killingley, J.S, and M.A, Re,\, 1985, Mode of larval devel- 
opment in some deep-sea gastropods indicated b\ oxygen- 
18 values of their carbonate shells, Deep-Sea Research 32; 
809-818, 

Labour, M,V, 1934, The eggs and larvae ot some British 
Turridae Journal of the Marine Biological .Association of 
the United Kingdom. 19; 541-554. 

Rex, M.A. 1977 Zonation in deep-sea gastropods; the im- 
portance of biological interactions to rates ot zonation. 
European Symposium on Marine Biology 11; 521-530. 

Rex, MA, 1983, Geographic patterns of species diversity in 
the deep-sea benthos In. Rowe, G.T, (ed) The sea. Vol. 
8. Wiley, N.Y. p. 453-472. 

Rex, M.A. and A. Waren. 1982. Planktotrophic development 
in deep-sea prosobranch snails from the western North 
Atlantic. Deep-Sea Research 29; 171-184. 

Rex, M.A., R.J. Etter and P.W. Nimeskern, Jr 1990. Density 
estimates for deep-sea gastropod as.semblages Deep-Sea 
Research. 37; 555-569 "" 

Rex, M.A., C.A. Van Linnnersen and R D Turner 1979 Re- 
productive pattern in the abyssal snail Rcnihoiiclla tenella 
(Jeffreys). In Stancyk, S.E. (ed). Reproducti\e ecology of 
marine invertebrates. Universil\ of South (Carolina Press, 
Columbia, p, 173-188, 

Robertson, R 1974 Marine prosobranch ga.stropods; larval 
studies and systematics, Thalassia Jugoslavica 10; 213-238, 

Shimek, R,L 1986, Biolog)' of the Northeastern Pacific Tur- 
ridae, W Demersal development, synchronous settlement 
and other aspects of the lar\al hiolog) of Ocnopota levi- 
(tcnsis International Journal of Invertebrate Beproduclioii 
and Development 10; 313-333. 

Sokal, R,R. and F J Kolill 1981. Biometry Freeman. San 
Francisco. 859 p. 

Siratlmiann, R.R. 1978. The evolution ami loss oi feeding 
larval stages of marine iiixcrtcbralcs E\(ilulion .32; 894- 
906, 

Tliiriol-(,^)uirvreu\, (^ 1972, \lnrosliiKturfs ilc roipiilles lar- 



E. Potter and M. A. Rex, 1992 Page 75 



vaires de prosobranches au microscope electronique a ba- Mcddelflser fra Konimissionen for Danmarks Fiskeri-og 

lavage. Archives de Zoologie Experimentale et General Havunders0gelser, Serie: Plankton 4, 523 p. 

1 13: 553-564 Thorson, G. 1950. Reproductive and larval ecology of marine 

Thorson, G. 1946. Reprotliiction and lar\al dtnelopment ot bottom invertebrates. Biological Reviews 25; 1-45. 

Danish marine bottom invertebrates, with special refer- Nance, RR 1973. On reproducti\e strategies in marine ben- 

ence to the planktonic larvae in The Sound (0resund) lliic- iiucrti-bratcs, .American Naturalist 107: 339-352. 



THE NAUTILUS 106(2):76, 1992 



Page 76 



Drillia macleani, New Name for Drillia sinuosa McLean and 
Poorman, 1971 (Gastropoda: Turridae) 



John K. Tucker 

Department of Biological Sciences 
Illinois State University 
Normal, IL 61761 USA 



During the preparation of a catalog of the family Tur- 
ridae, I found the name Drillia sinuosa McLean and 
Poorman, 1971, which was applied to a Recent eastern 
pacific species, to be preoccupied by Drillia sinuosa Bel- 
iardi, 1877, which was applied to an unrelated species 
from the Middle Miocene of Italy. In order to rectify 
this homonymy and to acknowledge the seminal work 
on turrids done by the senior author of the preoccupied 
taxon, Drillia macleani is here proposed as a replacement 
name for the eastern Pacific species. 

Drillia macleani Tucker, nomen novum 

Drillia sinuosa McLean and Poorman, 1971:96, fig. 16., 
McLean in Keen, 1971 :# 1622. 

Not Drillia sinuosa Bellardi, 1877:126, pl.4, figs. 18a,b. 



Remarks: The holotype is in the collection of the Los 
Angeles County Museum of Natural History (LACM 
1491). The type locality is off the southernmost coast of 
Isla Santa Cruz, Galapagos Islands, Ecuador (0°47'S, 
90°21'W), in 150 m, collected 10 June 1968 by Andre 
and Jacqueline DeRoy. 

I thank Dr. James H. McLean and an anonymous 
referee for reading the manuscript. 

LITERATURE CITED 

Bellardi, L. 1877. I molluschi dei terreni Terziarii del Pied- 
monte e della Liguria, 2. Torino. 364 p., 7 pis. 

Keen, A. M. 1971. Sea shells of tropical west America, second 
edition. Stanford University Press, Stanford, 1064 p. 

McLean, J H and R Poorman, 1971 New species of tropical 
eastern Pacific Turridae. The Veliger 14(1):89-113. 



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rHE NAUTILUS 



Volume 106, Number 3 
August 21, 1992 
ISSN 0028-1344 

A quarterly devoted 
to malacology. 



.t- 



AUG 2 C 1992 



Woods Hols, Mass. 




EDITOR-IN-CHIEF 
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Division of Mollusks 
National Museum of 
Natural History 
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Field Museum of 
Natural History 
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Department of Biology 
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Charleston, SC 29424 



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Department of Living Invertebrates 

The American Museum of Natural 

History 

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Division of Mollusks 
National Mu.seum of 
Natural History 
Smithsonian Institution 
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Division of Mollusks 
National Museum of 
Natural History 
Smithsonian Institution 
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Museum of Comparative Zoology 
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THEt^NAUTILUS 



CONTENTS 



Volume 106, Number 3 

August 21, 1992 

ISSN 0028-1344 



'larine Rioi 



IBRAR 



James F. Quinn, Jr. 



New species of Calliostoma Swainson, 1840 (Gastropoda: 

Trochidae), and notes on some poorK known species from 

the western Atlantic Ocean ^ ' ;?;'5; Hol 



r.i,n.;.. . 71 



James H. McLean 



A new species of Pseudorimula {Fissurellacea: 
Clypeosectidae) from hydrothermal vents of the Mid- 
Atlantic Ridge 



115 



Jov C. Goodsell 


Prodissoconch I and II length in Mercenaria Taxa 


119 


Arnold G. Eversole 






Harald A. Rehder 


Harpa cabriti Fischer. 1860, a replacement name for 

Harpa ventricosa Lamarck, 1816 (Gastropoda: Harpidae) 


123 


James H. McLean 


Obituary: Rae Baxter, 1929-91 


125 



THE NAUTILUS 106(3):77-114, 1992 



Page 77 



New Species of CaUiostoma Swainson, 1840 
(Gastropoda: Trochidae), and Notes on Some 
Poorly Known Species from the 
Western Atlantic Ocean 



James F. Quinn, Jr. 

Florida Marine Research Institute 
Department of Natural Resources 
100 Eighth Avenue, S. E. 
St Petersburg, FL 33701-5095, USA 



ABSTRACT 

Twent) -seven new species of CaUiostoma Swainson {sensii lata) 
are described from the western Atlantic Ocean. Trochus (Zi- 
zyphiniis) stirophorus Watson is confirmed to be a CaUiostoma. 
with C. arestum Dall as a synonym, and Trochus (Margarita) 
dnopherus Watson is transferred to CaUiostoma New records 
and observations are presented for C. apicitium Dall, C. in- 
diana Dall, C. orion Dall, C. aurora Dall, C jernandezi Princz, 
C, atlantis Clench & Aguayo, C, axelolssoni Quinn (new name 
for C. olssoni Bayer), and C. adspersun^ (Philippi). External 
morphologies of the head-foot of 13 species are briefly de- 
scribed, and illustrations of the radulae of 13 species are pre- 
sented, along with pertinent remarks. 

Key tvurds: Trochidae, Calliostomatinae, CaUiostoma. systeni- 
atics, new species, new records, radulae 



INTRODUCTION 

The subfamily Calliostomatinae is by far the most spe- 
ciose trochid subfamily in the western Atlantic Ocean. 
Species occur from the subarctic waters of Canada to the 
subantarctic waters of southern Argentina and the Ant- 
arctic waters of South Georgia Island (Powell, 1951; 
Clench & Turner, 1960), Calliostomatines inhabit a great 
variety of habitats, both biotic and abiotic, in depths 
ranging from the shallow subtidal to se\eral hundred 
meters (Clench & Turner, 1960; Reed & Mikkelsen, 1987; 
Quinn, in press). 

The greatest number of species of Calliostomatinae is 
assigned to the rather ill-defined genus CaUiostoma 
Swainson, 1840. Clench and Turner (1960) monographed 
the western Atlantic species of the genus and concluded 
that 41 names represented valid species (including 5 new 
species), 24 names were synonyms, and 8 names were 
not referrable to the genus. Subsequent studies (Merrill, 



1970; Bayer, 1971; Princz, 1978; Quinn, 1979, in press; 
Rios, 1985, 1990) have revealed one new synonym and 
two new species, reinstated two species excluded by 
Clench and Turner, and resurrected three species from 
synonymy. Another 15 nominal species and subspecies 
not treated by Clench and Turner (1960) have been 
described from off southern Argentina (Philippi, 1845 
[in 1842-1851]; Martens, 1881; Smith. 1881, 1915; Roch- 
ebrune & Mabille, 1889; Strebel, 1905, 1908); a few of 
these species were discussed or mentioned by Powell 
(1951), who later (1960) published a complete list, but 
most of these species remain essentially unknown. In- 
cluding the 27 new species described in this paper, the 
2 here reassigned to the genus, and at least 7 that are 
currently unnamed, almost 100 valid Recent species of 
CaUiostoma are now known from the western Atlantic 
(Table 1), In contrast, only 36 species of CaUiostoma are 
known to occur in the eastern Pacific (Table 2) (see Dall, 
1909; McLean, 1970, 1971, 1984; Rehder, 1971; McLean 
& Andrade, 1982). The number of valid Recent species 
of CaUiostoma now known in the Americas is somewhat 
greater than the number of species described from the 
Neogene formations of the Americas. 

At least six other calliostomatine species occur only off 
Argentina and in the Falkland and South Georgia Islands 
(Powell, 1951, 1960). Powell assigned two species each 
to Falsimargarita Powell, 1951, and Photinastoma Pow- 
ell, 1951, and one species each to Venustatrochus Powell, 
1951, and Photiniila Adams and Adams, 1854. Clench 
and Turner (1960) assigned CaUiostoma blakei Clench 
and Aguayo, 1938, to Photitnila, but characters of the 
animal and radula show that the species should be as- 
signed to the Lirulariinae Hickman and McLean, 1990 
(Quinn, in preparation). 

During the preparation of monographs of the Tro- 
chidae of the Gulf of Me.xico (Quinn, in press) and the 



Page 78 



THE NAUTILUS, Vol. 106, No. 3 



adjoining Straits of Florida (Quinn, 1979), I examined 
specimens of many calliostomatine species collected from 
throughout the Caribbean Sea and southwestern Atlantic 
Ocean, most obtained subsequent to (blench and Turners 
1960 monograph. Many of the species represented in 
that material were clearly undescribed, some were found 
to be species complexes, and others represented signifi- 
cant range extensions of pre\iously described but poorly 
known species. The purposes of this paper are to provide 
descriptions of the new species, to clarify the identities 
of some species, and to report new distributional data 
for species of Calliostoma in the western Atlantic Ocean. 
The external morphologies of the head-foot of 13 of the 
species are briefly described, and radulae of 13 species 
are illustrated. However, because so few species of west- 
ern .\tlantic Calliostoma have had their anatomy and 
radula illustrated, and because character states are so 
poorly understood for species worldwide, speculations on 
relationships based on the present observations are pre- 
mature. Of particular interest would be optical micro- 
scope observations to clarif\ characteristics of these rad- 
ulae as espoused by Hickman (1977). 

Numerous genus-level taxa have been proposed for 
perceived species groups within Calliostomatinae. Al- 
though some authors incorporated features of the exter- 
nal anatomy and radula (e.g., Powell, 1951; Clench & 
Turner, 1960), most of these names were based on shell 
characters alone, and some names seem to reflect a re- 
gional bias of the authors; however, the morphological 
limits of these nominal taxa are nebulous, and a world- 
wide survey of genus-level taxa is needed. Therefore, the 
species discussed in the present paper are not formally 
assigned to nominal subgenera, but the following species 
groups are recognized as relatively distinct: the C. ptil- 
chrum group [Calliostoma s.s., or Eucasta Dall, 1889a); 
the C. jujulnnum group {Elmerlinia Clench & Turner, 
1960); the C. bairdii group (Kombologion Clench & Tur- 
ner, 1960); the C. sayantim group; the C. circumcinctum 
group (?=Otuhaia Ikebe, 1942); the C. atlantis group. 
Many western Atlantic species cannot be satisfactorily 
assigned to any species group, but a forthcoming revision 
of the southwestern Pacific calliostomatine species (B. A. 
Marshall, in litt.) may help clarify some of these uncer- 
tainties. 

For brevity, authors and dates of publication for spe- 
cies included in Tables 1 and 2 are not repeated in the 
Remarks section of the species accounts. Species not in- 
cluded in those tables are cited with their authors and 
dates whenever mentioned in the text. Bibliographic ref- 
erences of species included in Tables 1 and 2 are not 
included in the Literature Cited section unless the species 
are also included in the text; most of these citations are 
readily available in Clench and Turner (1960), Keen 
(1971), and Quinn (in press). 

Acronyms for specimen repositories are as follows: 
AMNH (American Museum of Natural History, New 
York); ANSP (Academy of Natural Sciences of Phila- 
delphia); BM(NH) (The Natural History Museum, Lon- 
don); DMNH (Delaware Museum of Natural History, 



Wilmington); FSBC I (Invertebrate Collection, Florida 
Marine Research Institute, St. Petersburg); HMNS (Hous- 
ton Museum of Natural History, Texas); LACM (Natural 
History Museum of Los Angeles County, California); 
MCZ (Museum of Comparative Zoology, Harvard Uni- 
versity, Cambridge, Massachusetts); MORG (Museu 
Oceanographico da Fundafao L'niversidade do Rio 
Grande, Rio Grande, Rio Grande do Sul, Brazil); UF 
(Florida Museum of Natural History, University of Flor- 
ida, Gainesville); UMML (Rosenstiel School of Marine 
and Atmospheric Science, University of Miami, Florida); 
USNM (National Museum of Natural History, Smithson- 
ian Institution, Washington, DC). 



SYSTEMATICS 

Superfamily Trochoidea Rafinesque, 1815 

Family Trochidae Rafinesque, 1815 

Subfamily Calliostomatinae Thiele, 1924 

Genus Calliostoma Swainson, 1840 

Type species (subsequent designation; Herrmannsen, 

1846:154): Trochus conulns Linnaeus, 1758. 

Calliostoma hilare new species 
(figures 1, 2) 

Description: Shell medium-sized, attaining height of 
14.5 mm, width of 11.5 mm. conical, nonumbilicate, 
finely sculptured. Protoconch 375 nm maximum diam- 
eter, 1 whorl. Teleoconch whorls 8.6, flat-sided; first w horl 
with weak axial riblets and 2 strong spiral cords; axial 
riblets disappearing on second whorl; spiral sculpture 
increasing by intercalation to 7 cords and 4 threads; 
supraperipheral spiral cords finely beaded, beads crowd- 
ed, rounded; periphery of first 4 whorls sharply carinate, 
composed of abapical spiral cord set w ith sharpK conical 
beads, periphery of subsequent w horls narrowK round- 
ed, composed of two adpressed spiral cords set with 
rounded beads. Base weakly convex, with 15 narrow, 
finely beaded spiral cords, cords becoming broader and 
more coarseK beaded adaxialK . Aperture subquadrate. 
lips thin; columella almost straight, thickened, slightly 
twisted, terminating in weak, rounded denticle. Shell 
ground color golden yellow with greenish iridescence, 
with light golden brown and white spots alternating along 
periphery; w hite spots spiralK elongate, brow n spots ax- 
ialK elongate. 

Holotype: USNM 860249, height 14.5 mm. width 11.5 
tnm. 

Type locality: Off northwestern tip of Little Bahama 
Bank, John Elliott Pillsbury Station P-198, 27°30'N, 
79°10'W, in 242 m. 

Distribution: This species is known onl\ from the ho- 
lotype, an empt\- shell, collected in 242 m off the north- 
western tip of the Little Bahama Bank. 

Remarks: The holotype of C hilare is most similar to 



J. F. Quinn, Jr., 1992 



Page 79 



Table 1. Cieograpliif anil hatliynietric distributions of calliostoinatine species in the western Atlantic Ocean. 

Species 

C'alliostoma Swainson, 1840 
jujulnnuui (CInielin, 1791) 
javanicnm (Lamarck, 1822) 
occidentale (Mighels & A. Adams, 1842) 
gemmosum (Reeve, 1842) 
nudum nudum (Philippi, 184.5) 
tampaense (C^onrad, 1846) 
jucunduni (Gould, 1849) 
pulchriim (C B Adams. 18.50) 
adspcrsum (Pliilippi, 1851) 
euglyptuni (A, Adams, 18.55) 
dnophcrum (Watson, 1879) 
stirophorum (Watson, 1879) 
coppingeri Smith, 1880 
bairdii Verriil & Smith, 1880 
apicinum Dall, 1881 
circumcinctum Dall, 1881 
echinatum Dall, 1881 
orion Dall, 1881 
roscoluni Dall. 1881 
sapidum Dall, 1.881 
ijucatccanum Dall, 1881 
consimilis (Smith, 1881) 
aurora Dall, 1888 
benedicti Dall, 1889 
briinneum (Dall, 1889) 
cinctellum Dall, 1889 
Indiana Dall, 1889 
psyche Dall, 1889 
sayanum Dall, 1889 
dozei Rochebrime & Mabille, 1889 
nudum roseotinclum Rochebrune & Mabille, 1889 
optimum Rochebrune & Mabille, 1889 
senilis Rochebrune & Mabille, 1889 
irisans Strebel, 1905 
kophamcli Strebel, 1905 
mocbiusi Strebel, 1905 
nudum flaviducarnea Strebel, 1905 
marionae Dall, 190(5 
militare Ihering, 1907 
andersoni Strebel, 1908 
falklandicum Strebel, 1908 
modestulum Strebel, 1908 
nordenskjoldi Strebel, 1908 
venustulum Strebel, 1908 
depiclum Dall, 1927 
hahbrectum Dall, 1927 
hendersoni Dall, 1927 
sarcodum Dall, 1927 
amazonicum Finlay, 19.30 
bigelowi Clench & Aguayo, 1938 
schroederi Clench & Aguayo, 1938 
hassler Clench & Aguayo, 19.39 
atlantis Clench & Aguayo, 1940 
carcellesi Clench & .Aguayo, 1940 
cubanum Clench & Aguayo, 1940 
torrei Clench & Aguayo, 1940 
fascinans Schwengel & McGinty, 1942 
barbouri Clench & Aguayo, 1946 
adelae Schwengel, 1951 
bullisi Clench & Turner, 1960 



)eptli (m) 


Distribution 


0-10 


4 


0-40 


4 


.30-1,800 


1,2- 


10-100 


6 


? 


7 


0-10 


3- 


20-30 


7 


0-70 


3, 4, 5, ?6 


0-40 


5,6 


10-30 


3 


640 


6* 


700 


3-,4- 


10-80 


6,7 


70-450 


2 


100-200 


4- 


200-300 


4- 


70-1.50 


4 


20-60 


4 


40-80 


3 


100-250 


?3-, 4 


1.5-60 


3 


? 


7 


180-1,0.50 


4 


365 


3-* 


20-70 


4 


?-300 


3-, 4* 


.50-100 


4 


100-400 


3- 


150-300 


3- 


? 


7 


? 


7 


? 


7 


? 


7 


? 


7 


? 


7 


90 


7 


? 


7 


40-120 


3 


30-65 


6,7 


? 


7 


1.5-2.50 


7 


65-350 


7 


? 


7 


40 


7 


1-7 


6 


535 


3* 


150-3.50 


3- 


1-10 


4 


■? 


7 


,375-4.30 


4* 


250-450 


4- 


65 


6* 


?-600 


4- 


50 


7* 


900 


4* 


700 


4* 


70-120 


3 


25-100 


3-, 4 


0-2 


3- 


70 


5* 



Page 80 



THE NAUTILUS, Vol. 106, No. 3 



Table 1. ('ontiiiiied. 



Species 



Depth (m) 



Distributii 



jeanneae Clench & Turner, 1960 
Oregon Cleiieh & Turner, 1960 
roseicateri ( blench & Turner, 1960 
springvri Clench & Turner, 1960 
fcrnandezi Princz, 1978 
alternttm Quinn, 1992 
argcnttim Quinn, 1992 
allantoides Quinn, 1992 
(lulinim Quinn, 1992 
axclolssoni Quinn, 1992 
hertmulcnsc Quinn, 1992 
hruniicophttan Quinn, 1992 
ciiithjphiltini Quinn, 1992 
coronatnm Quinn, 1992 
cuhense Quinn, 1992 
defnle Quinn, 1992 
dentatum Quinn, 1992 
fucosum Quinn, 1992 
guesti Quinn, 1992 
hihirc Quinn, 1992 
hirttini Quinn, 1992 
moscatcUii Quinn, 1992 
ptirpureurn Quinn, 1992 
rota Quinn, 1992 
rude Quinn, 1992 
rugosum Quinn, 1992 
scalenum Quinn, 1992 
scurru Quiim, 1992 
senii.sudvr Quinn, 1992 
serratnhim Quinn, 1992 
tenebrosum Quinn, 1992 
vinosum Quinn, 1992 
oiscardii Quinn, 1992 

Venitstatrochus Powell, 1951 

georgianus Powell, 1951 
Falsimargarita Powell, 1951 

gemma (Smith, 1915) 

iris (Smith, 1915) 

Photinula Adams & Adams, 185-1 
coeridescens (Kiufi, & Broderip, 1831) 

Photinaatoma Powell, 1951 
tacniata taeniata (Wood, 1828) 
taeniata nivea (Cooper & Preston, 1910) 
'^gamma (Rorhehrune & Mahiile. 1889) 



200-350 
274-641 
200-450 

4.5-145 

5.5-146 
350-450 
400-600 

10-50 
200-300 

82 

50-80 

150-350 

770-800 

0-20 

? 

15-55 
5-9 
183-219 
250 
520 
50-80 
0-60 
27 

60-90 
? 

25-80 

20-90 

70-155 

120-190 

? 

18-22 
40-45 

120-200 

200-400 
225-450 

0-200 



0-80 
0-100 

? 



4* 

3- 

4- 

3- 

5 

5 

4 

4* 

5 

4-,6- 

3* 

6* 

3 

6* 

4- 

4* 

3- 

5* 

4* 

4* 

4* 

6 

5- 

6* 

5- 

4* 

3 

5 

5- 

5- 

6* 

6 

6* 



* Known only from the type lot or locality; 1 = northern boreal; 2 = "Virginian" (Cape Cod to Cape Hatteras); 3 = Carolinian; 
4 = Caribbean; 5 = northern ,South America; 6 = Brazil-northern Argentina; 7 = Patagonian; 8 = Subantarctic; '— ' indicates 
restricted range within region. 



shells of C. rude in shape, size, and sculpture, but it 
differs by having a slightly larger protoconch (375 ^m 
Ds. 350 nm); by having the ()eriplierul spiral cord strong 
and sharply beaded on whorls 2-4, becoming weaker 
and with rounded beads on later whorls, whereas the 
peripheral cord of C. rude is inconspicuous on the first 
three whorls, becoming very strong and coarsciv beaded 
on subsequent whorls; and by having more numerous 



but weaker spiral cords. Calliostoma hilarc is also similar 
to C. serratulum, but the shell of the former has a smaller 
protoconch (375 nm vs. 400-425 ^m), is proportioiiateK- 
narrower, has more numerous spiral cords that are more 
weakly beaded, and has a periphery that is composed of 
two subequal, finely beaded, adpressed cords rather than 
a strong, coarsely beaded upper cord and a weaker lower 
cord. 



J. F. Quinn, Jr., 1992 



Page 81 



Table 2. (ieograpliic ami balln metric distrihiitions of calliostomatine species in the eastern Pacific Ocean. 



Species 



Depth (m) 



Distrilmtiiin 



Calliostoma Swainson, 1840 
anniildtum (Liglitfoot. ITcSfi) 
caridlUulatum (Lightfoot, 1786) 
anlonii (Koch, 1843) 
exiniium (Reeve, 1843) 
ligatum (Gould. 1849) 
leatmm (C. B. Adams, 1852) 
fonkii (Philippi, 1860) 
gemrnulalum Carpenter, 1864 
splcndcns Carpenter, 1864 
supragranosum Carpenter, 1864 
varirgatum C^arpenter, 1864 
aequisculptnm Carpenter, 1865 
triculor Gahh, 1865 
gloriosum Dall, 1871 
palmeri Dall, 1871 
platinum Dall, 1890 
tiirhiuum Dall, 1895 
iridium Dall, 1896 
nephcloidc Dall, 1913 
honita Strong, Hanna. & Hertlein, 1933 
rema Strong, Hanna, & Flertlein. 19.33 
marshalli Lowe, 1935 
mcleani Shasky & Campbell, 1964 
gordanum McLean, 1970 
jacquelinae McLean, 1970 
keenae McLean, 1970 
sanjaimense McLean, 1970 
santacruzanum McLean, 1970 
veleruae McLean, 1970 
insignis Olsson, 1971 
joanneae Olsson, 1971 
pillsburyae Olsson, 1971 
chilena Rehder. 1971 
delli McLean & Andrade, 1982 
bernardi McLean, 1984 
titanium McLean, 1984 



"offshore 

25-750 

0-3 

0-40 

0-3 

0-3 

?-457 

0-3 

"offshore ' 

0-3 

25-750 

1-3 
15-60 
0-40 
0-45 
80-750 
60-120 
230-280 
50-125 
35-75 
20-45 

.5-10 
128 
146 

.5.5-110 
137 
45 

60-100 
50-80 
53 

57-64 
200-7.50 
200-4.50 
12.5-175 
200-300 



offshore' 



1 

1 

3 

3 

1 

3 

3,4 

2 

2 

2 

1 

3 

2,3 

2 

3 

1 

2 

3 

3 

3 

3 

3 

3 

2* 

3* 

2,3 

2* 

3* 

3 

3 

3* 

3* 

3,4 

4- 



* Known link from the t\pe lot or locality; 
restricted range within region. 



rthern boreal; 2 = Californian; 3 = Panamic; 4 = Peruvian; 



indicates 



Calliostoma serratulum new species 
(figures 3-6) 

Calliostoma sp. 4: Bayer et al. 1970;A29, A137, A146. 

Description: Shell medium-sized, attaining height of 
16.7 iTim, width of 16,4 mm, conical, nonumbilicate, 
coarsely sculptured. Protoconch 400-42.5 p.m ma.ximum 
diameter, 1 whorl. Teleoconch whorls about 7, flat; first 
2 whorls with low, rounded axial riblets and 2 strong, 
beaded spiral cords; a.\ial riblets gradualK disappearing 
on whorls 3-5; spiral sculpture increasing by interca- 
lation to 5-8 cords and 0-5 threads, of which abapical 
2 cords forming periphery; periphery sharply carinate; 
adapical peripheral cord strong, projecting, set with 
strong, conical beads, beads crossed by 2 fine spiral threads 
producing squarish apices; abapical peripheral cord 
weaker, weakly beaded. Base weakly convex, with 13- 
17 narrow, weakly beaded spiral cords, adaxial 2-3 cords 



stronger, more widely spaced, and more distinctly bead- 
ed. Aperture subquadrate, lips thin, weakly crenulate; 
columella concave above, straight below , thickened, ter- 
minating in weak, rounded denticle. Shell ground color 
golden yellow with spots and axial streaks of brown; 
beads white; basal spiral cords with elongate spots of 
brown, darkest on adaxial 2-3 cords. 

Holotype: USNM 860250, height 12.5-F mm, width 
12.3 mm. 

Type locality: Southw est of Cartagena, Colombia, John 
Elliott Pillsbiry Station P-375, 9°59.0'N, 76°02.0'W, 
in 135-130 m. 

Paratypes: 1, UMML 30.3539; off Santa Marta, Colom- 
bia, John Elliott Pillsbiry Station P-399, 9°01.3'N, 
76°40.2'W, 119-179 m; 17 Julv 1966; 10-ft otter trawl— 
1, UMML 30.6935; 1, FSBC'l 44069; off Peninsula de 



Page 82 



THE NAUTILUS. Vol. 106, No. 3 





.A 



/ 



ji 



"\ 



S, 



V 



/ 




■•'1 



v. 



■k_ 



^' 






8 





N 




10 



Figures 1, 2. Calliostotna hilare new species, liolotype, I'SNM 860249; height 14 3 mm, width 1 1.5 mm Figures 3-6. Calliostoma 
serratuhim new species, 3, i. Holotype, USNM 860250; height 12.5+ mm, width 12, .i mm, 5. 6. Parat\ pe, L'MML 30,35.39; height 
10.1 mm, width 100 mm. Figures 7, 8. Calliostoma rude new species, ho]()t>pe, L'SNM 860251; height 13.3 mm, width 10 6 mm. 
Figures 9. 10. Calliostoma sp, cf. rude, specimen Irom off Barbados, Sunderland collection; height 9,7 mm, width 7.3 mm. 



Paraguana, Venezuela, John Elliott Pillsbi ry Station 
P-757, 11°39.6'N, 69°22.1'W, 161-187 m; 27 July 1968; 
10-ft otter trawl. 

Other material: 4 broken, UMML 30.8366; off Santa 
Marta, Colombia, John Elliott Pillsbury Station 
P-785, 11°16.9'N, 74°17.0'\V, 176-165 m; 31 July 1968; 
10-ft otter trawl. 



Distribution: Calliostoma serratuhim is known only 
from empty shells traw let! from off the nortiieastern coast 
of Colombia and northwestern coast of Venezuela in 1 19- 
187 m. 

Remarks: Shells of C. serratuhim are most similar to 
those of C. rude but differ by having a much larger 
protoconch (400-425 nm vs. 350 fim), by being propor- 



J. F. Quinn, Jr., 1992 



Page 83 



tionately wider, and b\ having the adapical peripheral 
cord set with uniformly sized beads that are squarish in 
profile. 

Calliostoma rude new species 
(figures 7, 8) 

Description: Shell medium-sized, attaining height of 
13.3 mm, width of 10.6 mm, conical, nonumbilicate, 
coarsely sculptured. Protoconch 350 ixm maximum di- 
ameter, 1 whorl. Teleoconch whorls 8.3, flat; first w horl 
with low, rounded axial riblets and 2 strong, beaded spiral 
cords; axial sculpture absent after first w horl; spiral sculp- 
ture increasing by intercalation to 5-6 cords and 1-2 
threads, of which abapical 2 cords forming periphery; 
periphery sharpK carinate; adapical peripheral cord 
strong, strongly beaded, beads triangular, every third or 
fourth bead markedK- larger; abapical peripheral cord 
weaker, strongK undulate. Base weakly convex, with 11- 
13 strong, narrow, fineK beaded spiral cords .-Aperture 
subcjuadrate, lips thin, fineK crenulate; columella almost 
straight, thickened, terminating in weak, rounded den- 
ticle. Shell ground color ivor\ w ith greenish and reddish 
iridescence; streaks and flammules of golden brown pres- 
ent above periphery; periphery with spots of darker 
brow n; base with light goklen-brown spots arranged on 
cords to form obscure, crescentic radial streaks. 

Holoiype: USNM 860251, height 13.3 mm, width 10.6 
mm 

Type locality: Off Cayenne, French Guiana, John El- 
liott PiLLSBURY Station P-650, 6°07.0'N, 52°19.0'W, in 
84-91 m. 

Paratypes: 2, UMML 30.5844; 1, FSBC I 44076; from 
same lot as holotype. — 2, MCZ 302593; off Georgetow n, 
Guyana, Chain Cruise 35, Station 35 + 36, 8°10.5'- 
8°lb.0'N, 57°48'W, 53-60 m; 28 .\pril 1963. 

Distribution: Calliostoma rude is known only from two 
collections, one off Georgetown, Guyana, in 53-60 m, 
and the other from the type localit} off Cayenne, French 
Guiana, in 84-91 m; the latter collection contained a 
single live-collected specimen. The field notes on the type 
localit) cite shell rubble, and the associated invertebrates 
suggest that hard-bottom outcrops are nearby. 

Remarks: Comparisons of shells of Calliostoma rude 
with those of the similar C. serratuhim and C. hilare 
are discussed in the remarks for the latter two species. 
The adapical peripheral cord of shells of C rude is sub- 
equal to the other spiral cords on the first three whorls, 
but it becomes markedly stronger on the fourth and 
subsequent whorls. This characteristic separates C. rude 
from all other peripherally carinate species in which 
peripheral carination is present on the first w horls and 
either remains strong on all whorls (e.g., C. cincteltum) 
or disappears on the last one or two whorls (e.g., C. 
echinatum). The alternation of two or three small beads 
with a much larger triangular bead is also unique to C. 
rude. Four specimens collected from off the western coast 



of Barbados (Figures 9, 10; Sunderland collection, ex F 
Sander) are very similar to the types of C, rude but are 
relativeK smaller and narrower and have some subtle 
sculptural differences that suggest that the Barbados shells 
may be a separate species. 

Calliostoma cnidophilum new species 
(figures 11-14, 107) 

Description: Shell medium-sized, attaining height of 
14.1 nmi, width of 12.1 mm, conical, nonumbilicate, 
finely sculptured. Protoconch 375 /um maximum diam- 
eter, 1 whorl. Teleoconch whorls about 8, flat; first whorl 
with low, rounded axial riblets and 2 strong, beaded spiral 
cords; axial riblets disappearing on second whorl; spiral 
sculpture increasing b\ intercalation to 10-12 cords and 
0-2 threads, cords fineK beaded; peripher\ of first 2-2.5 
w horls sharpK carinate, set with sharply conical beads; 
periphery becoming narrowly rounded on subsequent 
whorls, beads becoming finer, rounded; peripheral cord 
splitting into 2 subequal, adpressed cords on fifth w horl; 
periphery obscureK multiangulate when viewed from 
base. Base weakly convex, with 14-20 narrow, sharp, 
very finely beaded spiral cords. Aperture subquadrate, 
lips thin, fineK' crenulate; columella short, weakly con- 
cave, thickened. Shell ground color golden yellow with 
greenish iridescence; periphery with regularly spaced, 
spirally elongate pinkish spots. Animal (in alcohol) white; 
cephalic tentacles long, slender, gradualK tapering, right 
longer than left, ocular peduncles long, slender, with 
large, black eye at tips; epipodium with 4 pairs of ten- 
tacles decreasing in size posteriorly, neck lobes well-de- 
veloped, thin, semicircular, smooth. 

Holotype: USNM 860265, height 13.7 mm, width 11.4 
mm. 

Type locality: Off Ilets-a-Goyaves, Guadeloupe, Lesser 
.\ntilles, Johnson-Sea-Link II Station JSL-lI-1845, 
16°10'00"N, 61°49'08"W, in 306 m. 

Paratypes: 2, UMML 30.8370; 1, FSBC 1 44070; off 
Dominica., John Elliott Pillsbury Station P-931, 
15°31.2'N, 6ri2.3'W, 174-357 m; 15 July 1969; 5-ft 
Blake trawl. 

Other material: 1, Sunderland collection; off St. James, 
Barbados, 152 m; ex F. Sander. 

Distribution: Calliostoma cnidophilum is known from 
the Lesser Antilles in depths of 152-357 m; living spec- 
imens were collected in 306 m and 174-357 m. 

Remarks: Shells of Calliostoma cnidophdum are most 
similar to the holotype of C. hdare but differ by being 
proportionately broader; by having an obscurely mul- 
tiangulate peripheral profile when viewed from the base; 
by having finer, more numerous spiral cords above the 
periphery; by having the spiral cords more finely beaded; 
and by having a shorter, thicker columella. The radula 
(formula 00.5.1. 5. oo; figure 107) differs from that illus- 
trated by Clench and Turner for C. pulchrum (1960:pl. 



Page 84 



THE NAUTILUS, Vol. 106, No. 3 









16 





19 







J. F. Quinn, Jr.. 1992 



Page 85 



3, fig. 3) by having the rachidian teeth with much broader 
bases and cusps and b\' having narrower cusps on the 
lateral teeth. 

Observations made from the submersible 
Johnson-Se.\-Link II indicate that C. cnidophiliim may 
feed on octocorals (M. G. Harasewych. personal com- 
munication; see also Harasewych ct al.. 1992. fig. 12). 

Calliostoma semisuave new species 
(figures 15. 16) 

Description: Shell medium-sized, attaining height of 
13.4 mm. width of 11.4 mm. conical, nonumbilicate. 
finely sculptured. Protoconch 400 ^"1 maximum diam- 
eter. 1 whorl. Teleoconch whorls 8.5. flat; first 4 whorls 
with low, rounded axial riblets and 2-4 strong beaded 
spiral cords; axial sculpture disappearing on fifth whorl; 
spiral sculpture increasing b\ intercalation to 11 cords 
and 2-3 threads, cords strongly beaded on first 5 whorls, 
beads gradually weakening on subsequent whorls; pe- 
riphery sharply carinate. with single strong, smooth, pe- 
ripheral cord. Base weakly con\ex, with 16-18 narrow, 
smooth to ver\ weaklv beaded spiral cords. Aperture 
subquadrate, lips thin; columella almost straight, thick- 
ened, terminating in sharp denticle. Shell ground color 
cream to light golden \ ellow , with diffuse spots and flam- 
mules of light golden brown; peripher\ with distinct, 
squarish spots of orange-brown. 

Holotype: USNM 860252, height 13.4 mm, width 11.4 
mm. 

Type locality: West of Isla La Tortuga, N'enezuela, John 
Elliott Pillsbury Station P-7o6, 10°57 0'\. 65°52.0'W. 
in 69-155 m. 

Paratype: 1. UMML 30.5915; off Pointe Isere. French 
Guiana. John Elliott Pillsbiry Station P-65S. 7°10.0'N. 
53°36.0'W. in 135-126 m; 9 July 1968; 10-ft otter trawl. 

Distribution: Calliostoma semisuave is known from dead 
shells collected off eastern X'enezuela and northwestern 
French Guiana in depths of 69-155 m. 

Remarks: The shells of Calliostoma semisuave are most 
similar to those of C. cnidophiliim but are proportion- 
ately narrower, have a larger protoconch. have the bead- 
ing of the spiral cords becoming very weak on the last 
several whorls, and have a smooth peripheral cord on 
the last several whorls. 

Other organisms listed on the field data sheets for the 
two collections of C. semisuave include sponges, alcyo- 
narians. Millepora. Haliotis, and chitons, suggesting that 
this species li\es on or near hard-bottom outcrops in 
depths of about 70-150 m. 



Calliostoma hrunneopictum new species 

(figures 17-20) 

Description: Shell small, attaining height of 8.5 mm, 
w idth of 6.9 mm. conical, narrowly umbilicate or non- 
umbilicate. finely sculptured, Protoconch 365-375 ^i.va 
maximum diameter. 1 whorl. Teleoconch whorls 6.7. flat 
to weakly convex; first whorl with low, rounded axial 
riblets and 2 strong, beaded spiral cords; axial sculpture 
absent after first whorl; spiral sculpture increasing b%' 
intercalation to 8-12 cords and 1-3 threads, cords finely 
beaded, beads conical; peripher\ rather sharp on first 5 
whorls, becoming narrowly rounded on last 2 whorls. 
Base weakly convex. v\ith 12-16 narrow, fineh' beaded 
spiral cords, adaxial 2-3 cords slightly stronger and more 
w idel\ spaced. Umbilicus ver\ narrow, chink-like, par- 
tialK or completeK filled by columella. Aperture 
subquadrate to subovate. lips thin, finely crenulate; col- 
umella concave, thickened, terminating in obscure, 
rounded denticle. Shell ground color ivory with greenish 
and reddish iridescence, irregular spots and flammules 
of brown above periphery, squarish spots of brown reg- 
ularly spaced on peripher\ . and obscure, crescentic streaks 
of brown on base. 

Holotype: MORG 29.291. height 8.5 mm, width 6.9 
mm. 

Type locality: Off Santana Island. Estado de Rio de 
Janeiro. Brazil, in 50-80 m. 

Paratypes: 1, MCZ 302592; 1. FSBC I 44068; from same 
lot as holot\pe. 

Other material: 2. Coltro collection; from same lot as 
holot\pe. — 1. Sunderland collection; off Rio de Janeiro. 
Brazil, 50-70 m; trawled by shrimpers. 

Distribution: At present. Calliostoma hrunneopictum 
is know n onl\ from off Rio de Janeiro. Brazil, in depths 
of 50-80 m. Living specimens are also known from 50- 
80 m. 

Remarks: Shells of Calliostoma hrunneopictum are most 
similar to those of C. roseolum but differ by having a 
larger protoconch (365-375 (um r.s. 325 nm); b\ having 
more numerous, more finely beaded spiral cords; b\ hav- 
ing an umbilical chink; and by having crescentic streaks 
of brown on the base. Differences between C. hrunneo- 
pictum and the similar C. viscardii are discussed in the 
Remarks section of the latter species. 



Calliostoma viscardii new species 
(figures 21. 22. 108) 



Figures 11-14. Calliostoma cnidophilum new species. 11, 12. Holotype, USNM 860265; height 13,7 mm, width 11.4 mm 13, 
14. Specimen from off Barbados, Sunderland collection; height 11,2 mm, width 9,2 mm. Figures 15, 16. Calliostoma semisuave 
new species. holot\pe. USNM 860252; height 13,4 mm, width 11,4 mm. Figures 16-20. Calliostoma hrunneopictum new species. 
17, 18. Holotype, MORG 29 291; height 8.5 mm. width 6.9 mm. 19. 20. Paratype, FSBC I 44068; height 7.5 mm, width 6 mm 
Figures 21. 22. Calliostoma viscardii new species, holotype. MORG 29,292; height 10,9 mm. width 8,2 mm. 



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THE NAUTILUS, Vol. 106, No. 3 



Description: Shell small to medium-sized, attaining 
height of 13.0 mm, width of 10.8 mm, conical, non- 
umbilicate, finely sculptured. Protoconch 300-335 ^m 
maximum diameter, 1 whorl, tip violet. Teleoconch 
whorls 8.25, flat to weakly convex; first whorl with 2 
strong, smooth to weakly beaded spiral cords, initially 
lacking axial sculpture but developing low, rounded rib- 
lets near end of whorl, riblets persisting on subsequent 
whorls as low folds between spiral cords; spiral sculpture 
increasing b\- intercalation to 7-8 cords and 2-6 threads, 
of which abapical 2 spiral cords forming periphery, cords 
rather coarsely beaded, beads rounded, slightly spirally 
elongate on last 2 whorls; periphery rather sharp on first 
5 whorls, becoming narrowly rounded on last 2 whorls, 
peripheral cords adpressed. Base weakly convex, with 
13-16 narrow, weakly beaded spiral cords. Aperture 
subquadrate to subovate, lips thin, finely crenulate; col- 
umella concave, thickened, terminating in obscure, 
rounded denticle. Shell ground color tan with greenish 
and reddish iridescence, patches and flammules of brown 
and occasional white spots on and above periphery, and 
elongate spots of brown on basal cords. Animal (in al- 
cohol): foot dark brown with white papillae; mantle edge 
with irregular band of brown, mantle mottled black and 
white posteriorly; head mottled black and white; cephalic 
tentacles short, stout, tapering rapidK to narrow tip; eye- 
stalks short (but about 40% tentacle length), stout, with 
large, black eye at tips; snout mottled with brown and 
white, longer than broad, slightly longer than cephalic 
tentacles, slightly tapered at tip, tip with fringe of fine, 
rather long papillae; epipodium with 4 pairs of short, 
stout tentacles; left neck lobe seems to be smooth, right 
lobe finely digitate. 

Holotype: MORG 29.292, height 10.9 mm, width 8.2 
mm. 

Type locality: Between Ihla de Sao Sebastiao and Ihla 
de Buzias, off Sao Paulo, Estado de Sao Paulo, Brazil, in 
40-45 m. 

Paratypes: 1, USNM 860253; 1, FSBC 1 44066; from 
same lot as holotype. 

Other material: 3, Coltro collection; off Guarapari, Es- 
tado de Espirito Santo, Brazil, 20-30 m; trawl; July 
1991. — 6, Coltro collection; off Ihla de Sao Sebastiao, 
Estado de Sao Paulo, Brazil, 30-35 m; dredge; J. & M. 
Coltro & L. F. Viscardi, collectors. 

Distribution: Calliostoma viscardii is known from 
southeastern Brazil (Guarapari to Sao Paulo) in depths 
of 20-45 m; living specimens have been collected in 
depths of 30-45 m. 

Remarks: Shells of this species resemble those of Cal- 
liostoma brunneopictiim but differ by having a smaller 
protoconch (300-335 nm vs. 365-375 ^m) with a violet 
tip; by having stronger, more widely spaced spiral cords 
whose beads are large and rounded rather than small 
and conical; by having the whorl periphery composed 
of a pair of close-set spiral cords abapical to the cord 



that is the continuation of the abapical cord on the first 
whorl, whereas the abapical cord on the first whorl of 
C. brunneopictiim forms the periphery on all whorls; 
and b\ having less well-defined beading on the basal 
spiral cords. Calliostoma viscardii is also similar to C. 
roseolum, but the shells have a larger protoconch (335- 
350 (um vs. 300-335 fim); lack the prominent, broadly 
rounded whorl periphery; have stronger, more widely 
spaced spiral cords above the periphery; and have slightly 
larger but less discrete beading. The radula of C. viscardii 
(formula oo.5.1.5.oo; figure 108) does not differ signifi- 
cantl)' from that of C. piilchrum illustrated b\ Clench 
and turner (1960:pl. 3, fig. 3). 

This species is named viscardii to recognize the con- 
tributions of Sr. Luiz Francisco Viscardi to Brazilian mal- 
acology. 

Calliostoma bermiidense new species 

(figures 23, 24) 

Description: Shell small to medium-sized, attaining 
height of 12.6 mm, width of 8.9 mm, conical, nonum- 
bilicate, finely sculptured. Protoconch 315-325 ^m max- 
imum diameter, 1 whorl. Teleoconch whorls about 8, flat 
to weakly concave; first whorl with low, rounded axial 
riblets and 2 strong, beaded spiral cords; axial sculpture 
weakening rapidly at end of second whorl, absent there- 
after; spiral sculpture increasing by intercalation to 8- 
10 cords and 1-4 threads, of which abapical 2-3 cords 
forming periphery, cords strongly beaded; periphery of 
first 3-4 whorls sharply angulate, that of subsequent 
whorls broadly rounded, not distinctly set off from base. 
Base flat to weakly convex, with 10-14 narrow, weakly 
beaded spiral cords of which 1-3 markedK weaker. .Ap- 
erture subquadrate to ovate, thickened within, lips thin, 
crenulate; columella short, almost straight, thickened. 
Shell ground color yellowish brown, spiral cords brown, 
often broken into dashed lines on base, periphery often 
with diffuse patches of brown to red-brown, apical 3-4 
whorls pinkish brown. 

Holotype: DMNH 187591, height 11.9 mm, width 8.9 



Type locality: Off Castle Roads, Bermuda, in 82 m. 

Paratypes: 9, DMNH 187592; 2, USNM 860269; 1, FSBC 
I 42541; all from same lot as holot\pe. 

Distribution: Calliostoma bermiidense is known only 
from the type localit> ; no living specimens are known. 

Remarks: Shells of Calliostoma bermiidense are most 
similar to those of C. roseolum but differ by being rel- 
atively broader; by having narrower, more widely spaced 
and more finely beaded spiral cords; b\- having a flatter 
whorl profile after the fourth whorl; b\ having a pro- 
portionateK- shorter and thicker columella; b\ having 
brown or pinkish-brown apical whorls; and by having 
brown spiral cords. 



J. F. Quinn, Jr., 1992 



Page 87 



Calliostoma deniatum new species 
(figures 25, 26) 

Description: Shell small, attaining height of 8.4 mm, 
width of 7.4 mm, conical, nonumbilieate, finely sculp- 
tured. Protoconch 335-350 /jm ma.ximum diameter. 1 
whorl. Teleoconch whorls 8, flat to weakly concave; first 
2 whorls with low, rounded axial riblets and 2 strong, 
beaded spiral cords; axial sculpture on remaining whorls 
of low, rounded, discontinuous axial threads, most dis- 
tinct at peripher\; spiral sculpture increasing by inter- 
calation to 4-8 strongly beaded cords, of which abapical 
2-3 spiral cords forming periphery; periphery angulate, 
adapical peripheral cord strongest, abapical 2 (when 3 
present) weaker, closeK' appressed, set with axially elon- 
gate beads. Base flat, with 10-14 narrow, finely beaded 
spiral cords. Aperture subquadrate, lips thin, crenulate; 
columella straight, thickened. Shell ground color ivory 
with diffuse axial flammules of golden brown above pe- 
riphery, and 2-3 spiral row s of discrete, spirally elongate 
spots of golden or reddish brown on base. 

Holotype: USNM 859358, height 5.7 mm, width 5.2 
mm. 

Type locality: Off Freeport, Texas, approximately 
28°05'N, 94°35'W, in 51 m. 

Paratypes: 1, MCZ 297052; 1, LACM 2293; 1, HMNS 
3946; N\V Gulf Survey station, off Freeport, Texas, 51 
m; trawl; A. Kight, collector.— 1, HMNS 15866; 2, FSBC 
I 32301; XW Gulf Survey station, 2 mi south of #1 buoy, 
Heald Bank, 22 m; 22 August 1966; H. Geis and W. 
Pierce, collectors. Other paratypes and material (44 lots) 
are listed by Quinn (in press). 

Distribution: Calliostoma dentatum is confined to the 
northwestern Gulf of Mexico, from the Mississippi River 
to off Port Isabel, Texas, and is usually collected from 
depths of 15-55 m; living specimens are known from 
depths of 12-51 m. 



Turner (1960:pl. 3, fig. 3) by having four rather than five 
lateral teeth. 

Calliostoma tenebrosum new species 
(figures 27, 28) 

Description: Shell medium-sized, attaining height of 
13.6 mm. width of 11.5 mm, conical, nonumbilieate, 
finely sculptured. Protoconch slightly chipped, about 325 
;um maximum diameter, 1 whorl. Teleoconch whorls 7.9, 
flat; first 3 whorls with low, rounded axial riblets and 2- 
4 strong, beaded spiral cords; axial riblets weakening on 
fourth whorl, becoming weak, irregular folds on subse- 
quent whorls; spiral sculpture increasing by intercalation 
to 7 cords and 1 thread, of which abapical 3 cords form- 
ing periphery, beads on cords strong, rounded; periphery 
rather broadly rounded, adapical 2 peripheral cords 
strongest, subequal, separated by rather broad groove. 
Base very weakly convex, with 13 narrow, finely beaded 
spiral cords. Aperture subquadrate, moderately thick- 
ened within, lips thin, crenulate; columella very weakly 
concave, thickened, terminating in rounded tubercle. 
Shell ground color tan with spots of white and spots and 
axial streaks of dark brown. 

Holotype: ANSP 300356, height 13.6 mm, width 11.5 
mm. 

Type locality: Off Potengi River, Natal, Estado Rio 
Grande do Norte, Brazil, depth unrecorded. 

Distribution: This species is known only from the ho- 
lotype, a hermit-crabbed shell, collected from a bottom 
of black mud in an unrecorded depth. 

Remarks: Calliostoma tenebrosum is readily distin- 
guished from other species of the C. pulchrum group by 
the relatively large, darkly colored shell that has the 
whorl periphery composed of three widely spaced spiral 
cords; the beads on the spiral cords are unusually large 
for this species group. 



Remarks: Calliostoma dentatum is closely related to 
C. pulchrum, C. roseolum. and the Plio-Pleistocene C. 
bowdenense Woodring, 1928. Shells of C. dentatum dif- 
fer from those of C. pulchrum by having a larger pro- 
toconch .X = 346 |um vs. 300 ^m; Quinn, in press); by 
having fewer spiral cords both above and below the pe- 
riphery; and by having the beads on the cords stronger, 
more sharply defined, and lateralK' compressed. Shells 
of C. dentatum differ from those of C. roseolum by 
having a larger protoconch x = 346 ^m vs. 324 jum), by 
being relatively narrower, b\' having a narrower and 
more angulate periphery, and by having laterally com- 
pressed beads on the spiral cords. Shells of C. dentatum 
differ from those of C. bowdenense by having more 
numerous spiral cords above the periphery; by having a 
narrower, more sharply angulate periphery; and by hav- 
ing weaker, more weakly beaded spiral cords on the base. 
The radula of C. dcntatun} (formula go. 4.1. 4. oo) differs 
from that of C. pulchrum illustrated by Clench and 



Calliostoma moscatellii new species 
(figures 29, 30, 109) 

Calliostoma carcellesi. Rios, 1970:25, pi. 4 (partim); 1975:23, 
pi. 6, fig. 61 (partim). (Non Calliostoma carcellesi Clench 
& .'^guayo, 1940). 

Calliostoma {Neocalliostoma) carcellesi: Rios, 1985:22, pi, 10, 
fig, 88 (partim). 

Description: Shell medium-sized, attaining height of 
27.7 mm, width of 17.9 mm, conical, nonumbilieate, 
rather coarsely sculptured. Protoconch 375 ^m maxi- 
mum diameter, 1 whorl. Teleoconch whorls about 10, 
flat to weakly convex; first whorl with weak, rounded 
axial riblets and 2 strong, beaded spiral cords; axial riblets 
replaced on subsequent whorls by low, rounded, crowded 
plicae in interspaces between spiral cords, plicae strong- 
est near suture and below peripheral cord; spiral sculp- 
ture increasing by intercalation to 5-8 cords and 4-9 
threads, of which abaxial 2-3 strong cords and 2-3 weak 



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THE NAUTILUS, Vol. 106, No. 3 






27 




28 







32 



J. F. Quinn, Jr., 1992 



Page 89 



cords or threads forming peripher\ , cords rather coarsely 
beaded, beads rounded, spirally elongate; periphery nar- 
rowly rounded. Base weakK convex, w ith 24-27 narrow, 
smooth to weakly beaded spiral cords and threads. Ap- 
erture subquadrate, lips thin, creiiulate; columella short, 
straight, thickened. Shell ground color light yellowish or 
pinkish tan with greenish iridescence, diffuse patches of 
tan to reddish brown, and occasional discrete spots of 
light brown; 2-5 basal spiral cords with elongate spots 
of brown; protoconch and first 2 whorls lilac. Animal (in 
alcohol) white except brown sides of snout; cephalic ten- 
tacles long, broad, tapering abruptK- near tip, ocular 
peduncles short (about 20% tentacle length), with large, 
black eye at tips; snout broad at base, tapering to rounded 
tip, tip with fringe of ver\ small, short papillae; epipo- 
dium with 1 tentacle on left, 2 on right, left larger than 
right, neck lobes well de\eloped, thin, semicircular, 
smooth. 

Holotype: MORG 29.293, height 25.9 mm, width 17.7 
mm. 

Type locality: Off Ilha de Santana, Estado de Rio de 
Janeiro, Brazil, in approximately 50-80 m. 

Paratypes: 1, USNM 860254; 1, MCZ 302591; 1, ANSP 
389337; 1, FSBC I 44075; 1, UP 189458; all from same 
lot as holotype. 

Other material: 5, Coltro collection; from same lot as 
holotype. 

Distribution: Numerous specimens of C. moscatellii, 
including live-collected specimens, have been obtained 
from depths of 50-200 m off Rio de Janeiro, Brazil, by 
shrimpers. 

Remarks: Shells of Calliostoma moscatellii are ver\' 
similar to those of C. carcellesi (figures 31, 32) but differ 
by being markedly narrower; by having a smaller pro- 
toconch (375 ^m vs. 400-425 fim); by having fewer basal 
spiral cords, most of which are weakly but discretely 
beaded; by having more strongly spirally elongate beads 
on the supraperipheral spiral cords; by having lilac apical 
whorls; and by having a distinct color pattern. Callios- 
toma moscatellii is also similar to C. jucundum but has 
narrower shells with more numerous, narrower spiral 
cords that bear much smaller, spirally elongate beads. 
The radula of C. moscatellii (formula 00.6.I.6.00; figure 
109) resembles that of C. pulchrum illustrated by Clench 
and Turner (1960:pl. 3, fig. 3) but has a broader rachidian 
tooth, and the cusps of the rachidian and lateral teeth 
are more finelv denticulate. 



This species is named for Sr. Renato Moscatelli, in 
recognition of his support of malacological publications 
in Brazil. 

Calliostoma apicii^um Dall, 1881 
(figures 33, 34) 

Catliosionm apicinum Dall, 1881:46; 1889b:162, pi. 24, figs. 3, 
3a.— Piisbrv, 1890:379, pi. 60, figs. 1, 2.— Johnson, 1934: 
69. 

Calliostoma {Calliostoma) apicintim. Dall, 1889a:366, pi. 24, 
figs. 3, 3a. 

Calliostoma roseolum. Clench & Turner, 1960:19, pi. 15, figs. 
1,2 (partim).— Abbott, 1974:43 (partim).— Quinn, 1979: 
26 (partim),— Sander & Lalli, 1982:table 4. (Non Cal- 
liostoma roseolum Dall, 1881). 

Description: See Dall (1881:46). 

Lectotype: (by implication of holotype; Clench & Tur- 
ner, 1960): USXM 95013, height 7.7 mm, width 6.7 mm. 

Type locality: Off Barbados, Blake station (number and 
coordmates unrecorded), in 183 m. 

Paralectotypes: 1, USNM 95012; off Barbados, Blake 
Station 290, 13ni'54"N, 59°39'45"W, 134 m; 9 March 
1879. — 2, MCZ 7564; Blake station (number and coor- 
dinates unrecorded), 183 m. 

Other material: 1, ANSP 353528; 1, FSBC I 44077; off 
Holetown, Barbados, 175-225 m; ex F. Sander. — 5, Sun- 
derland collection; off St. James, Barbados, 100 m; dredge; 
ex F. Sander. — 1, Sunderland collection; off St. James, 
Barbados, 152 m; 1988; ex F. Sander. — 6, Sunderland 
collection; off St. James, Barbados, 175-225 m; ex F. 
Sander. — Several other specimens, all from Barbados, in 
private collections. 

Distribution: This species is known only from off Bar- 
bados in depths of 100-225 m; 1 have only seen one live- 
collected specimen from a depth of 100 m. 

Remarks: This species was synonymized with Callios- 
toma roseolum b\ Clench and Turner (1960), who 
thought that the shells described by Dall as C. apicinum 
were merely juveniles of C. roseolum. Shells of C. api- 
cinum. however, are distinguished from those of C. ro- 
seolum by retaining flat-sided, peripherally subangulate 
whorls as adults (7-1- whorls) rather than by ha\ing con- 
cave whorls with broadly rounded peripheries after the 
third or fourth whorl; by having a larger protoconch 
(360-375 ^lm vs. 300-330 ^m) that has a purplish-brown 
tip; by having the apical two teleoconch whorls brown 
or purplish brown; by having more widely spaced, smooth 



Figures 23, 24. Calliostoma bermudctise new species, holotype, DMNH 187591; height 11.9 mm, width 8.9 mm. Figures 25, 
26. Calliostoma dentatum new species, hoiot\pe, I'SXM 859358; height 5.7 mm, width 5,2 mm. Figures 27, 28. Calliostoma 
tcncbrosum new species, holot\'pe, ANSP 300356; height 13,6 mm, width 11,5 mm. Figures 29, 30. Calliostoma moscatellii new 
species, holotype. MORG 29 293; height 25,9 mm, width 17 7 mm Figures 31, 32. Calliostoma carcellesi Clench & ,\guayo, 1940, 
holotype, MCZ 104719; height 20,5 mm, width 17.5 mm. 



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THE NAUTILUS, Vol. 106, No. 3 












Figures 33, 34. Calliostoma apicintim Dall, 1881, specimen from off Barbados, FSBC I 44077; height 10.0 mm, width 8.2 mm. 
Figures 35, 36. Calliostoma delnle new species, holotype, USNM 860255; height 8. 1 mm, width 7.0 mm. Figures 37, 38. Calliostoma 
Indiana Dall, 1889, specimen from off Honduras, FSBC I 44071; height 10.4 mm, width 9.0 mm. Figures 39, 40. Calliostoma 
orion Dall, 1881, specimen from John Elliott Pillsbury ,Station P-425, UMML 30.;3635; height 16 2 mm, width 14 4 nmi. 



or very weakly beaded spiral cords on the outer two- 
thirds of the base; by often having a chink-like umbilicus; 
and by having a shorter columella that often has a sub- 
conical, medial swelling. Calliostoma apicinum seems 
to be most closely related to the sympatric C. debile (see 
Remarks under the latter species for comparisons). Spec- 
imens that I can assign unequivocally to C. apicintim 
seem to come only from Barbados. Some other specimens 
w ith similar morphologies from the Bahama Islands (MCZ 
uncatalogued) and northwestern Cuba (M("Z 7566) are 
not here considered conspecific with the Barbados spec- 



imens. A paralectotype of C. apicinum (MCZ 7565) from 
off Havana, Cuba, is definitely not this species; the spec- 
imen has a uniquely undulate peripheral spiral cord on 
several of the adapical teleoconch whorls and probably 
represents an undescribed species. 

Calliostoma dehilc new species 
(figures 35, 36) 

Description: Shell small to medium-sized, attaining 
height of 10.3 mm, width of 8.7 mm, conical, nonum- 



J. F. Quinn, Jr., 1992 



Page 91 



bilicate, finely sculptured. Protoconch 350-365 A^ni max- 
imum diameter, 1 whorl. Teleoconch whorls 7.7, flat; 
first 2 whorls with low, rounded a.xial riblets and 2 strong, 
beaded spiral cords, abapical cord stronger on first 4 
whorls, set with strong, conical beads; a.\ial sculpture 
absent on subsequent whorls; spiral sculpture increasing 
by intercalation to 7-10 cords and 0-3 threads, of which 
aba.xial 3, sometimes 2, cords forming periphery, supra- 
peripheral cords fine, sharp, finely beaded; periphery 
subcarinate, slighth projecting, peripheral cords stronger 
than supraperipheral cords, set with conical beads. Base 
flat, with 10-14 narrow, flattened, smooth to weakly ru- 
gose spiral cords and 2-3 threads or weak cords near 
periphery. Aperture subquadrate, lips thin, weakly cren- 
ulate; columella straight, thickened. Shell ground color 
ivory, periphery light pinkish brown or with pale to 
bright spots of pinkish or \ ellowish brou n or clear rose. 

Holotype: USNM 860255, height 8.1 mm, width 7.0 
mm. 

Type locality: Off St. James, Barbados, in 175-225 m. 

Paratype: 1, FSBC I 45776; off St. James, Barbados, 152 
m; dredge; ex F. Sander. 

Other material: 4, Sunderland collection; off St. James, 
Barbados, 152 m; 1988; dredge; ex F. Sander. — 6, Sun- 
derland collection; off St. James, Barbados, 175-225 m; 
dredge; ex F. Sander. 

Distribution: Calliostoma debile is known only from 
empt\ shells collected off Barbados in 152-225 m. 

Remarks: Shells of Calliostoma debile are most similar 
to those of the sympatric C. apicinitm but differ by 
having a slightK smaller protoconch; b\ lacking the brow n 
or purplish-brown apical whorls; by having a more pro- 
jecting periphery, usually composed of three (sometimes 
two) close-set spiral cords; by usually having two to three 
spiral threads or weak cords between the circumbasal 
cord and the outermost strong basal spiral cord; by having 
distinct radial threads that finely bead the basal spiral 
cords; by having a less-thickened columella that lacks 
any indication of a medial swelling; and by lacking strong 
ridges within the aperture. 



Calliostoma indiana Dall, 1889 
(figures 37, 38) 

Calliostoma (Eucasta) indiana Dall, lS89a;368, pi. 32, figs, 3, 

5.— .Abbott, 1974:46, fig. 333, 
Calliostoma indiana: CHench & Turner. 1960:52, pi 34, fig. 1. 

Description: See Dall (1889a:368) and Clench and Tur- 
ner (1960:52). 

Holotype: USNM 214273, height 7.9 mm, width 6.9 



Type locality: Off Grenada, Lesser Antilles, Blake Sta- 
tion 247, 12°05'25"N, 61°47'15"W, in 311 m. 

Other material: 1, FSBC 1 44071; off Honduras, Cape 



Hatteras station, 15°32.36'N, 8r39.80'W, 50-85 m; 8 
April 1987; rock dredge. 

Distribution: This species is now known from two spec- 
imens, the holotype from Grenada and the Cape 
Hatteras specimen from off Honduras. Based on know 1- 
edge of the distributions of other species of Calliostoma 
(Clench & Turner, 1960; Quinn, herein, in press), I be- 
lieve C. indiana probabK inhaliits deep fore-reef areas 
(50-150 m) of Central America and the Greater and 
Lesser Antilles. 

Remarks: The new shell reported here is larger than 
the holotype (height 10.4 mm, width 9.0 mm), has five 
rather than four strong supraperipheral cords, and has 
ten rather than seven basal spiral cords; otherwise the 
two shells are very similar. Based on shell characters, this 
species clearly belongs to the C. pidchrum species com- 
plex and is most similar to C. echinatiim. C. orion, C. 
sopidiim, and C. cinctetliim. 



Calliostoma orion Dall, 1889 
(figures 39, 40) 

Calliostoma orion Dall, lS89a:367, pi. 28, fig. 2; 1889b:162.— 
Pilsbrv, 1890:383, pi. 48. fig. 18.— Johnson, 1934:70.— 
Clench & Turner, 1960:54, pi. 35, figs. 1, 2.— Quinn, 1981: 
151, figs, 1-13. 

Description: See Dall (1889a;367) and Quinn (1981: 
151). 

Holotype: USNM 214272, height 4.6 mm, width 4.3 
mm. 

Type locality: Off Havana, Cuba, Blake Station 62, in 
146 m. 

Other material: 2, FSBC I 31518; off western end of 
Isla de Roatan, Honduras, 19.8 m; 15 September 1982; 
SCUBA collection by W. C. Jaap. — 1, Crnkovic collec- 
tion; off western end of Isla de Roatan, Honduras, ap- 
proximatelv 30 m; 1990; SCUBA collection bv L. A. 
Crnkovic — 1, UMML 30.5688; off Isla de Roatan, Hon- 
duras, John Elliott Pillsbury Station P-629, 15°58.2'N, 
86°09.0'W, 40 m; 21 March 1968; 40-ft otter trawl— 1, 
UF 36324; Isla de Providencia, Colombia (off Nicara- 
gua), "shallow water"; August 1971; C. R. Gilbert, col- 
lector.— 1, UMML 30.3635; off Punta Manzanillo, Pan- 
ama, John Elliott Pillsbury Station P-425, 9°38.9'N, 
79°15.3'W, 70-64 m; 19 July 1968; 10-ft otter trawl.— 
3, Sunderland collection; off Barbados; ex F. Sander. 

Distribution: Calliostoma orion is now known from 
almost the entire Caribbean coast of Central America, 
Cuba, the Bahama Islands, and Barbados. 

Remarks: 1 discussed the ecology, shell and radular 
morphologies, and other distributional records in a pre- 
vious paper (Quinn, 1981). The present specimens in- 
crease the maximum known size to 20.2 mm high, 17.1 
mm wide; extend the maximum depth from w hich living 
specimens have been obtained from 43 m to 64-70 m; 



Page 92 



THE NAUTILUS, Vol. 106, No. 3 



and add new collection localities off Honduras, Nicara- 
gua, and Panama. 

Calliostoma jernandezi Princz, 1978 
(figures 41, 42) 

Calliostoma jernandezi Princz, 1978:1.52-154, figs. 1, 2. 

Description: Shell large, attaining height of 28. .5 mm, 
width of 25.5 mm, conical, umbilicate, finely sculptured. 
Protoconch 350-375 ^m maximum diameter, 1 whorl. 
Teleoconch whorls more than 10, flat to concave; first 2 
whorls flat-sided, with low, rounded a.xial riblets and 2- 
4 finely beaded spiral cords; axial sculpture absent on 
subsequent v\horls; spiral cords increasing by intercala- 
tion to 17-22, of which abapical 3-4 cords forming pe- 
riphery, all cords subequal and with well-defined, close- 
set, rounded beads; periphery narrowly rounded. Base 
flat to weakly convex, with 17-28 finely beaded spiral 
cords, those near periphery narrow, sharp, cords becom- 
ing broader and flatter adaxially, often splitting into 2 
subequal cords; circumumbilical 1-2 cords strongest, tu- 
berculate. Umbilicus funnel-shaped, 17%-24% maxi- 
mum shell diameter, wall almost vertical, white. Aper- 
ture subquadrate, thickened within and with strong, sharp 
ridges, lips thin, crenulate; columella weakly sigmoid, 
thickened, terminating in blunt, rounded tubercle. Shell 
ground color cream to straw with irregular, diffuse patch- 
es of light brown above periphery; periphery with reg- 
ular series of squarish, rose-brown to brown spots; inter- 
spaces of supraperipheral spiral cords golden brown; base 
finely mottled with cream and yellow-brown. 

Holotype: Collection of "Familia Fernandez en Ma- 
racay, Estado Aragua, Venezuela" (Princz, 1978:152), 
height 14.7 mm, width 15.8 mm. 

Type locality: "Grottos" at Las Cuevas Beach, Trinidad, 
depth not reported. 

Other material: 1, UMML 30.7162; off Peninsula de la 
Guajira, Colombia, John Elliott Pillsbury Station 
P-769, 12°31.0'N, 71°41.0"W, 14.3-146 m; 28 July 1968; 
10-ft otter trawl.— 1, UMML 30.6424; off Peninsula de 
Paria, Venezuela, John Elliott Pillsbury Station P-709, 
11°08.8'N, 62°46.1'W, 46 m; 19 July 1968; 10-ft otter 
trawl.— 1, UMML 30.6454; off Peninsula de Paria, Ven- 
ezuela, John Elliott Pillsbury Station P-708, 11°24.7'N, 
62''40.5'W, 69-73 m; 19 July 1968; 10-ft otter trawl.— 
1, UMML 30.6373; off Peninsula de Paria, Venezuela, 
John Elliott Pillsbury Station P-707, ir21'N, 62°21'W, 
78 m; 19 July 1968; 10-ft otter trawl— 1, UMML 30.6373; 
off Peninsula de Paria, Venezuela, John Elliott 
Pillsbury Station P-705, 10°45'N, 62°00'W, 77-86 m; 18 
July 1968; 10-ft otter trawl.— 2, MCZ 273512; George- 
town, Guyana, Ch.mn Cruise 35, Station 354-36, 8°10.5'- 
8°10.0'N, 57'='48'W, 97-110 m; 28 April 1963—2, USNM 
866503; 1, FSBC I 44072; 1, UF 189459; 5, UMML 
30.5840; off Pointe Isere, French Guiana, John Elliott 
Pillsbury Station F'-650, 6°07'N, 52°19'W, 84-91 m; 8 
July 1968; 10-ft otter trawl. 



Distribution: Calliostoma jernandezi occurs from Su- 
riname northward and westward to off Cabo de la \'ela, 
Colombia, in depths of about 45- 245 m; living specimens 
have been collected from depths of 84-146 m. 

Remarks: Although the description presented by Princz 
(1978) is incomplete, the specimens recorded here seem 
to be referrable to Calliostoma jernandezi. The shells 
of this species are very similar to those of C. scalenum 
but differ by having a larger protoconch (375 ^m vs. 
320-325 fim); by having fewer, stronger, more evenly 
sized spiral cords; by having stronger, more closely spaced, 
and more s\ mmetrical beading; by having an umbilical 
wall that is almost vertical rather than sloping steeply 
inward; and by having a different, lighter color pattern. 
The spiral cords at the periphery and on the base of shells 
of C. jernandezi tend to increase in number by fission 
of existing cords rather than by intercalation of new 
cords; this tendency is rare among species of Calliostoma 
that I have examined. 

Data obtained from the original field data sheets in- 
dicate that C. jernandezi occurs on shell-hash and coral- 
rubble bottoms, often in association with sponges and 
octocorals. 



Calliostoma scalenum new species 
(figures 43, 44, 110, 111) 

Description: Shell large to ver\- large, attaining height 
of 40.6 mm, width of 34.2 mm, conical, umbilicate, finely 
sculptured. Protoconch 320-325 yum maximum diameter, 
1 whorl. Teleoconch whorls about 10, first 4-5 whorls 
flat-sided, subsequent whorls weakly to strongly concave; 
first 3 whorls with low, rounded axial riblets and 2-6 
finely beaded spiral cords; axial sculpture absent on sub- 
sequent whorls; spiral cords increasing by intercalation 
to 25-30, of which abapical 8-12 cords forming periph- 
ery, alternating in size, finely beaded, beads close-set, 
rounded, conical; periphery narrowly rounded. Base flat 
to weakly convex, with as many as 42 narrow, weakly 
beaded spiral cords, those on abaxial half alternating in 
size, those on adaxial half subequal except 2-3 stronger, 
tuberculate circumumbilical cords. Umbilicus funnel- 
shaped, 14%-23% maximum shell diameter, wall smooth, 
white, often with \ ellow-brown flush, .\perture subquad- 
rate, thickened within and with strong ridges, lips thin, 
crenulate; columella sigmoid, thickened, terminating in 
blunt, rounded tubercle. Shell ground color light chestnut 
to red-brown with irregular, diffuse axial flammules of 
darker ground color and white; light and dark flammules 
more distinct and closely spaced on periphery, .\nimal 
(in alcohol) white; foot with numerous, scattered, small, 
opaque, white spots; cephalic tentacles long, tapered, 
bases with short ocular peduncles bearing small, black 
eye at tips; epipodium with 4 pairs of tentacles, neck 
lobes well-developed, thin, semicircular, smooth-edged. 

Holotype: USNM 859356, height 31.5 mm, width 25.9 



J. F. Quinn, Jr., 1992 



Page 93 



Type locality: Stetson Bank, southeast of Galveston, 
Texas, 28°09'54"N, 94°18'00"W, in 21-27 m. 

Paralypes: 2, HMNS 15022; 2, USNM 859357; 1, MCZ 
297051; 1, ANSP 367152; 1, AMNH 225980; 1, LACM 
2292; 1, UF 110222; 2, FSBC I 32315; 34, HMNS 15850; 
all from same lot as holot>pe.— 3, ANSP 338470; off 
Freeport, Texas, NW Gulf Survey station, 28°10'N, 
94°55W, 51 m; dredge; A. Kight, collector 

Other material: More than 200 lots listed b> Quinn (in 
press). 

Distribution: Calliostoma scalenum inhabits offshore 
waters on calcareous substrates from North Carolina 
southward through the Florida Keys and throughout the 
Gulf of Mexico in depths of 25-80 m. 

Remarks: Shells of Calliostoma scalenum have been 
identified pre\iously as C, jujuhinum \\ ithoiit exception 
(see Quinn, in press, for complete synonymy). Callios- 
toma scalenum differs from C. jujuhinum by having 
shells that are larger and more narrowly conical; that 
have a slightK' larger protoconch (320-325 ^m vs. 315 
jj.m)\ that lack fine, coUabral threads after the fourth or 
fifth whorls; that have two rather than three distinct sizes 
of spiral cords with discrete, symmetrical rather than 
spiralK elongate beads; that have a greater number of 
spiral cords, particular!} on the base (35-42 vs. 25-30 
total); and that lack very dark-colored apical whorls. 
Shells of C. scalenum are also similar to those of C. 
tampaense but differ b\ having straight-sided apical 
whorls that lack a strong, sharply beaded peripheral ca- 
rina; by having finer, more numerous, and more fineK' 
beaded spiral cords; and b\ being proportionately much 
narrower. Both C. jujuhinum and C. tampaense are also 
shallow-water species, neither species occurring alive in 
depths exceeding 11m (Quinn, in press). The radula of 
C. scalenum (formula 00.6.I.6.00; figures 110, 111) is 
very similar to that of C. tampaense illustrated by Clench 
and Turner (1960:pl. 5, fig. 2, as C. jujuhinum) and C. 
vinosum (figure 114 herein) in having six pairs of lateral 
teeth, of which the outer two pairs lack cusps. Radulae 
of C. jujuhinum (figure 112) and C. cuhense (figure 1 13) 
are also very similar in morphology but differ in having 
only five pairs of lateral teeth. One character common 
to all of these radulae is that the cusps of the inner lateral 
teeth arise from the outer corner of the tooth bases; this 
characteristic has not been explicitly noted before and 
is not clear in anv of the previously published line draw- 
ings (Clench & Turner, 1960;pl. 5,'figs. 1, 2; Calvo, 1987: 
fig. 26). 

Calliostoma cuhense new species 
(figures 45, 46, 113) 

Description: Shell medium-sized, attaining height of 
16.6 mm, width of 14.3 mm, conical, umbilicate, rather 
coarsely sculptured. Protoconch about 350 ^m maximum 
diameter, 1 whorl. Teleoconch whorls 7-8. flat to weakK 
concave; first 2-3 whorls with low , sharp axial riblets and 



2-4 strong, beaded spiral cords; axial riblets replaced by 
low, crowded, rounded axial threads in interspaces be- 
tween primar\ spiral cords; spiral cords increasing by 
intercalation to 6-12, of which abapical 1-2 beaded cords 
and broad, smooth cord forming periphery, usually al- 
ternating in size, primary cords strongly beaded, sec- 
ondary cords smooth or finely beaded by axial threads; 
periphery narrowly rounded, smooth peripheral cord of- 
ten dissected by 1 (rarely 3-5) fine, spiral stria. Base flat 
to weakly convex, with 10-13 narrow, smooth or weakl) 
beaded spiral cords. Umbilicus funnel-shaped, 18%-21% 
maximum shell diameter, w all w hite. Aperture subquad- 
rate. thickened w ithin and w ith strong ridges, lips thin, 
crenulate; columella sigmoid, thickened, terminating in 
moderately strong, rounded denticle. Shell ground color 
tan to pinkish brown with numerous white dots, prin- 
cipally on indi\ idual beads of primary spiral cords but 
broader on peripher\ ; interspaces between spiral sculp- 
ture golden brown, usually at periphery and on base, 
occasionally above periphery. Animal (reconstituted in 
trisodium phosphate) uniformly tan; cephalic tentacles 
long, slender, gradually tapering; eyestalks short (about 
10%-15% tentacle length), stout, with large black eye at 
tips; snout longer than broad, with fringe of fine, short 
papillae; epipodium w ith 3 pairs of long tentacles; neck 
lobes well-developed, left lobe finely digitate, right lobe 
smooth-edged. 

Holotype: ANSP 196933, height 14.7 mm, width 13.7 
mm. 

Type locality: Off Pini Pini, Cardenas Bay, Matanzas, 
Cuba, in 1 1 m. 

Paratypes: 4, USNM 438265; ToM.is Barrera Expe- 
dition Station 224, Cabo San Antonio, Pinar del Rio, 
Cuba,— 12, ANSP 196926; off Cardenas Bay, Matanzas, 
Cuba, 11 m; 11 February 1954; V. Conde. collector. — 
9, ANSP 357161; 2. FSBC 1 32405; from same lot as 
holotype. — 2. MCZ 204705; Peninsula de Hicacos, Ma- 
tanzas, Cuba; 1940; V. Conde, collector.— 4, MCZ 216120; 
Cayo Fragoso, Villa Clara, Cuba; R. Humes, collector. — 
1, MCZ 235536; off Cayo Fragoso, Villa Clara, Cuba; R. 
T. Abbott, collector.— 8, MCZ 129623; Cayo Frances, 
Villa Clara, Cuba; ex P. J. Bermudez collection. — 3, ANSP 
216122; Caibarien, Villa Clara, Cuba; R. Humes, collec- 
tor. 

Distribution: This species seems to be restricted to the 
northern coast of Cuba; live-collected specimens were 
obtained from off Pini Pini in 11 m. 

Remarks: Shells of Calliostoma cuhense most resemble 
those of C. adelae in shape and size and in having rather 
coarse sculpture relative to other species of the C. ju- 
juhinum species complex. Shells of C. cuhense differ 
from those of C adelae by having more numerous, un- 
equally sized spiral cords both above and below the pe- 
riphery; b\ having the basal spiral cords closeK spaced, 
separated b\ narrow. \'-shaped grooves, rather than 
widely spaced and separated by broad, flat-bottomed 



Page 94 



THE NAUTILUS, Vol. 106, No. 3 








/\ 







51 




J, F, Quinn, Jr., 1992 



Page 95 



grooves; by having distinct, fine collabral threads be- 
tween the supraperipheral spiral cords; and by having a 
rather broad and smooth rather tlian narrow and strongly 
beaded peripheral cord. The shells of C cuhense from 
the ToM.4s Barrera Expedition (USNM 438265) are ap- 
parently those that Henderson (1916:185) recognized as 
"an apparently new Calliostoma" from dredgings off 
Cabo Cajon (near Cabo San Antonio) on a bottom of 
coral sand with patches of seagrass and sponge, the area 
swept by strong currents. 

The intestine of one specimen [ex FSBC 1 32405) was 
packed with amorphous organic material, hydroid stalks, 
and several different forms of foraminiferans. 

Calliostoma ptirpurciim new species 
(figures 47-30) 

Description: Shell medium-sized, attaining height of 
23.9 mm, width of 20.2 mm, conical, umbilicate, rather 
coarsely sculptured. Protoconch unknown. Teleoconch 
whorls about 9, flat to weakK convex; first 1-2 whorls 
worn on ail specimens but apparently with axial riblets 
and 2-4 spiral cords; axial sculpture absent on subsequent 
whorls; spiral sculpture increasing by intercalation to 8- 

14 cords and 1-8 fine threads, of which abapical 3-5 
cords forming periphery; peripher\ broadly rounded. 
Base flat to weakly convex, with 13-18 coarsely beaded 
spiral cords. Umbilicus funnel-shaped, 18%-26/o maxi- 
mum shell diameter, wall white to intense violet. Ap- 
erture subquadrate, thickened within and with strong, 
sharp ridges, lips thin, crenulate; columella weakly sig- 
moid, thickened, often with 1-3 small, sharp denticles 
or ridges on lower third, terminating in strong, rounded 
tubercle. Shell ground color tan to reddish brown, mot- 
tled with cream to yellowish brown. 

Holotype: USNM 860247, height 21.1 mm, width 19.6 
mm. 

Type locality: Off Cabo de la Vela, Colombia, in 61- 
91 m. 

Paratypes: 3, USNM 860248; 1, FSBC I 44074; from 
same lot as holotype. 

Other material: 5, Deynzer collection; from same lot 
as holotype.— 1, UMML 30.7204; off Cabo de la Vela, 
Colombia, John Elliott Pillsbury Station P-773, 
12°17,0'N, 72°15.0'W, 60-64 m; 29 July 1968; 10-ft otter 
trawl— 3 broken, FSBC I 32742; about 5 km north of 
La Guardia, Isia de Margarita, Venezuela, beach drift; 
19 November 1987; W. G. & C. B. Lyons, collectors.— 

15 broken, FSBC I 32747; 6 km north of La Guardia, 



Isla de Margarita, Venezuela, beach drift; 18 November 
1987; W. G. & C. B. Lyons, collectors. — 1, UMML 
30.7269; off Galera Point, Trinidad, John Elliott 
Pillsbury Station P-840, 10°40.5'N, 60°37.5'W, 33-37 
m; 1 July 1969; 10-ft otter trawl. 

Distribution: Calliostoma purpiireum is known from 
off northeastern Colombia, Isla de Margarita, Venezuela, 
and Trinidad; most specimens have been collected from 
33-91 m, but broken shells are also known from beach 
drift. 

Remarks: Shells of Calliostoma purpiireum are very 
similar to those of C. jujubinum but differ by lacking 
collabral threads between the spiral cords, by having the 
spiral cords narrower and more finely beaded, and by 
having the umbilical wall almost vertical rather than 
strongK sloped. Most specimens of C purpureum have 
one or more small, sharp denticles on the columella, 
whereas specimens of C. jujubinum very rarely have 
only a single denticle; I know of no other species that 
has a denticulate columella. Additionally, whereas spec- 
imens of C. jujubinum may have the umbilical wall 
flushed with pinkish brown, I have never seen any spec- 
imen of that or any other species of western Atlantic 
Calliostoma with the violet color or the intensity of any 
other color displayed on the umbilical wall of specimens 
of C. purpureum except the red-brown in C. brunneum 
and C. barbouri. Shells of C. purpureum may have been 
reported from Curasao by de Jong and Coomans (1988) 
as C, tampaense. 

Almost all of the specimens collected at Isla de Mar- 
garita consisted of only the last one to three whorls; 
because the damage to these shells is almost identical to 
that illustrated by Vermeij (1978: fig. 2.10), this damage 
may have been the result of predation by xanthid crabs. 
This species has been collected principally in areas with 
large concentrations of sponges and was probably the 
species from Islas Los Roques, Venezuela, that Work 
(1969) observed eating an encrusting sponge. 

Calliostoma fiicosum new species 

(figures 51, 52) 

Description: Shell medium-sized, attaining height of 
12.8 mm, width of 12.6 mm, conicoturbinate, umbilicate, 
rather coarseK sculptured. Protoconch eroded. Teleo- 
conch whorls about 7, weakly concave; first whorl eroded, 
next whorl with 3 strong, strongly beaded spiral cords; 
spiral cords increasing by intercalation to 7, of which 
abapical 2 cords forming periphery; periphery broadly 
rounded, adapical peripheral cord beaded, abapical pe- 



Figures 41, 42. Calliostoma jernandezi Princz, 1978, specimen from John Elliott Pillsbury Station P-650, UMML 30.5840; 
height 21.5 mm, width 19,;3 mm. Figures 43, 44. Calliostoma scalenum new species, holotype from Stetson Bank, SE of Galveston, 
Texas, USNM 859356; height 31.5 mm, width 25.9 mm. Figures 45, 46. Calliostoma cubense new species, holotype from Pini 
Pini, Cuba, ANSP 196933; height 14.7 mm, width 13.7 mm. Figures 47-50. Calliostoma purpureum new species. 47, 48. Holotype, 
USNM 860247; height 21.1 mm, width 19.6 mm. 49, 50. Paratype, USNM 860248; height 15.8 mm, width 15.4 mm. Figures 51^ 
52. Calliostoma fucosum new species, holotype, USNM 860259; height 12.8 mm, width 12.6 mm. 



Page 96 



THE NAUTILUS, Vol. 106, No. 3 



ripheral cord smooth with shallow medial stria. Base 
weakK convex, with 9 broad, coarseK beaded and 2 
narrow, smooth spiral cords; circumumbilical cord 
strongest and most strongly beaded. Umbilicus 20% max- 
imum shell diameter, wall white with faint flush of pink. 
Aperture subquadrate, lips thin, weakK crenulate; col- 
umella weakly sigmoid, somewhat thickened, terminat- 
ing in sharp denticle. Shell ground color predominantly 
orange-brown with spots and flammules of white and 
darker orange-brown. 

Holotype: USNM 860259, height 12.8 mm, width 12.6 
mm. 

Type locality: Off Cabo de la Vela, Colombia, John 
Elliott Pillsbury Station P-774, 11°56.5'N, 72°17.9'W, 
in 5-9 m. 

Distribution: This species is onl\ known from the ho- 
lot\pe shell. 

Remarks: The holotype of Calliostoma fucosiim resem- 
bles a small C. euglypfum, but the shell is umbilicate, 
proportionately broader, has stronger and coarser sculp- 
ture, and has a much brighter color pattern. 

The specimen of C. fucosum was trawled in shallow- 
water from a bottom composed principally of calcareous 
red algae (UMML, unpublished). 

Calliostoma vinosiim new species 
(figures 53-56, 114) 

Calliostoma (Elmerlinia) Inillisi: Rios, 1970:24, pi. 5. (Non 

Calliostoma bidlisi Clench & Turner, 1960). 
Calliostoma barbouri. Rios, 197.5:23, pi. 5, fig. 60; 1985:22, pi. 

10. fig. 87. (Non Calliostoma barbouri Clench & Aguavo, 

1946). 
Calliostoma javanicum: Leal, 1991:45, 353 (partim). (Non Tro- 

cliiis javanicum Lamarck, 1822), 

Description: Shell medium-sized, attaining height of 
19.8 mm, width of 21.3 mm, conical, umbilicate, finely 
sculptured. Protoconch 325-350 /ttm maximum diameter, 
1 whorl. Teleoconch whorls 8, flat to weakly convex; first 
whorl initially with 2 strong spiral cords and low, round- 
ed axial riblets; axial sculpture absent after first one-third 
whorl; spiral sculpture increasing by intercalation to 9- 
15 beaded cords and 0-6 smooth to finely beaded threads, 
of which abapical 2 cords forming periphery, beads 
rounded; surface of first 4-5 whorls microscopically frost- 
ed; periphery narrow, subcarinate, abapical peripheral 
cord often with shallow, median groove. Base flat to 
weakly concave, with 12-14 narrow , finely beaded spiral 
cords and 0-3 spiral threads. Umbilicus funnel-shaped, 
18%-22% maximum shell diameter, wall smooth, white 
or flushed with yellow. Aperture subquadrate, lips thin, 
crenulate; columella weakly sigmoid, weakly thickened, 
terminating in sharp denticle. Shell ground color deep 
red-brown with crescentic streaks of yellow-brown above 
periphery; base golden brown with 6-9 spiral lines of 
red-brown. Animal (in alcohol): foot red-brf)w ti w ith large, 
white papillae; mantle with narrow band of white and 



red-brown spots at edge behind which is series of dark 
brown spots extending posteriorly as gradually fading 
streaks; cephalic tentacles red-brown with darker median 
stripe, white near base, left tentacle stout, about length 
of snout, right tentacle longer and more slender; snout 
long, broad, mottled red-brown and w hite, with anterior 
fringe of long papillae; epipodium with 4 pairs of red- 
brown tentacles, neck lobes well-developed, semicircu- 
lar, left lobe smooth, right lobe finely fringed. 

Holotype: MORG 29.294, height 13.5 mm, width 15.3 
mm. 

Type locality: Off Guarapari, Estado de Espirito Santo, 
Brazil, in 18-22 m. 

Paratypes: 1, USNM 860256; 1, FSBC I 44067; both 
from same lot as holotype. — 1, MORG 15.043; off Cabo 
Orange, Estado de Amapa, Brazil, .Almir.^nte Saldanh.\ 
Station 2029, 103 m; 30 November 1968. 

Other material: 1, MNHN uncatalogued; 
Marion-Dufresne Cruise MD-55, Station DC-40, 
20°40'S, 34°41'W, 60 m; 17 May 1987; dredge— 2, 
MNHN uncatalogued; Marion-Dufresne Cruise MD- 
55, Station DC-47, 20°42'S, 32°13'W, 94-105 m; 19 May 
1987; dredge. — 2, MNHN uncatalogued; Marion- 
Dufresne Cruise MD-55, Station DC-42, 20°55'S, 
34°01"W, 60 m; 17 May 1987; dredge —7, MNHN un- 
catalogued; Marion-Dufresne Cruise MD-55, Station 
DC-15, 2r37'S, 40°18'W, 37 m; 11 May 1987; dredge. 

Distribution: Calliostoma vinosttm is known in north- 
eastern Brazil from the Amazon River to just north of 
Rio de Janeiro, in 18-105 m. 

Remarks: Shells of Calliostoma vinosum are most sim- 
ilar to those of C. barbouri in shape, size, and color but 
differ b\ having a somewhat smaller protoconch (325- 
350 ^m vs. 350-400 nm); b\ having flat-sided rather than 
convex whorls; by having fewer, stronger, more coarsely 
beaded spiral cords, particularly on the base; by having 
axial riblets that disappear early on the first whorl rather 
than persisting onto the second whorl; b\ lacking col- 
labral threads between the primary spiral cords; and by 
having the first 4-5 whorls more distincth frosted. 

Most of the specimens examined for this study were 
trawled from bottoms on which bryozoans and coralline 
algae were abundant (J. & M. Coltro, personal com- 
munication; Leal, 1991). 



Calliostoma alternum new species 
(figures 57, 58) 

Description: Shell medium-sized, attaining height of 
15.6 mm, width of 13.9 mm, conical, umbilicate, fineK 
sculptured. Protoconch about 350 /um maximum diam- 
eter, 1 whorl. Teleoconch whorls about 8.5, flat; first 4 
whorls with low, rounded axial riblets and 2-7 beaded 
spiral cords; axial sculpture reduced to fine collabral 
threads or absent on subsequent whorls; spiral sculpture 



J. F. Quinn, Jr., 1992 



Page 97 



increasing by intercalation to 7-11 beaded cords and 5- 
6 threads, of which abapical 3 cords forming periphery; 
periphery narrowly rounded. Base weakly convex, with 
17 narrow, finely beaded spiral cords. Umbilicus 22%- 
24% maximum shell diameter, wall almost vertical, white. 
Aperture subquadrate, lips thin, weakly crenulate; col- 
umella weakly sigmoid, rather thin, terminating in small, 
rounded denticle. Shell ground color uniformK ivory or 
v\ ith \erv faint peripheral light brow n maculations. 

Holotype: USNM 860257, height 15.6 mm, width 13.9 
mm. 

Type locality: Northeast of Caracas, Venezuela, John 
Elliott Pillsbury Station P-737, 10°44.0'N, 66°07.0'W, 
in 60-73 m. 

Paratypes: 1, UMML 30.8374; off Peninsula de la Gua- 
jira, Colombia, John Elliott Pillsbi ry Station P-769, 
12°31.0'N, 71°41.0'\V, 143-146 ni; 28 Julv 1968; 10-ft 
otter trawl. — 1, UN4ML 30.6155; off Suriname, John El- 
liott Pillsbury Station P-684, 7°19.0'N, 56°51.0'W, 55- 
59 m; 14 July 1968; 10-ft otter trawl. 

Distribution: Shells of this species are known from 
northeastern Colombia, Venezuela, and Suriname, in 55- 
146 m. 

Remarks: Shells of Calliostoma alternitm are most sim- 
ilar to those of C. aulicum but differ b\ being evenly 
conical rather than coeloconoid; by having a more round- 
ed, less strongly projecting periphery; by having fewer, 
stronger, more coarsely beaded spiral cords, most of which 
are separated by a fine spiral thread; by having axial 
riblets that persist on the first four whorls rather than 
only on the first two; and by having the abapical of the 
two primary spiral cords remaining conspicuously strong 
on first five whorls. 

The specimens of C. altcrnum were collected on bot- 
toms composed of carbonate rock covered by calcareous 
algae and sponges (UMML, unpublished). 



Calliostoma aulicum nev\ species 
(figures 59-62) 

Description: Shell medium-sized, attaining height of 
15.9 mm, width of 16.1 mm, conical, umbilicate, finely 
sculptured. Protoconch about 350 m maximum diameter, 
1 whorl. Teleoconch whorls 7.3, weakly concave; first 
two whorls with low, rounded axial riblets and 2-3 bead- 
ed spiral cords; axial sculpture absent after second whorl; 
spiral cords strongly beaded, increasing by intercalation 
to 10-12, of which abapical 3 strongest cords forming 
periphery on last whorl, beads rounded; periphery nar- 
rowly rounded. Base flat to weakly convex, with 17-19 
narrow, finely beaded spiral cords. I'mbilicus funnel- 
shaped, 17%-20% maximum shell diameter, wall white. 
Aperture subquadrate, thickened and stronglv ridged 
within, lips thin, crenulate; columella weakly sigmoid, 
thickened, terminating in narrow denticle. Shell ground 
color ivory with few, faint patches of golden brown 



Holotype: USNM 860258, height 15.9 mm, width 16.1 
mm. 

Type locality: Off Colon, Panama, John Elliott 
Pillsbury Station P-451, 9°22.0'N, 79°56.0'W, in ap- 
proximately 12 m. 

Paratype: 1, UMML 30.8375; off Suriname, John El- 
liott Pillsbury Station P-669, 6°39.0'N, 55°15.5'W, 33 
m; 10 July 1968; 10-ft otter trawl. 

Other material: 1 fragment, UMML 30.7238; off Ori- 
noco River, Venezuela, John Elliott Pillsbury Station 
P-835, 9°36.0'N, 60°10.0"W, 48 m; 30 June 1968; 10-ft 
otter trawl.— 1 fragment, UMML 30.7306; off Galera 
Point, Trinidad, John Elliott Pillsbury Station P-840, 
10°40.5'N, 60°37.5'W, 33-37 m; 1 July 1968; 10-ft otter 
trawl. 

Distribution: Calliostoma aulicum is known from off 
Panama and eastern Venezuela in 12-48 m. 

Remarks: Shells of this species are very similar to those 
of Calliostoma tampaense but differ by having a slightly 
larger protoconch (350 nm vs. 325 nm); bv' having a 
more broadlv' rounded, more strongly projecting periph- 
ery; by being much thicker; by having more strongly 
beaded basal spiral cords; and by lacking a distinct color 
pattern. If the holotype of C. aulicum represents the size 
of a fully grown adult, as the thickened and ridged ap- 
erture seems to indicate, then C. aulicum is much smaller 
than C. tampaense, shells of which attain a height of 
about 30 mm (Quinn, in press). 



Calliostoma hirtum new species 
(figures 63, 64) 

Description: Shell medium-sized, attaining height of 
17.9 mm, width of 19.8 mm, conical, umbilicate, finely 
sculptured. Protoconch missing. Teleoconch whorls about 
7, flat to weakly concave; first remaining v\horl worn 
smooth; next whorl with 4 beaded spiral cords, cords 
increasing to 13, of which abapical 2 cords forming pe- 
riphery on last whorl, beads conical; periphery carinate, 
adapical peripheral cord stronger than abapical periph- 
eral cord. Base weakly concave to weakly convex, with 
17-18 narrow, weakly beaded spiral cords. Umbilicus 
funnel-shaped, 20% maximum shell diameter, wall al- 
most vertical, white or yellow-brov\n. Aperture subquad- 
rate, lips thin, crenulate; columella weakly sigmoid, thin. 
Shell ground color light yellow-brown with diffuse flam- 
mules of cream and light brown; interspaces between 
several spiral cords golden brown. 

Holotype: USNM 860260, height 17.9 mm, width 19.8 
mm. 

Type locality: Off Navidad Bank, northeast of the Do- 
minican Republic, John Elliott Pillsbury Station P- 

1160, 20°01'N, 68°51'W, in 521 m. 

Paratype: 1, UMML 30.8371; from same lot as holotype. 



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THE NAUTILUS, Vol. 106, No. 3 




53 














J. F. Quinn, Jr., 1992 



Page 99 






V^ ) 



-;7 



Figures 63, 66. Calliostonia atlantis C\euch & Aguayo, 1940, holotype, MCZ 135164; height 32.7 mm, width 33.6 mm. 



Distribution: This species is known only from the type 
locality. 

Remarks: The shells of Calliostoma hirtnni are most 
similar to those of C. javanicum but differ by having a 
stronger, more sharply beaded peripheral cord; by lack- 
ing axial sculpture in the interspaces between the spiral 
cords; and by having more numerous, sharper spiral cords, 
particularly on the outer part of the base. 

Calliostoma atlantis Clench & Aguayo, 1940 
(figures 65, 66) 

Calliostoma (Calliostoma) atlantis Clench & Aguavo, 1940: 

81, pi. 13, fig. 4. 
Calliostoma atlantis: Clench & Turner, 1960:62, pi. 44; Har- 

asewych, 1989:27, pi. 17. 
Calliostoma (Kombologion) atlantis: Abbott, 1974:4.5. 

Description: See Clench and Aguayo (1940:81) and 
Clench and Turner (1960:62). 

Holotype: MCZ 135164, height 32.7 mm, width 33.6 
mm. 

Type locality: Northwest of Mariel, Cuba, Atlantis 
Station 3306, 23°04'30"N, 82°37'00"W, in 604 m. 

Other material: 1, USNM 869000; off Great Inagua 
Island, Bahama Islands, Johnson-Sea-Link I Station JSL- 
1-2323, 21°01'45"N, 74°43'48"W, 628 m; 15 October 1988. 

Distribution: The specimen of this species reported here 
is only the second collected and extends the range from 



northwestern Cuba to off Great Inagua Island northeast 
of the eastern tip of Cuba. This specimen was collected 
by M. G. Harasewych, using the submersible 
Johnson-Sea-Link I from a large boulder in 628 m, a 
depth very similar to that from which the holotype was 
collected (604 ni). The animal was reported to be bright 
red (Harasewych, 1989:27). 

Remarks: Based on shell characters, Calliostoma at- 
lantis, C. torrei, C. dnopherum, and the new species C. 
rugosum, C. atlantoides, C rota, and C. coronatum 
form a distinctive species group. All seven species have 
shells w ith strongly bicarinate whorls, at least on the first 
several whorls; have, at the beginning of the first teleo- 
conch whorl, an adapical spiral cord that becomes weak 
or disappears on subsequent whorls; have a weak an- 
gulation or strong spiral cord below the peripheral cord; 
have the inner lip of the columella weakly or strongly 
reflected over the umbilicus, or thickened and filling the 
umbilicus with callus; and have the columella rounding 
into the outer lip. Other species that may belong to this 
group include C. cuhanum Clench & Aguayo, 1940, and 
the eastern Atlantic C. grimaldii Dautzenberg & Fischer, 
1896; C. leptophyma Dautzenberg, 1927; C. normani 
Dautzenberg, 1927; and C. caroli Dautzenberg, 1927. 

Calliostoma dnopherum (Watson, 1879), 
new combination 
(figures 67-69) 

Trochus (Margarita) dnopherns Watson, 1879:711; 1886:90, 
pi. 5, fig. 3. 



Figures 53-56. Calliostoma vinosum new species. 53, 54. Holotype, MORG 29.294; height 13.5 mm, width 15.3 mm. 55, 56. 
Paratype, \40RG 15 043; height 10.2 mm, width 11.9 mm. Figures 57, 58. Calliostoma alternum new species, hoiotvpe, USNM 
860257; height 15.6 mm, width 13,9 mm. Figures 59-62. Calliostoma auliciun new species, 55, 56. Hoiotvpe, USNM 860258; 
height 15,9 mm, width 16,1 mm 57, 58. Parat\pe from John Elliott Pillsbiry Station P-669, L'MML 30,8375; height 10.8 mm, 
width 11.2 mm. Figures 63, 64. Calliostoma hirtiim new species, holotype, USNM 860260; height 17,9 mm, width 19.8 mm. 



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THE NAUTILUS, Vol. 106, No. 3 




67 

Figures 67-69. Calliostoma dnopherum (Watson, 1879). Lectotype of Trochus {Margarita) dnophcrus. BM(NH) 1887.2.9.333; 
height 7.2 mm. width 8.8 mm. 



Margarites dnopherus: Lange de Morretes, 1949:58. 
"Margarites" dnopherus: Rios, 198.5:19, pi. 8, fig. 72. 

Description: See Watson (1879:711; 1886:90). 

Lectotype (here selected): BM(NH) 1887.2.9.333, height 
7.2 mm, width 8.8 mm. 

Type locality: Southeast of Recife, Brazil, Challenger 
Station 122, 9''05'S, 34°49'W, in 640 m. 

Paralectotypes: 4 (broken or fragments), BM(NH) 
1887.2.9.334-335; from same lot as holotype. 

Other material: 1, Coltro collection; off Ihla de Sao 
Sebastiao, Estado de Sao Paulo, Brazil, 600 m; dredge; 
1991. 

Distribution: This species is now known from off Recife 
to off Sao Paulo, Brazil (a range extension of appro.xi- 
mately 2000 km), in 600-640 m. 

Remarks: The shape and sculpture of the shells of this 
species are very similar to those of C rota, C. atlan- 
toides, and C. coronatum {q. v.); therefore, the species 
is here transferred to Calliostoma sensu lata. Shells of 
C. dnopherum differ from those of C. rota by being 
somewhat higher and more globose, by having a larger 
protoconch (825-850 ^m vs. 400-425 /im), by having a 
strong spiral cord midway between the shoulder spiral 
cord and the circumbasal cord, by liavirig a strong spiral 
cord just beneath the circumbasal cord, and by having 
an open umbilicus. Differences between C. dnopherum 
and C. atlantoides and C. coronatum. are discussed in 
the Remarks in the species accounts of the latter two 
species. 

The syntype lot originally contained five specimens, 
one large, live-collected specimen and four very small 
shells. The large shell is in excellent condition, although 
the animal was removed at some point and the oper- 
culum glued to cotton in the aperture. Of the four small 
shells, one has completely disintegrated, and the other 
three are in various stages of disintegration. 

The recently collected specimen (Cloltro collection) is 
very similar to the lectotype but is slightly .smaller (7.0 
mm height, 8.5 mm width), lacks fine threads between 
the supraijeripheral spiral cords, has one fewer basal 



spiral cord, and has the inner lip of the columella rather 
strongly expanded to almost cover the umbilicus. 

Calliostoma rota new species 
(hgures 70-73, 117, 118) 

Description: Shell small, attaining height of 8.1 mm, 
width of 9.9 mm, depressed turbinate, nonumbilicate. 
Protoconch 400-425 ^m maximum diameter, 1 whorl. 
Teleoconch whorls 5.5, rapidly expanding, carinate; first 
whorl initially with 2 smooth spiral cords; adapical cord 
strong on first 2 whorls, gradually weakening to fine 
thread on next 3 whorls, finely undulate to beaded on 
last 3 whorls; abapical cord strong, smooth to weakly 
undulate on all whorls, forming carinate whorl shoulder; 
subsutural cord appearing on first quarter-whorl, becom- 
ing strong, finely beaded by third whorl; last whorl u ith 
strong, smooth, circumbasal spiral cord forming periph- 
eral carina; interspaces between spiral cords concave with 
fine, smooth or finely beaded spiral threads (last whorl 
with 11-12 between subsutural and shoulder cords. 8- 
14 between shoulder and peripheral cords); axial sculp- 
ture absent except fine growth lines. Base almost flat, 
with 17-21 weak to strong, flat, smooth spiral cords and 
0-4 fine threads in interspace between peripheral cord 
and outermost ba.sal cord; outermost cord sometimes 
torniing very weak subperipheral angulation. Umbilicus 
tilled with callus. Aperture subquadrate; outer lip thin, 
angulate; inner lip thickened, forming umbilical plug; 
columella rather short, concave in upper third, straight 
l)elov\ . Shell ground color ivory to |)inkisli tan with green 
and pink iridescence, with regularly spaced spots of light 
to dark yellow-brown on subsutural, shoulder, periph- 
eral, and 1-2 inner basal cords, basal cords sometimes 
lacking spots. 

iloloiypc: MOiU; 29.295, height 8.1 nun, width 9.3 
mm. 

Type loraliiy: Off illuibcki, illui de Sao Sebastiao, Es- 
tado (Ic S;h) I';ui1(i, Hr;i/ii, in 20 30 ni, 

Paratype: I, MCZ 258057; oil Hio de J.ineiro, Estado 
de Rio de Janeiro, Brazil, from stom;uli ol st;irfish, in 
ai)oul 16 m; Bernard 'I'ursch colleetor. 



J. F. Quiim. Jr . 1992 







^^ 



Figures 70-73. Calliostoma rota new species 70. 71. Holotr>^. MORG 29.295; height S.l mm. width 9.3 mm. 72, 73. Parat>^, 
MCZ 258057; height 7.6 mm. width 9 9 mm. Figures 74. 75. Calliostoma atlantoides new species, holot\pe, USNM S60261; height 
9.1 mm, width 10 mm Figures 76. 77. Calliostoma cvronatum new species, holotype, MC2 2T-1.56S; height 4.2 mm. width 
4.2+ mm 



Other material: 1, Coitro collection; from same lot as 
holot\-pe. 

Distribution: This species is known only from off Rio de 
Janeiro and Sao Paulo. Brazil, in 20-46 m; the single 
li\lng specimen was from 46 m. 

Remark*: Shells of Calliostoma rota are most similar 
to those of C. dnopherum but differ by ha\ing more 
whorls, b> ha\ ing a much smaller protoconch, by lacking 
a strong spiral cord between the shoulder and circum- 



basal cords, by lacking a strong subperipheral spiral an- 
gulation, and by having the umbiliciis filled with callus. 
The radula of C. rota i^formula 12-13.7.1.7.12-13; figures 
117, lis) has bvo principal characters that depart from 
the typical calliostoma tine morphology: 1" the reduced 
number of elements in the marginal tooth field; and 2^ 
the single, imcusf)ed tooth base representing the outer 
lateral tooth. The number of pairs of calliostomatine 
marginal teeth is rarely reported, but in those instances 
where the number of pairs is known ^e.g., Calvo, 1987). 



Page 102 



THE NAUTILUS, Vol. 106, No. 3 



the number is 25 or more except in Astele Swainson, 
1855, the type species of which has about 20 pairs (Clench 
& Turner, 1960:76); however, in C. rota there are only 
12 or 13 pairs. The presence of a single, uncusped outer 
lateral tooth has not been reported pre\iously in the 
Calliostomatinae, although this condition bridges the gap 
between the majority of species that have all lateral teeth 
cusped and those in the C. jujubimim species group that 
have the outer two pairs of lateral teeth uncusped. How- 
ever, although the anterior two-thirds of the radula ex- 
amined here had the uncusped lateral tooth, a few teeth 
from the posterior third retained a membranaceous, ru- 
dimentary cusp. The holotype and Coltro specimen were 
dredged from muddy sand bottom. 

Calliostoma atlantoides new species 
(6gures74, 75, 115, 116) 

Description: Shell small, attaining height of 9.1 mm, 
width of 10.0 mm, turbinate, narrowK umbilicate. Pro- 
toconch 1.10 mm maximum diameter, 1 whorl. Teleo- 
conch whorls 3.4, rapidly expanding, carinate; first whorl 
initially with 2 smooth, spiral cords, adapical cord dis- 
appearing b\ end of first whorl, abapical cord strength- 
ening and forming strong midwhorl carination on sub- 
sequent whorls; third spiral cord appearing between 
adapical cord and suture near beginning of first whorl, 
strengthening and forming rather strong subsutural an- 
gulation; both subsutural and midwhorl angulations set 
with narrow, rounded, spirally elongate beads; last whorl 
with strong, smooth spiral cord forming periphery; in- 
terspaces between spiral angulations weakly concave, with 
fine, smooth spiral threads (last whorl with 2 between 
suture and subsutural angulation, 16 between subsutural 
and midwhorl angulations, and 12 between midwhorl 
and peripheral angulations); axial sculpture of low, 
rounded riblets restricted to first 1.25 whorls. Base di- 
vided into narrow, weakly concave peripheral zone and 
broad, weakly convex central zone by strong spiral cord; 
peripheral zone with 6 spiral threads, central zone with 
about 18 weak, smooth spiral cords. Umbilicus funnel- 
shaped, broad, about 25% maximum shell diameter, wall 
strongly constricted to narrow pore. Aperture obscurely 
subquadrate, almost elliptical; outer lip thin, slightly an- 
gulate; inner lip thin and expanded above to partially 
cover umbilicus, becoming narrower and thicker below; 
columella concave in upper and lower quarters, almost 
straight in middle half, narrow above but broad and 
weakly concave at base, with prominent adapical nacre- 
ous tongue. Shell ground color silvery white w ith brilliant 
pink and green iridescence. 

Holotype: USNM 860261, height 9.1 mm, width 10 
mm. 

Type locality: West of St. Lucia, Lesser Antilles, John 
Elliott Pillsbury Station P-904, 13°45.5'N, 61°05.7'W, 
in 417-589 m. 

Distribution: This species is known only from the t\ pe 
locality. 



Remarks: In shell shape and sculpture, Calliostoma 
atlantoides is most similar to C. coronatum new species 
but differs by having a larger protoconch, by having the 
beads on the subsutural spiral cord weaker and more 
widely spaced, b\ having numerous spiral threads in the 
interspaces, by having a strongK beaded rather than 
finely undulate shoulder spiral cord, by having weaker 
and more numerous basal spiral cords, and by having 
the inner columellar lip expanded to partially cover the 
umbilicus. The shell of C. atlantoides is also ver\ similar 
to those of C. rota and C. dnophenim. It differs from 
that of C. rota by being relatively higher and more 
globose, by having a much larger protoconch, by having 
a more coarsely beaded subsutural spiral cord, b\' having 
a strongly beaded rather than smooth shoulder spiral 
cord, by having an open umbilicus, and by being silvery 
white rather than slightly brassy. The shell of C. atlan- 
toides differs from those of C. dnophenim by being 
relatively higher; by having a larger protoconch; b\ hav- 
ing a single, beaded spiral cord rather than two smooth 
spiral cords below the subsutural cord; by having finer, 
more numerous spiral threads in the interspaces; by hav- 
ing more numerous basal spiral cords; and b\ having the 
inner lip of the columella thinner and more strongly 
expanded to partially cover the umbilicus. Calliostoma 
atlantoides is similar to C. atlantis but has a much small- 
er, more globose shell; has a much larger protoconch; 
has the shoulder spiral cord located much higher on the 
whorl; has a much stronger subperipheral angulation; 
and has the inner lip of the columella thin and partially 
covering the umbilicus rather than being thickened and 
completely filling the umbilicus. The radula of C. at- 
lantoides '({ormula <20.5?.1.5^.<20; Figures 115, 116) 
is very small (length 3.1 mm) and delicate, and there 
seem to be five, thin-cusped laterals and fewer than 20 
pairs of marginal teeth, the innermost of which has a 
heavily buttressed and rather finely denticulate cusp. The 
seemingly degenerate radula of C. atlantoides is similar 
to that illustrated for an undescribed species of Callios- 
toma illustrated by Hickman and McLean (1990:fig. 71C). 

Calliostoma coronatum new species 
(figures 76, 77) 

Description: Shell small, attaining height of 4.2 mm, 
width of more than 4.2 mm, depressed turbinate, um- 
bilicate. Protoconch 850 nm maximum diameter, 1 whorl. 
Teleoconch whorls about 2.2, rapidK expanding, cari- 
nate; first whorl initially with 3 smooth spiral cords, adap- 
ical cord disappearing at whorl 1.3, midwhorl cord 
strengthening, becoming weakly undulate at whorl 1.3 
and forming strong midwhorl angulation, abapical cord 
remaining weak on all whorls; fourth spiral cord ap- 
pearing between adapical cord and suture at whorl 0.3, 
rapidly strengthening, undulate on last half of first whorl, 
undulations becoming strong, closel\- .set, upturned, tri- 
angular beads; last whorl with strong, smooth spiral cord 
forming periphery; interspaces smooth except for weak 
spiral cord between midwhorl and peripheral cords; axial 



J. F. Quinn, Jr.. 1992 



Page 103 




Figures 78, 79. Calliostotna rugosum new species, holot\pe. USNM 860262; height 23.5 mm, width 24. S mm Figures 80, 81. 

Calliostoma torrei Clench & Aguayo, 1940, holotype, MCZ 135165; height 40.8 mm, width 36.8 mm. 



sculpture absent except for \o\\. rather broad, rounded 
folds on last whorl. Base divided into narrow, smooth, 
concave peripheral zone and broad, almost flat central 
zone by strong, smooth spiral cord; central zone with 12 
narrow, smooth spiral cords, adaxial 3 somewhat stronger 
and more widely spaced. Umbilicus rather narrow, less 
than 20% maximum shell diameter, constricted v\ithin 
to small pore. Aperture probably obscurely subquadrate 
or ovate, outer lip broken; inner lip thin, narrow, slightly 
reflected over umbilicus; columella convex, thin. 

Holotype: MCZ 274568, height 4.2 mm, width 4.2 + 
mm. 

Type locality: Off Joao Pessoa, Brazil. Ch.\in Cruise 35, 

Station 12, 7°09.0'S, 34°25.5'W, in 768-805 m. 



Distribution: 

locality. 



This species is known only from the type 



Remarks: Calliostoma coronatum is similar to C. dno- 
pherum but differs by having a relatively smaller, lower- 
spired shell that has a much weaker spiral cord between 
the shoulder and peripheral cords, that lacks spiral threads 
between the primary spiral cords, and that has a pro- 
portionately narrower umbilicus. 

Calliostoma rugosum new species 
(figures 78, 79) 

Description: Shell medium-sized to large, attaining es- 
timated height of 25 mm. estimated width of 31 mm. 
trochoid, nonumbilicate. Protoconch worn, at least 550 
Hm maximum diameter. 1 whorl. Teleoconch whorls 
about 7.5 (last 0.5 whorl broken), flat to weakly concave; 
first 2 whorls worn and partially decorticated; whorls 3- 
4 with 5 strong, subequal, beaded spiral cords, adapical 
cord forming weak subsutural angulation, abapical cord 
forming upper limit of flat periphery bearing 1 spiral 
thread; subsutural and upper peripheral cords each split- 
ting into 2 subequal spiral cords on whorls 5-6; spiral 
cords between subsutural and upper peripheral cords 
increasing to 7 by end of seventh whorl; peripheral zone 
with 4 spiral cords by end of seventh whorl, periphery 



becoming rounded on last whorl; all cords set w ith rather 
strong, rounded, generally spirally elongate beads. Base 
weakly convex, with 19 strong, smooth, subequal spiral 
cords. Umbilical area filled with irregularly ridged callus. 
.Aperture subquadrate; outer lip broken; iiuier lip thick- 
ened; columella concave, broken basallv. 



Holotype: USNM 860262, 
24.8+ mm. 



height 23.5+ mm, width 



Type locality: Straits of Florida, J.^mes M. Gillis Cruise 
7307, Station 13, depth and exact locality unknown. 



Distribution: 

localitv . 



This species is known only from the type 



Remarks: The holoty pe of Calliostoma rugosum is very 
similar to that of C. torrei Clench and Aguayo, 1940 
(figures 80, 81), from off Cardenas, Cuba, but differs by 
being relativ ely much broader; by having a weaker, more 
abapical upper peripheral cord; by having coarser, more 
rounded spiral cords; and by having larger, rounded, 
spirally elongate beads on the cords. The protoconch of 
C. rugosum (estimated to be more than 550 ^m) may 
be larger than that of C. torrei (about 525 ^m), and the 
subsutural and upper peripheral cords of C. rugosum 
split into two rather coarse cords rather than three fine, 
sharp cords as in C. torrei. Both C. rugosum and C. 
torrei, based on similarities of shape and sculpture, seem 
to be closely related to C. caribbcanum \\'eisbord, 1962, 
a fossil species from the Pleistocene Mare Formation of 
\enezuela. 

Calliostoma argentum new species 

(figures 82-85) 

CaliiiKloma tnarionac. Sander & Laili, 1982:table 4. (Non Cal- 
liostoma marionae Dail. 1906). 

Description: Shell medium-sized, attaining height of 
25.6 mm, width of 28.2 mm, conical, umbilicate, finely 
sculptured to almost smooth. Protoconch about 375 jum 
maximum diameter, 1 whorl Teleoconch whorls about 
9.5, flat to verv weakly convex; first 2 whorls with low, 
rounded axial riblets and 2-3 finely beaded spiral cords; 



Page 104 



THE NAUTILUS, Vol. 106, No. 3 




> 



J 









Figures 82-85. CUitliostoma argentuni new species. 82. 83. Holotype, USNM 860263; height and width 18.1 mm. 84, 85. 
Specimen from off Barbados, Sunderland collection; height 22.5 mm, width 24.2 mm Figures 86, 87. Calliostoma jeanneae Clench 
& Turner, 1960, holotype, MCZ 228370; height 11.2 mm, width 13.2 mm. Figures 88, 89. Calliostoma axelolssoni. new name, 
specimen from off Mostardas, Brazil, MORG 18.738; height 30 4 mm, width 34 2 mm 



axial sculpture absent on subsecjuent w liorls; spiral cords 
increasing by intercalation to 9-1 1, beading disappearing 
on fourth or fifth whorl but sometimes reappearing on 
eighth or ninth whorl; periphery sharply carinate, smooth, 
sometimes reflected adapically. Base flat to weakly con- 
vex, with 13-15 weak, broad, smooth spiral cords, adaxial 
2-3 cords slightly stronger, weakly beaded; circumum- 
bilical cord strong, coarsely beaded. Umbilicus funnel- 



shaped, 14%-17% maximum shell diameter, wall almost 
vertical, white. Aperture subquadrate, slightly thickened 
within and with several low, rounded ridges, lips thin; 
columella rather long, weakK concave, slightly thick- 
ened, terminating in small, rounded tubercle. Shell ground 
color cream to light tan w ith pale orange-brown patches 
below suture and spots on periphery; base uniformly 



J. F. Quinii, Jr., 1992 



Page 105 



Holotype: USNM 860263, height and width 18 1 mm. 

Type locality: Arrovvsmith Bank, (^uintana Roo, Mex- 
ico, John Elliott Pillsbury Station P-594, 21°00.5'N, 
86°23.0'W, in 307-329 m. 

Paratypes: 1, UMML 30.5583; off Arrowsmith Bank, 
Mexico, John Elliott Pillsbury Station P-584, 
21°02.0'N, 86°24.0'W, 353-347 m; 23 May 1967; 10-ft 
otter trawl.— 1, ANSP 353529; off Holetovvn, Barbados; 
F. Sander, collector. 

Other material: 1 fragment, UMML 30.8372; off St. 
Vincent, John Elliott Pillsbury Station P-877, 
13°16.7'N, 6r05.6"W, 348-466 m; 6 July 1969; 5-f t Blake 
trawl. — 3, Sunderland collection; off St. James, Barbados, 
175 m; dredge; F. Sander, collector. 

Distribution: Shells of Calliostoma argentum are known 
from off the Yucatan Peninsula. Mexico, and St. Vincent 
and Barbados, Lesser Antilles, in depths of 175-466 m. 

Remarks: Shells of Calliostoma argentum are most sim- 
ilar to the holotype of C. jeanneae (figures 86, 87), from 
off Havana, Cuba, but differ by being relatively narrow- 
er; by having spiral cords that are strongly beaded until 
the middle of the fourth or fifth whorls rather than be- 
coming smooth near the beginning of the third whorl; 
b> lacking a strong spiral cord just above the peripher\ 
on the third through sixth whorls; by having an umbilicus 
that is open rather than filled with callus; by having a 
strongly beaded rather than smooth circumumbilical cord; 
and by having a longer, less thickened and oblique col- 
umella. Perhaps of less importance are differences in the 
color patterns: that of C. argentum consists of cream to 
light tan spiral cords separated by darker-colored striae 
and pale orange-brown patches below the sutures and 
somewhat darker spots on the periphery; that of C. jean- 
neae consists of rows of golden-brown spots on the spiral 
cords above and below the periphery and brighter spots 
of the same color on the periphery. Both C. argentum 
and C. jeanneae are similar to the eastern Pacific C. 
platinum, C. chilena, and C. titanium, and all five spe- 
cies may be closely related to C. metalium Woodring, 
1957, from the late Miocene to early Pliocene Chagres 
Sandstone of Panama. 



Calliostoma axelolssoni new name 
(figures 88, 89, 119) 

Calliostoma olssoni Ba\er, 1971:118, fig. 4 (left). — Kaicher, 

1980:card no. 2239. (Non Calliostoma olssoni Maury, 1925). 
Calliostoma (Kombologion) rosewateri: Rios, 1975:23, pi 5, 

fig. 59— Calvo, 1987:63, 65, fig. 28. (Non Calliostoma 

rosewateri Clench & Turner, 1960). 
Calliostoma [Kombologion) bairdi rosewateri: Rios, 1985:22, 

pl.9, fig. 85. 

Description: See Ba\er (1971:118) for description of 
shell. Animal (in alcohol) white; mantle edge smooth; 
cephalic tentacles moderately long, slender, gradually 
tapering, right longer than left, ocular peduncles rather 



long (about 35% tentacle length), broad, with large, black 
eye at tips; snout long, broad, slightly expanded at tip, 
tip fringed with small, short papillae; epipodium with 4 
pairs of tentacles decreasing in size posteriorly, neck lobes 
well-developed, semicircular, smooth. 

Holotype: USNM 700002, height 16.8 mm, width 21.0 



Type locality: Southwest of St. N'incent, Lesser Antilles, 
John Elliott Pillsbury Station P-876, 13°13.9'N, 
6r04.7'W, in 231-238 m. 

Other material: 1, UMML 30.8373; off Dominica, John 
Elliott Pillsbury Station P-931, 15°31.2'N, 61°12.3'W, 
174-357 m; 15 July 1969; 5-ft Blake trawl.— 1, MORG 
18.738; W. Besnard station, off Mostardas, Rio Grande 
do Sul, Brazil, 230 m; 1972. 

Distribution: This species is now known from off Dom- 
inica and St. Vincent in the Lesser Antilles, and off south- 
ern Brazil, in 174-357 m. 

Rema.»-ks: Calliostoma olssoni Bayer, 1971, is preoc- 
cupied by C. olssoni Maury, 1925, a species (probably 
assignable to Calliomphahis Cossmann, 1888) from the 
Mio-Pliocene formations of Trinidad (Maury, 1925; Jung, 
1969). With F. M. Bayer's permission, 1 offer a replace- 
ment name that preserves his original intent to honor 
the late Dr. Axel A, Olsson. 

The Brazilian specimen (misidentified as C. rosewateri 
by Rios, 1975, 1985) (figures 88, 89) is much larger (height 
30.4 mm, w idth 34.2 mm) than the holot> pe, has a nar- 
rower umbilicus that has been partially filled with callus, 
has three weak circumumbilical and six weak subperi- 
pheral spiral cords, and has a more diffuse and subdued 
color pattern except for the pink spots on the periphery. 
The Dominican specimen is very similar to the holotype 
but is shghtly larger (height 19.9 mm, width 24.1 mm), 
has a slightly narrower umbilicus, and has a much less 
vivid color pattern. The radula from this specimen (figure 
119) conforms well with Calvo's illustration (1987:fig. 
28) but the lateral tooth cusps are longer than he de- 
picted. 

Based on shell characters, C. axelolssoni seems to be 
closely related to C. bigeloivi, C brunneum, and C. 
hendersoni — the four species forming a rather distinct 
subgroup of Calliostoma. This subgroup may be most 
closely related to the C. bairdii species group. 

Calliostoma aurora Dall, 1888 
(figures 90, 91, 120) 

Calliostoma aurora Dall, 1888:68, fig. 285; 1889a:366, pi. 37, 
fig.2— Clench & Turner, 1960:64, pi. 45, figs. 1, 2. 

Description: Shell: see Dall (1889a:366) and Clench and 
Turner (1960:64). Animal (reconstituted in trisodium 
phosphate): foot mottled with red-brown, mantle edge 
with light, transverse band and long, median longitudinal 
streak of red-brown; cephalic tentacles very long, slen- 
der; eyestalks very short (less than 10% tentacle length) 



Page 106 



THE XAUTILIS. \ol. 106, No 3 





Figure? 90. 91. Calliostoma aurora Dall, 1888. specimen from off Guadeloupe, USXM 860504; height 25.4 mm, width 27 8 mm. 
Figures 92. 93. Calliostoma guesti new species, holotype, DMNH 96994; height and width 27.5 mm. 



with large, black eye at tips; snout slightly longer than 
broad, tip fringed by rather long, stout papillae; epipo- 
dium with 3 or 4 pairs of very long, slender tentacles; 
neck lobes very well-developed, seemingK smooth-edged. 

Holotype: MCZ 73808. height 21.0 mm, width 26,5 mm. 

Type locality: Off Grenada, Lesser .Antilles, Bl.\ke Sta- 
tion 26.5, 12°03'55"N, 6r49'40"W, in 1054 m. 

Other material: 1, USNM 860504; off Great Inagua 
Island, Bahama Islands. Johnson-Se.^-Link I Station JSL- 
1-2323, 2r01'45"N', 74''43'48"\V, 628 m; 15 October 
1988, — 1, Dan collection; off Barbados, Lesser .\ntilles, 
183 m. 

Distribution: Calliostoma aurora is now known from 
the southeastern Bahama Islands and from off Barbados, 
in depths of ia3-1054 m. 

Remarks: The present specimens agree well with the 
holotype in most shell characters, but are larger (Great 
Inagua; height 25.4 mm, width 27.8 mm; Barbados: height 
23.5 mm, width 27.2 mm;; have whorls with flatter pro- 
files; and have numerous very fine spiral striae on the 
outer two-thirds of the base, the striae becoming deeper 
on the adaxial third and forming several indistinct and 
three distinct spiral cords near the center. The ground 
color of the new shells is a dark golden brown with 
alternating spots of white and brown on the periphery; 
the base of the Barbados specimen has faint, crescent- 
shaped streaks of pale brown; and the umbilical callus 
of both specimens is white. 

.Although Clench and Turner (1960) compared the 
shell morphology of C, aurora to that of C. hairdii and 
C. psyche, the relationship of this species to other Cal- 
liostoma species is unclear. The radula of C. aurora 
(formula oo,10,l,10.oo; figure 120) indicates that the spe- 
cies is not very closely related to the C. hairdii species 
group or to any other western Atlantic species group. 
The most notable difference is that C, aurora has ten 
pairs of lateral teeth, whereas most other species have 
only four to seven pairs; only Calliostoma militare (22 
pairs; Castellanos & Fernandez, 1976; Calvo, 1987) and 
Venustatrochus georgianus (16 pairs; Powell, 1951) have 
more lateral tooth pairs. The inner marginal teeth of C. 
aurora sltc also rather unusual: the innermost marginal 



is rather slender with a sharply curved cusp and ten 
denticles, of which the terminal one is unusualK' long, 
and the ne.xt several teeth have a very long terminal 
denticle with four to six secondary denticles about a third 
of the wa> down the shaft. 

The Barbados specimen was collected alive from a 
depth of 183 m, which is shallower than, but comparable 
to, the provenance of the paratype lot (from Barbados 
in 256 m), suggesting that the occurrence of the holot>pe 
in 1054 m was adventitious. The specimen from Great 
Inagua was collected from atop a small, blue sponge (M. 
G, Harasewych, personal communication), and its intes- 
tine was packed with a dark brown, rather solid mass of 
organic material in which were embedded numerous 
sponge spicules and poK chaete setae, suggesting that this 
species feeds on sponges, 

Calliostoma guesti new species 

(figures 92, 93) 

Description: Shell medium-sized, attaining height and 
width of 27.5 mm, conical, nonumbilicate, finely sculp- 
tured. Protoconch about 425 nm maximum diameter, 1 
whorl. Teleoconch whorls more than 9.1, flat to weakly 
convex; first 2 whorls with low, rounded axial riblets and 
2-3 beaded spiral cords; axial sculpture absent on sub- 
sequent whorls; spiral cords increasing by intercalation 
to 12, finely beaded throughout; periphery sharp, nar- 
rowly bicarinate, peripheral cords adpressed, adapical 
peripheral cord distinctly stronger than abapical periph- 
eral cord. Base weakly convex, with numerous fine striae 
on outer three-fourths and 3-4 weak, weakly beaded 
spiral cords near center. Umbilicus filled with white cal- 
lus. Aperture subquadrate, lips thin, weakly crenulate; 
columella rather short, weakly concave, thickened. Shell 
ground color ivory with irregular, diffuse patches of gold- 
en brown above periphery, somewhat more discrete spots 
of same color on periphery; base with very faint, cres- 
cent-shaped, radial streaks of light brown and sometimes 
a few spots on 1-2 spiral cords, 

Holotype: DMNH 96994, height and width 27.5 mm. 

Type locality: 2.5 mi off south shore of Bermuda, in 
183-219 m. 



J. F. Quinn, Jr.. 1992 



Paee 107 






96 



97 



Figures 94-97. Calliostoma stirophorum (Watson. 1879). 94. 9.5. Holot\ pe of Trochus (Zizyphinus) stirophonis Watson. 1879. 
BM(\H) 1887.2.9,210; height 7.5 mm, width 6 6 mm 96. 97. Holotype of Calliostoma arestum Dall, 1927. ISN'M 108412; height 
5.4 mm. width 5.0 mm. 



Paratypes: 3. DMXH 1S75SS; 1. USXM S60270; from 
same lot as holot\ pe. 

Distribution: This species is known onK from the t\ pe 
locality. 

Remarks: Shells of C.alliostoma guesti are most similar 
to those of C. psyche but differ b\ being relati\ eK nar- 
rower; b\ having more fineK beaded spiral cords; b\ 
having the supraperipheral spiral cord on all whorls 
sharply beaded and stronger than the subsutural cord 
rather than being smooth on the first two to three whorls 
and weaker than the subsutural cord; b\ ha\ing the pe- 
riphery composed of tw o adpressed spiral cords, of which 
the adapical cord is distinctly stronger, rather than hav- 
ing the cords subequal and separated b\ a concave in- 
terspace that is as wide or w ider than the cords; h\ ha\ing 
much weaker spiral and radial sculpture on the base; 
and by having a color pattern of light brow n rather than 
rose, particularly on the peripher\ . 

Calliostoma guesti is named in honor of Mr .\rthur 
Guest for his contributions to the knowledge of the mol- 
luscan fauna of Bermuda. 

Calliostoma stirophorum (Watson, 1879) 
new combination 
(figures 94-97) 

Trochus {Zizitphinus) stirophorus Watson, 1879:695; 1886:59, 

pi. 6, fig. 2. 
Calliostoma arestum Dall, 1927b 127. — Johnson, 1934:70, — 

Clench & Turner, 196079 — .\bbott, 1974:46, 

Description: See Watson (1879:695; 1886:59), 

Holotypes: BM{NH) 1887.2.9.210. height 7.5 mm, width 
6,6 mm (Trochus (Zizyphinus) stirophorus); USNM 
108412, height 5.4 mm. w idth 5.0 mm (Calliostoma ares- 
tum). 

Type localities: Off Culebra Island. \'irgin Islands, 
c'h.allenger Station 24. 18''38'30"N, 64°05'3b"W. in 713 
m (Trochus (Zizyphinus) stirophorus); off southern 
Georgia, .-^lb.^tross Station 2415. 30°44'00"N, 
79°26'00"W. in 805 m (Calliostoma arestum). 



Distribution: This species is know n onlv from the two 
t\ pe specimens, one from off Fernandina. Florida, in 805 
m. and off Culebra Island, Virgin Islands, in 713 m. 

Remarks: Both Watson (1879, 1886) and Dall (1927b) 
based their respective species descriptions on unique 
specimens that w ere empt\ and somew hat damaged w hen 
collected. The specimen of Calliostoma arestum ifigures 
96. 971 is slightK smaller, has fewer whorls, and is more 
w eathered than that of Trochus (Zizyphinus) stirophorus 
vfigures 94, 95), but both shells seem to represent the 
same species. These two shells are most similar to those 
of C. circumcinctum but differ principalK b> ha\ing 
three strong, rounded, beaded spiral carinae above the 
suture rather than the two ver\ strong, blade-like carinae 
of C. circumcinctum; the shells of C. stirophorum are 
also proportionateK broader than those of C. circum- 
cinctum. The shells of both C. stirophorum and C. cir- 
cumcinctum are ver\ similar to those of Otukaia hlacki 
(Dell. 1956) from New Zealand: Calliostoma (Otukaia) 
delli McLean and .\ndrade. 19S2; from off Chile, and 
Otukaia eltanini Dell. 1990. from .Antarctica, and could 
be included in the genus or subgenus Otukaia Ikebe. 
1942. if that ta.xon gains general acceptance. Some recent 
authors (Powell, 1979; McLean & .\ndrade, 19S2; Dell, 
1990) are now using Otukaia at either the genus or sub- 
genus level. 



Calliostoma adspersum (Philippi, 1851) 
(figures 98-100, 121) 

Trochus eximius Philippi, 1844 Un 1842-1851). vol. 1. pt. 6: 
17. pi 4. fig 7 (Non Trochus eximius Reeve, 1843). 

Trochus adspersus Philippi. 1851 ^in 1846-1S55):217. pi. 32 
fig. 8. 

Calliostoma aspersum. Lange de Morretes. 1949:58. — Calvo. 
1987:63. fig. 27. (Unjustified emendation). 

Calliostoma adspersum: Calvo. 1987:63. fig. 27. 

Calliostoma [Koml>ologion) adspersum . Clench & Turner, 1960: 
46. pi. 30. figs. 1. 2 (partim).— Rios. 1970:25. pi. 4 (lower 
right) (partim); 1975:23. pi. 5. fig. 58; 1985:21. pi. 9, fig. 
84; 1990:9, 10 (photographs). 



Page 108 



THE NAUTILUS, Vol. 106, No. 3 




Figures 98-100. Calliostoma adspersum (Philippi, 1851), specimens from off Cabo de la Vela, Colombia. 98, 99. FSBC 1 34044; 
height 17.9 mm, width 18.2 mm, 100. Deynzer collection; height 16.3 mm, width 15.6 mm Figures 101, 102. Calliosloma 
depictum Dall, 1927, specimen from off Salvador, Brazil, FSBC I 44110; height 10.3 mm, width 9.9 mm. 



Description: See Clench and Turner (1960:46) and Rios 
(1990:9) for descriptions of shells. Animal (in alcohol): 
foot red-brown; mantle with narrow band of white at 
edge, behind which is broad, irregularly edged band of 
brown; cephalic tentacles long, slender, evenly tapered, 
ocular peduncles stout, about 25% tentacle length, with 
large, black eye at tips; snout long, broad, expanded at 
tip, red-brown, with anterior fringe of long papillae; 
epipodium w ith 3 pairs of red-brown tentacles, those on 
left larger, neck lobes well-developed, semicircular, left 
lobe finely fringed, right lobe smooth. 

Holotype: Philippi's type material could not be located 
in the Museum fiir Naturkunde der Humboldt-Univer- 
sitat zu Berlin (R. Kilias, in litt.), and although it may 
be present in Chile, I consider it to be lost. 

Type locality: Restricted by Clench and Turner (1960; 
47) to Praia do Leste, Ihia Guaiba, Estado de Rio de 
Janeiro, Brazil. 

Other material: 1, UMML 30.8368; off Cabo de la Vela, 
Colombia, John Elliott Pillsbury Station P-780, 
11°39.0'N, 73°08.5'W, 18-27 m; 30 July 1968; 10-ft otter 
trawl.— 1, FSBC I 34044; 21, Devnzer collection; off 



Cabo de la Vela, Colombia; 1990; shrimp trawlers. — 1, 
UMML 30.7156; off Cabo de la \'ela, Colombia, John 
Elliott Pillsbury Station P-767, 12°I6.1'N, 71°03.3'W, 
24-26 m; 28 July 1968; 10-ft otter trawl.— L UMML 
30.7040; off Peninsula de Paraguana, Venezuela, John 
Elliott Pillsbury Station P-758, 11°42.2'N, 69°40.0'W, 
15-18 m; 27 July 1968; 10-ft otter trawl— 1, UMML 
30.6888; off Tucacas, Venezuela, John Elliott Pillsbury 
Station P-750, 10°36.1'N, 68n2.2'W, 22-26 m; 25 July 
1968; 10-ft otter trawl —2, UMML 30.6617; off Isla de 
Margarita, N'enezuela, John Elliott Pillsbi:ry Station 
P-721, 11°06.5'N, 64°22.5'\V, 26-27 m; 21 July 1968; 10- 
ft otter trawl— 1, UMML 30.6183; off Georgetown, Guy- 
ana, John Elliott Pillsbury Station P-686, 7°00.0'N, 
57°08.0'W, 27-26 m; 15 July 1968; 10-ft otter trawl — 
2, UMML 30.5931; off Cayenne, French Guiana, John 
Elliott Pillsbury Station P-655, 6°07.0'N, 53°39.0'W, 
26 m; 9 July 1968; 10-ft otter trawl.— 4, FSBC I 44109; 
off Guarapari, Estado de Espirito Santo, Brazil, 18-20 
m; March 1991; ex J. & M. Coltro. 

Distribution: Calliostoma adspersum w as thought to be 
endemic to Brazil (Clench & Turner, 1960; Rios, 1970, 
1975) until Rios (1985) reported its occurrence in Suri- 



J. F. Quinn, Jr., 1992 



Page 109 






105 



106 



Figures 103-106. Calliostoma scurra new species. 103, 104. Holotype, USNM 860264; height 14.1 mm. width 13 3 mm. 105. 
106. Paratype from John Elliott Pillsblry Station P-834, FSBC I 44073; height 13.9 mm, width 12.7 mm. 




Figures 107-1 II. Radulae of Calliostoma species. 107. Callidstiiuta < nulnphilum new species, ex paratype, FSBC I 44070; 95 x. 
108. Calliostoma viscardii new species, ex holotype, MORG 29,292, 310 x . 109. Calliostoma moscatellii new species, ex specimen 
from off Rio de Janeiro, Brazil; 65 x , 1 10, 111. Calliostoma scalenum new species, ex specimen from Campeche Bank, Yucatan, 
FSBC I 32311; 235 x and 435 x, respectively 112. Calliostoma jujtibinum (Gmelin, 1791), ex specimen from Cat Island, Bahama 
Islands, FSBC I 15698; 125 x 113. Calliostoma cubense new species, ex paratype from off Peninsula de Hicacos, Cuba, FSBC 1 
32405; 245 X. 114. Calliostoma vinosum new species, ex holotype, MORG 29.294; 280x. 



Page 110 



THE NAUTILUS, Vol. 106, No. 3 




Figures 115-122. 



Radulae of Calliostoma species 



115, 



116. Calliostoma atlantoides new species, ex holotype, USNM 860261; 
75 X and 160 x, respectively 117, 118. Calliostoma rota new species, ex paratype, MCZ 2580.57; .50 x and 115x, respectively. 
119. Calliostoma axelolssoni new name, ex specimen from off Dominica, UMML 30.8.373; 65 x. 120. Calliostoma aurora Dall. 
1888, ex specimen from Guadeloupe, USNM 860504; 70x. 121. Calliostoma adspersum (Philippi, 1851), ex specimen from off 
Guarapari, Brazil, FSBC 1 44108; 95 x. 122. Calliostoma depietum Dall, 1927, ex specimen from off llhabela, Sao Paulo, Brazil, 
FSBC 1 44109; 190 X. 



name. The present material establishes the presence of 
the species along the entire northern coast of South Amer- 
ica, principally in depths of about 15-30 m. 

Remarks: As noted by Rios (1975, 1985, 1990), the 
names Calliostoma adspersum and C. depietum (figures 
101, 102) do not represent the same species, as was con- 
tended by Clench and Turner (1960). Shells of C. ad- 
spersum attain a larger size; have all whorls with a dis- 
tinct shoulder angulation and fiat-sided periphery; have 
more numerous, more unequally sized spiral cords; and 
have more diffuse color patterns than those of C. depie- 
tum. One juvenile specimen (UMML 30.7156; height 6.5 
nun, width 7.3 mm) differs from larger shells by having 
an open umbilicus whose wall narrows rapidlv from 1.6 
mm to a pore-like opening onK 0.3 mm wide; larger 
specimens have the umbilicus filled with callus. The rad- 
ula of C. adspersum (figure 121) differs from that of C. 
depietum (figure 122) by having a more broadly cusped 
rachidian, by having fewer pairs of lateral teeth (4 vs. 



5, respectively), and by having m.ore numerous (about 
40 vs. about 25) marginal tooth pairs whose cusps are 
narrower and have more slender denticles. 

This species seems to occur most frequentK in areas 
of "hard bottom" in the western part of its range (John 
Elliott Pillsbury field data log) and on sandy mud 
bottom in the eastern and southern part (Rios, 1985; J. 
& M. Coltro, personal communication). Calliostoma de- 
pietum lives in depths of 1-7 m in sandy areas aroimd 
rocks and corals (Rios, 1990), where it is often found 
living under an unidentified sea urchin (J. &. M. Coltro, 
personal communication); the species is also found on 
Sargassum (Rios, 1985, 1990). 



Calliostoma seurra new species 
(figures 103-106) 

Description: Shell medium-sized, attaining height of 
14.1 mm, width of 13,3 mm, conical, nonumbilicate. 



J. F. Qiiinii, Jr., 1992 



Page 111 



rather coarsely sculptured. Protoconch 40U— 125 ^lm ma.\- 
imum diameter, 1 whorl. Teleoconch whorls about 7, 
weakly convex; first 4 whorls with niunerous low , round- 
ed axial riblets and 2-5 fineK beaded spiral cords; axial 
sculpture absent on subsequent whorls; spiral cords in- 
creasing by intercalation to 10-15, usualK alternating in 
size, set with crowded, rounded beads; periphery weakly 
angulate, composed of pair of adpressed spiral cords, 
adapical cord stronger, beaded, abapical cord smooth. 
Base convex, with 12-19 strong, narrow, smooth spiral 
cords, interspace between abaxial 2 cords usually with 
1-5 fine spiral threads and next adaxial interspace usually 
with 1 thread. Umbilicus filled with callus, sliallnw de- 
pression sometimes present. Aperture subquadrate, 
thickened within, often with channel corresponding to 
periphery, lips thin, weakly crenulate; columella very 
weakly concave, greatly thickened, with tongue of nacre 
extending outward to umbilical callus. Shell ground color 
cream with spots and irregular patches ol reddish brow n 
above periphery and irregularly crescentic streaks of same 
color on base. 

Holotype: USNM 860264, height 14.1 mm, width 13.3 
mm. 

Type locality: Off mouth of Orinoco River, Venezuela, 
John Elliott Pillsburv Station P-834, 9°04.1'N, 
60°10.7'W, in 33-35 m. 

Paratypes: 1, USNM 860265; 1, FSBC I 44073; 1, UP 
189457; 1, MCZ 302590; 1, ANSP 389338; 1, LACM 
2253; 4, UMML 30.7225; all from same lot as holotype. 

Other material: 1, UMML 30.6529; off Isla de Mar- 
garita, Venezuela, John Elliott Pillsbury Station P-718, 
ll''22.5'N, 64°08.6'W, 60 m; 20 July 1968; 10-ft 
otter trawl.— 1, UMML 30.6374; off Peninsula de Paria, 
Venezuela, John Elliott Pillsblry Station P-705, 
10°45.0'N, 62°00.0'W, 77-86 m; 18 Julv 1968; 10-ft otter 
trawl.— 6, UMML 30.6359; Gulf of Paria, Venezuela, 
John Elliott Pillsbury Station P-704, 10°34.3'N, 
61°57.0'W. 18 m; 18 July 1968; 10-ft otter trawl.— 1, 
UMML 30.6339; off Orinoco River, Venezuela, John El- 
liott Pillsbury Station P-696, 8°38.0'N, 58°56.0'W, 55- 
59 m; 16 July 1968; 10-ft otter trawl.— 1 + 1 fragment, 
UMML 30.5979; off Parimaribo, Suriname, John El- 
liott Pillsbury Station P-663, 6°29.0'N, 54°41,0"W, 24 
m; 10 July 1968; 10-ft otter trawl. 

Distribution: Calliostoma scurra has been collected from 
off Suriname northwestward to off Isla de Margarita, 
Venezuela, in depths of 18-86 m; live-collected speci- 
mens are known from depths of 18-35 m 

Remarks: Calliostoma scurra does not seem to be close- 
K related to any other western Atlantic species except, 
perhaps, C. adspersum. Shells of the latter species differ 
from those of the former by having a broad, flat periph- 
ery; by having distinct axial rugae in the interspaces 
between the primary spiral cords; by having finer, more 
numerous, more distinctly beaded supraperipheral spiral 



cords; by having a more oblique columella; and by hav- 
ing a different color pattern. 

The field data indicate that this species lives on bottoms 
composed of mud, shell hash, and coral rubble. 

ACKNOWLEDGMENTS 

I thank the following for access to collections under their 
care: George M. Davis and Robert Robertson (ANSP); 
Richard S. Houbrick, M. G. Harasewych, and the late 
Joseph Rosewater (USNM); Kenneth J. Boss, Silvard P. 
Kool. and Ruth D. Turner (MCZ); Riidiger Bieler and 
Russell Jensen (DMNH); Nancy A. and the late Gilbert 
L. Voss (UMML); John D. Taylor and Kathie M. Way 
(BM[NH]); James H. McLean (LACM); Fred G. Thomp- 
son and Kurt Auffenberg (UF); Constance E. Boone and 
the late Thomas E. Pulle> (HMNS); and Eliezer de C. 
Rios (MORG). I am especialK grateful to Kevan and 
Linda Sunderland for their hospitality, for allow ing me 
to examine their collection that contains much important 
material, and for generously donating type or \oucher 
specimens. Jose and Marcus Coltro were instrumental in 
obtaining Brazilian material and generousK' donated type 
material. Jose Leal (UMML) also provided material col- 
lected from seamounts off southeastern Brazil. Arthur 
Guest and Jack Lightbourn kindly provided the type 
material of the Bermudan species. Albert E. and Beverly 
A. Deynzer generously donated material of Calliostoma 
purpureum. Harry G. Lee, Donald Dan, and Leslie A. 
Crnkovic kindly allowed me to examine specimens in 
their collections. Rudolf Kilias (Museum fiir Naturkunde 
der Humboldt-Universitat zu Berlin) provided infor- 
mation on Philippi's type material. Sally D. Kaicher kind- 
ly provided photographs of the types of Watson's species. 
Thomas H. Perkins. Da\ id K. Camp. William G. Lyons 
(all Florida Marine Research Institute), and two anon- 
ymous review ers provided helpful comments on previous 
versions of this paper. 

LITERATURE CITED 

.'\bbott, R. T. 1974. American seasheils. 2nd ed. Van Nostrand 
Reinhold. New York. 663 p. 

Adams, C. B. 1850. Descriptions of supposed new species of 
marine shells which inhabit Jamaica. Contributions to 
Conchology l(5):69-75. 

Adams. H. and A, .\dams. 1854, The genera of Recent Mol- 
lusca. Vol ] John Van Voorst, London, 484 p. 

Bayer, F. M. 1971. New and unusual mollusks collected bv 
R/V JOHN ELLIOTT PILLSBURY and R/V GERDA in 
the tropical western .Atlantic. Bulletin of Marine Science 
21(l):lll-236. 

Bayer, F. M., G. L. Voss, and C. R. Robins. 1970. Bioenvi- 
ronmental and radiological safety feasibility studies .\t- 
lantic-Pacific interoceanic canal. Report on the marine 
fauna and benthic shelf- slope communities of the isthmian 
region. Processed report (University of Miami). Battelle 
Memorial Institute. 99 -I- vi -I- 311 -I- [70] p. 

Calvo, I. S. 1987. Radulas de gastropodes marinhos brasileiros. 
Rio Grande do Sul. Editora da Funda(jao llniversidade do 
Rio Grande, 201 p. 



Page 112 



THE NAUTILUS, Vol. 106, No. 3 



Castellanos, Z. and D. Fernandez 197(5. Los generos Cal- 
liosloma y Neocalliostoma del Mar Argentine. Revi.sta del 
Museo de La Plata 12(116):135-156. 

Clench, W. J. and C. J. Aguayo. 1938. Notes and descriptions 
of new species of Calliostoma, Gaza and Columbarium 
(Mollusca) obtained by the Harvard-Habana E.xpedition 
off the coast of Cuba. Memorias de la Sotiedad de Historia 
Natural 12(5):375-384. 

Clench, VV. J. and C. J. .\guayo. 1940. Notes and descriptions 
of new deep-water Mollusca obtained by the Harvard- 
Habana E.xpedition off the coast of Cuba. III. Memorias 
de la Sociedad de Historia Natural 14(l):77-94. 

Clench, W. J. and R. D Turner. 1960. The genus Ca//ios<oma 
in the western Atlantic. Johnsonia 4(40): 1-80. 

Conrad, T. A. 1846. Descriptions of nineteen new species of 
fossil and Recent shells and corals of the United States. 
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THE NAUTILIIS 106(3):115-118, 1992 



Page 115 



A New Species of Pseiidori77iula 
(Fissurellacea: Clypeosectidae) from 
Hydrothermal Vents of the 
Mid-Atlantic Ridge 



James H. McLean 

Los Angeles C^iuinh Miiseiiin of 

Natural Histor\ 
900 Exposition Houle\ard 
Los Angeles, CA 90007, USA 



ABSTRACT 

Pseudorimula midatlantica new species is described from the 
Snake Pit hydrothermal lield on the Mid-Atlantic Ridge It is 
the second member of its genus, the type species being known 
Irom the Mariana Trough h)drothermal vents in the mid-Pa- 
cific It differs from the type species in its hypertrophied de- 
velopment of the gonad, which displaces part of the space 
normally occupied by the foot on the left side; correspondingly, 
the posterior shell muscle of the type species is merged with 
the right shell muscle in P. midatlantica. Other differences are 
that it has three rather than six pairs of epipodial tentacles. 
The new species also provides evidence of faunal interchange 
between widely separated ridge systems. 

Key words: .'\rchaeogastropoda, Fissurellacea, Clypeosectidae; 
hydrothermal- vent limpets; Mid-Atlantic Ridge. 



INTRODUCTION 

The slit limpet genus Pseudorimula McLean, 1989, was 
based on a single species from hydrothermal vents at the 
Marina Trough in the mid-Pacific. Here I add to the 
genus a second species from the Mid-Atlantic Ridge, a 
ridge system for which other components of the fauna 
are largely undescribed. First indications of biota on the 
Mid-Atlantic Ridge came from camera tows and dredg- 
ings by the NOAA vessel Researcher at a hydrothermal 
field at 26°N (Rona et a/., 1986); moilusks were not re- 
ported. Mollusks from the Mid-Atlantic Ridge were first 
collected in 1988 by observers on the deep-submersible 
Nautile at the Snake Pit hydrothermal field at 23°N. 
Spreading centers at these two sites on the Mid-Atlantic 
Ridge are diverging at a slower rate than those of the 
East Pacific (Rona et a/., 1986; Tunnicliffe, 1991). 

Unusual features of the biota of the Snake Pit vents 
were noted by Mevel et al. (1989): "The characteristic 
feature of these hydothermal sites is the amazing density 
of shrimps agglutinated on the chimneys; around the 
vents, the fauna consists of sea anemones, polychaetes. 



gastropods, galatheids, mussels and zoarcid fish. Tiie Snake 
Pit differs from the Pacific sites mostly by the absence 
of vestimentiferan worms, alvinellid and serpulid poly- 
chaetes and cephalopods. ' 

This new species of Pseudorimula came to my atten- 
tion after the original paper (McLean, 1989) was in press. 
It adds new limits to the morphology known in the family 
and provides an example of interchange between widely 
separated ridge systems. It is also the first mollusk to be 
documented from the Mid-Atlantic Ridge. 



MATERIALS AND METHODS 

Specimens were collected by the French expedition HY- 
DROSNAKE to the Mid-Atlantic Ridge, June-July 1988, 
and forwarded to me by Michel Segonzac of the Centre 
National de Tri d'Oceanographie Biologique (CENTOB, 
IFREMER, Brest). 

The illustrated radula was extracted from a preserved 
specimen after dissolution of tissues with room temper- 
ature 10% NaOH for 48 hours, washed in distilled water, 
air dried and coated with gold palladium for SEM ex- 
amination. 

Abbreviations for museums mentioned in the text are 
MNHN, Museum National d'Histoire Naturelle, Paris 
LACM, Los Angeles County Museum of Natural History 
LISNM, National Museum of Natural History, Washing- 
ton. 



Suborder VETIGASTROPODA Salvini-Plawen, 1980 
Superfamily FISSURELLACEA Fleming, 1922 
Family CLYPEOSECTIDAE McLean, 1989 

Clypeosectids differ from fissurellids in having a distinct 
radular plan, a reduced epipodium, a different pattern 
of shell musculature, and differences in the internal anat- 
omy, as discussed in more detail by McLean (1989) and 
Haszprunar (1989). Haszprunar (1989) provided the an- 



Page 116 



THE NAUTILUS, Vol. 106, No. 3 




Figures 1-7. PseudorUmda midatlantica McLean, sp. nov. NaiUile dive HSIO. Snake Pit hydrothermal field, Mid-Atlantic Ridge, 
3,478 ni. Hololype, MNHN. Anterior at top in vertical views. 1-3. External, internal, and left lateral views of shell. Length 8.1 
mm. 4. Dorsal view of detached body showing left and right shell muscles. 5. Ventral view of body attached to shell, showing 
protruding gonad on left side of foot. Epipodial tentacles are concealed by the foot in this view 6, 7. SEM views of radula of 
paratype. Scale bar for 6 = 20 ^m; scale bar for 7 = 10 fim. 



atomicai evidence to justify the erection of a second 
family in the superfamily Fissurellacea. 

On radular characters, the Plssurellidae differ from 
the Clypeosectidae in having a massive piuricuspid tooth 
tfiat separates the lateral field of teeth from the marginal 
teeth. The piuricuspid tooth is lacking in the C'lypeosec- 
tidae, in which the lateral and marginal teeth are strik- 
ingly similar in morphology. Clypeosectus McLean, 1989, 
was established for two species from eastern Pacific vents 
that have shells with an open slit deflected to the right. 
Pseudorimula. which is convergent in shell morphology 
with the fi.ssurellid genus Rimtda de France, 1827, has 
the slit closed at the margin. 

Genus Pseudorimula McLean, 1989 

Pseudorimula McLean, 1989:22. Type species; Pseudo- 
rimula marianae McLean, 1989. 



Pseudorimula midatlantica McLean, spec. nov. 
(figures 1-7) 

Pseudorimula sp. McLean; Martin & Hessler, 1990:10; 
Tunnidiffe, 1991:349. 

Description: Shell (figures 1-3) relatively large for fam- 
ily, maximum length 8. 1 mm. Surface coated with rusty 
mineral deposits, under which periostracum yellowish 
brown, tightly adhering, projecting slightly past shell 
margin. Outline of aperture oval, margin of aperture 
nearly in same plane; highest elevation of shell at about 
one-half its length. Profile moderately high, height of 
h()lot\pe 0.34 times length. Apical whorl at two-thirds 
shell length from anterior end, slightly deflected to right; 
protoconch diameter 150 nm. First teieoconch whorl 
smooth, rounded; slit arising two protoconch diameters 
away. Juvenile shells with open slit; in mature shells slit 
open about one-third length of anterior slope, strongly 



J. H. McLean, 1992 



Page 117 



deflected to right. Borders of foramen raised, except an- 
teriorly, where slit is sealed in mature shells and its trace 
slightly depressed. Selenizone weakly depressed below 
slit border, additions to selenizone extending straight 
across. Sculpture of about 30 well-defined primary ribs 
with one to three secondary ribs of lesser prominance 
arising in interspaces. Concentric sculpture of fine growth 
lines, raised into sharp lamellar scales on crossing primary 
ribs. Shell interior opaque. Muscle scar well marked on 
shell interior; left and right arms swollen at anterior tips, 
but of uneven thickness posteriorly; right arm longer and 
retaining its thickness to its termination near midpoint; 
left arm shorter anteriorly and posteriorly, connecting to 
right arm through narrow band. Anterior pallial attach- 
ment scars well marked, extending close to suture bring- 
ing two anterior portions into contact anterior to fora- 
men. Suture with zig-zag outline. Shell strengthened by 
thickened callus adjacent to suture and surrounding fo- 
ramen. 

Dimensions of holotype: Length 8.1, width 6.4, height 
2.5 mm. 

External anatomy (figures 4, 5): Anterior end of foot 
with double anterior edge marking opening of anterior 
pedal gland; foot posterior rounded; left side of foot 
displaced posterio-laterally by projecting gonad. Ce- 
phalic tentacles contracted from preservation. Two pos- 
terior pairs of epipodial tentacles, on body wall midway 
between foot edge and thick border of mantle margin 
(posterior tentacles of right side in cavity abutting pro- 
jecting gonad); single anterior pair, all with thick, joined 
bases, each with narrow projecting tips. Mantle skirt 
deeply emarginate, corresponding to foramen and seam 
in shell, edge of emargination with projecting papillae. 
Mantle skirt above head thin, nearly transparent. Shell 
muscles without inturned hooks; left muscle short in re- 
lation to right and truncate, right muscle extending more 
anteriorly than left, having thick posterior arm extending 
not quite to midpoint; left and right muscles joined by 
thin connective muscle near margin. Right ctenidium 
smaller than left; both ctenidia reduced, having filaments 
only on inner sides of axes. 

Radula (figures 6, 7): Radular ribbon nearly symmet- 
rical. Rachidian tooth with long overhanging tip, edges 
deeply serrate; shaft of rachidian short but broad at base. 
Four pairs of lateral teeth, similar in morphology to 
rachidian except much narrower; outer edges with fine 
serration, inner edges smooth; size of overhanging tips 
decreasing gradually away from rachidian. Marginal teeth 
numerous, with broad tips, edges finely and sharply ser- 
rate, serrations similar to those of laterals. Marginals and 
outer laterals with one long denticle on outer edge of 
shaft near overhanging tip. 

Type locality: Snake Pit hydrothermal field, Mid-At- 
lantic Ridge (23°22'N, 47°57'W), 3,478 m. 

Type material: Holotype and 20 paratypes from Natt- 
tile dive HSIO, 28 June 1988. Holotype MNHN; 16 para- 
types MNHN; 2 paratypes LACM 2424, 2 paratypes 



USNM 859485. The holotype is the only specimen in 
which both body and shell are in good condition, al- 
though the shell is heavily coated with mineral deposits. 
All other paratypes are smaller and completely decal- 
cified so that they can not be measured. Bodies are mostly 
separated from the shell remnants and are somewhat 
mangled, although it is evident that the gonad protrudes 
on the left side in each. The radula was prepared from 
a detached body. 



DISCUSSION 

Pseiidorimula midatlantica differs from P. marianae in 
having the right shell muscle longer than the left rather 
than having three separate muscles, a hypertrophied go- 
nad that displaces the foot on the left side, a single pair 
rather than two pairs of anterior epipodial tentacles and 
two rather than four pairs of posterior epipodial tentacles. 
The two species can hardly be distinguished in shell 
profile or sculpture, although the holotype shell of P. 
midatlantica has an opaque interior, in contrast to the 
transparent condition in P. marianae. This, however, 
may be due to differences in preservation, including the 
treatment conditions that led to the decalcification of all 
paratype specimens of P. midatlantica. 

In the original description of the genus Pseudorimula, 
I noted that the outer lateral teeth are morphologically 
similar to the inner marginal teeth, so much so that it is 
difficult to determine which are lateral teeth and which 
are marginal teeth. For P. midatlantica (figures 6, 7) I 
identify four pairs of lateral teeth by their more acute 
tips, compared to the broader terminations of the mar- 
ginal teeth. Earlier (McLean, 1989: figs. 13C,D) I stated 
that P. marianae has five pairs of lateral teeth, but now 
revise that to four pairs based on re-examination of the 
original illustration. 

Both of the two known species of Pseudorimula have 
differing features that set each apart from all other lim- 
pets: In P. mariana the shell muscle is inexplicably di- 
vided into three units, quite unlike the usual horseshoe- 
shaped muscle configuration in limpets of many families. 
In P. midatlantica the gonad is so hypertrophied that it 
displaces the foot on the left side; the shell muscle on 
the right side, where the foot remains large, is corre- 
spondingly longer. Which, then, of these two conditions 
is the more derived? We can assume that an ancestral 
species would have the usual horseshoe-shaped shell mus- 
cle and a gonad of normal size contained within the body 
cavity. Indeed, such a species may yet be discovered 
living on unexplored ridge systems. It is easy to under- 
stand the origin of the asymmetrical muscle pattern of 
P. midatlantica as an adjustment to gonad hypertro- 
phication (in response to need for greater reproductive 
output) that eliminates space for muscle attachment on 
the left side, but there is no easy explanation for the 
presence of three separate shell muscles in P. marianae. 
If, however, the hypertrophied gonad of P. midatlantica 
were to revert to a normal size, the large right muscle 
would already be in place and the posterior muscle could 



Page 118 



THE NAUTILUS, Vol. 106, No. 3 



then be pinched off from the posterior tip of the right 
muscle, once there was no longer the need for a large 
right muscle. 

The occurrence of the two species of Pseudorimula at 
such widely separate habitats as the Mariana Trough and 
the Mid-Atlantic Ridge is noteworthy but not unique. 
Martin and Hessler (1990) discussed "a growing body of 
evidence that there is a faunal connection between the 
Mariana vent area and the northern Mid- Atlantic Ridge." 
They mentioned similarities in the bythograeid crabs and 
gave three examples of genera represented in the two 
habitats, including Pseudorimula (as a personal com- 
munication from me), the shrimp genus Chorocaris 
(therein proposed), and a new genus of mussel (reported 
by Grassle, 1989). Martin and Hessler proposed that "the 
hydrothermal areas of the western Pacific and northern 
Mid-Atlantic Ridge were at one time connected via a 
series of active vent areas, not necessarily active simul- 
taneously, that extended from the Mid-Atlantic Ridge 
south to the Atlantic-Indian Ocean Ridge, north along 
the Southwest Indian Ocean Ridge, Mid-Indian Ocean 
Ridge, and Southeast Indian Ocean Ridge, and finally 
north through the various spreading centers of the Indo- 
West Pacific." 

Faunal connection between the Mariana Vents and 
those of the eastern Pacific were discussed by Hessler 
and Lonsdale ( 1991a, b). For vent limpets this applies 
only at the family level in the Clypeosectidae, as well as 
the Neomphalidae (McLean, 1990), but is exemplified 
at the species level by Lepetodrilns elevatus McLean, 
1988, reported by McLean (1990) to occur at the Mariana 
vents as well as the east Pacific vents. 

ACKNOWLEDGMENTS 

I thank Michel Segonzac (CENTOB, Brest), and Phi- 
lippe Bouchet of the Museum National d Histoire Na- 
turelie, Paris, for allowing me to describe the new species. 
(.1. (Clifton Coney operated the SEM to produce the rad- 
ular illustrations. I thank Anders Waren, Swedish Mu- 
semn of Natural History, and two anonymous reviewers 
for reading the manuscript. 



LITERATURE CITED 

Grassle, J. F. 1989. A plethora of une.xpected life. Oceanus 
31:41-46. 

Haszprunar, G. 1989. New slit-limpets (Scissurellacea and 
FLssurellacea) from hydrothermal vents. Part 2. Anatomy 
and relationships. Natural History Museum of Los Angeles 
County, no. 408, 17 p. 

Hessler, R. R. and P. F. Lonsdale. 1991a. Biogeography of 
Mariana Trough hydrothermal vent communities. Deep- 
Sea Research .38:185-199. 

Hessler, R. R. and P. F. Lonsdale. 1991b. The biogeography 
of the Mariana Trough hydrothermal vents. In: J. Mauch- 
ling and T Nemoto (eds. ), Marine biology, its accomplish- 
ment and future prospect. Kokusen-sha (Japan), p 165- 
182. 

Martin, J. W and R R Hessler. 1990. Chorocaris vandoverae. 
a new genus and species of hydrothermal vent shrimp 
(Crustacea, Decapoda, Bresiliidae) from the Western Pa- 
cific. Natural History Museum of Los Angeles County, 
Contributions in Science, no. 417, lip. 

McLean, J. H. 1989. New slit-limpets (Scissurellacea and Fis- 
surellacea) from hydrothermal vents Part 1. Systematic 
descriptions and comparisons based on shell and radular 
characters. Natural History Museum of Los Angeles Coun- 
ty, Contributions in Science, no. 407, 29 p. 

McLean, J. H. 1990. A new genus and species of neomphalid 
limpet from the Mariana vents with a review of current 
understanding of relationships among Neomphalacea and 
Peltospiracea. The Nautilus 104(3):77-86. 

Mevel, C, J. Auzende, M. Cannat, J. Donval, J. Dubois, Y. 
Fouquet, P. Gente, D. Grimaud, J. A. Karson, M. Segonzac, 
and M Stievenard. 1989. La ride du Snake Pit (dorsale 
medio-Atlantique, 23°22'N) resultats preliminaires de la 
compagne HYDROSNAKE. Comptes Rendus des Seances 
de 1 Academic des Sciences, Paris, series 2, 308:545-552. 

Rona, P. A., G. Klinkhammer, T. A. Nelsen, J. H. Trefry, and 
H. Elderfield. 1986. Black smokers, massive sulphides 
and vent biota at the Mid-Atlantic Ridge. Nature 321:33- 
.37, 

Tunnicliffe, V 1991. The biology of hydrothermal vents: 
ecology and evolution. Annual Review of Oceanography 
and Marine Biology, Margaret Barnes, ed., 29:319-407. 



THE NAUTILUS 106(3):119-122, 1992 



Page 119 



Prodissoconch I and II Length in Mercenaria Taxa^ 



Joy G. Goodsell 
Arnold G. Eversole 

Department of Aqiiaculture, 

Fisheries and Wildlife 
Clemson University 
Clemson, SC 29634-0362, I'SA 



ABSTRACT 

Mercenaria mercenaria larvae raised from 71) aiul SO ^m di- 
ameter eggs were reared for 4.S hours, until the prodissoconch 
I boundary was clearly visible. Prodissoconch I lengtlis of larvae 
from 80 ^m eggs were significantly greater than tho.se from 70 
^m eggs Larvae from crosses using similarly sized eggs of A/. 
campechiensis and M. campechiensis texana females and sperm 
from M. campechiensts. M. cairipechiensis texana and M. mer- 
cenaria males were reared through settlement There were no 
significant differences in prodissoconch I lengths; however, 
prodissoconch II lengths of larvae sired by A/, mercenaria were 
significantly longer than those sired by either subspecies of M. 
campechiensis. Differences in shell growth (AP = prodisso- 
conch II — prodissoconch I length) also followed taxonomic 
lines. 

Key words: Prodissoconch, egg, Mercenaria mercenaria. Mer- 
cenaria campechiensis. Mercenaria campechiensis texana. 



INTRODUCTION 

Relationships between egg size and shell length at the 
first shelled stage (i.e., prodissoconch I or protoconch I) 
have been reported for both bivalves and gastropods 
(Thorson, 1950; Robertson, 1971; Jablonski & Lutz, 1983; 
Lima & Lutz, 1990). Ockelmann (1962) suggested the 
usefulness of the length of prodissoconch I for estimating 
the egg size of bivalve species whose eggs were unavail- 
able because of extinction or the inaccessability of their 
habitat. Estimates of egg size could also be used to cat- 
egorize the developmental strategy of a bivalve as lethici- 
trophic ( 150-200 /zm diameter eggs), mi.xed (90-140 ^m) 
or planktotrophic (40-85 ^m) (Ockelmann, 1962). Small- 
er differences in the diameter of eggs (< 50 ^m) within 
Mercenaria taxa and other planktotrophic species have 
been observed (Bricelj & Malouf, 1980; Barber & Blake, 
1983; Heffernan et al.. 1989; Goodsell, 1991). Intraspe- 
cific differences in egg size have been attributed to the 
gametogenic state of the spawning female (Gallagher & 



' Technical contribution No. 3188 of South Carolina Exper- 
iment Station, Clemson University. 



Mann, 1986). Small differences in egg size (<10 fim) 
may be reflected in the prodissoconch I length of Mer- 
cenaria, consequently allowing inferences from larval 
assemblages about the condition of the female parent 
and the survival potential of its eggs. 

Developmental markers, such as prodissoconch I, re- 
main visible with continued growth (figure 1) and their 
size can be compared to later shell features (e.g., prodis- 
soconch II). In Mercenaria these shell markers delineate 
shell margin at the first shelled stage and at metamor- 
phosis to a benthic existence. Although molluscan larvae 
have been shown to delay metamorphosis until suitable 
environmental conditions are found (Bayne, 1965; Crisp, 
1974) no corresponding increase in shell length associated 
with the delay has been noted (Loosanoff, 1959). 

Pechenik and Lima (1980) demonstrated an inverse 
correlation between growth rate and length of larval 
period of the common Atlantic slippersnail, Crepidula 
jornicata (Linnaeus, 1758). They hypothesized that length 
of larval life in the slippersnail is limited by the rate of 
development toward a "predetermined endpoint' and 
that such an endpoint might also exist in bivalves. Early 
studies by Loosanoff (1959) demonstrated that when the 




Figure I. Scanning electron photomicrograph of a M. cam- 
pechiensis juvenile 753 Mm in shell length illustrating prodis- 
soconch I (P I) and prodissoconch II (P II) boundaries. 



Page 120 



THE NAUTILUS, Vol. 106, No. 3 



length of the larval period was increased by decreasing 
the culture temperature of M. mercenaria (Linnaeus, 
1758) larvae there was no corresponding increase in shell 
length, supporting Pechenik and Lima s "endpoint" ar- 
gument. The endpoint of larval bivalve life may be as- 
sociated with the loss of feeding ability that occurs with 
the degeneration of the velum and ciliary feeding tracts 
(Bayne, 1965). Differences in prodissoconch II length 
may prove useful in detecting the difference in larval 
endpoints among Mercenaria taxa. 

During the course of investigations into the genetics 
of Mercenaria we noted that eggs produced by any sin- 
gle, laboratory conditioned female during a given spawn- 
ing event were remarkably consistent in diameter. The 
availability of these consistently sized eggs from known 
parents allowed us (I) to examine the relationship be- 
tween egg diameter and prodissoconch I length in M. 
mercenaria, (2) to compare the prodissoconch I and II 
lengths of offspring from crosses involving M. campe- 
chiensis (Gmelin, 1791) and M. campechiensis texana 
(Dall, 1902) females with M. campechiensis, M. cam- 
pechiensis texatia and M. mercenaria males; and (3) to 
determine whether or not prodissoconch II lengths reflect 
taxonomic differences in parental stocks. 

MATERIALS AND METHODS 

Collection of adult Mercenaria and controlled spawning 
were achieved using methods previously described by 
Goodsell et al. (1992). Briefly, adults were collected from 
geographically isolated areas. M. mercenaria were col- 
lected from Milford, Connecticut; M. campechiensis from 
a site near Appaiachicola, Florida; and M. campechiensis 
texana from a location near Galveston, Texas. Adults 
were conditioned at 19 °C for six weeks prior to spawning 
and were induced to spawn individually using thermal 
shock and the addition of pasteurized sperm. Eggs were 
counted (40 x ) and 100 eggs from each spawning female 
were measured with an ocular micrometer at 100 x 
using a compound microscope. Eggs categorized as 70 
fim had a minimum diameter of 70 tim, maximum di- 
ameter of 75 ^m, mean <73 and standard deviation 
<2.0. Those categorized as 80 ixm had a minimum di- 
ameter of 80 nm, maximum diameter of 85 /xm, mean 
<83 and standard deviation <2.0. Sperm concentrations 
were estimated by spectrophotometry (Bricelj, 1979). To 
insure optimal gamete condition, all eggs were fertilized 
within one hour of spawning at the rate of 1,000 sperm 
ml' (Goodsell, 1991). 

Egg Size and Prodissoconch I Length in 

M. MERCENARIA 

Fertilized 70 and 80 ixm eggs of M. mercenaria were 
reared for 48 hours in 1 ^m filtered, UV treated seawater 
at a density <20 eggs ml '. A mixed algal diet (25,000 
cells ml"') was added to the culture at 24 hours. After 
48 hours larvae were viewed with light microscopy ( 100 x ) 
to ensure that the prodissoconch I boundary was clearly 



visible. Larvae were collected on a 44 /um mesh sieve 
and treated with sodium hypochlorite and distilled water 
to disarticulate the valves and remove soft tissues. Dis- 
articulated valves from each species were mounted, shell 
margin downward, on double stick tape attached to a 
glass slide. Prodissoconch I lengths (n = 30) from each 
egg size category were measured with an ocular mi- 
crometer on a compound microscope (lOOx). Analysis 
of variance by general linear model was used to examine 
the effect of egg diameter on prodissoconch I length (SAS 
Institute, 1985). 



Prodissoconch I and II Length of 
Mercenaria Taxa 

Larvae were reared for 15 days after prodissoconch I 
was achieved at a density of one larva ml ', to minimize 
the effects of crowding. Simultaneous spawning attempts 
resulted in crosses between female M. campechiensis (n 
= 3) and M. campechiensis texana (n = 2) and male M. 
campechiensis (n = 2), M. campechiensis texana (n = 
1 ) and M. mercenaria (n = 3). No M. mercenaria females 
spawned during the trials. Egg diameters and prodisso- 
conch I and II lengths were measured. Egg diameters (n 
= 100) from each spawning female were compared with 
an analysis of variance by general linear model to insure 
that egg sizes did not vary significantly between species. 
A measure of growth after the deposition of the prodisso- 
conch I boundary was calculated for each larva (AP = 
prodissoconch II length — prodissoconch I length). Pro- 
dissoconch I and II lengths and AP for each parental 
cross were compared with analysis of variance by general 
linear model and differences among crosses were sepa- 
rated using least square difference (SAS Institute, 1985). 



RESULTS 

Egg Diameter and Prodissoconch I Length in 

M. MERCENARIA 

Embryos derived from 80 um eggs of M. mercenaria 
(mean = 80.9 ^"11, SE ± 0.32 um) produce larvae with 
significantly greater (P < 0.0001 ) prodissoconch I lengths 
(109.7 ^m ± 0.7; range 102-119) than prodissoconch I 
lengths (101.9 jum ± 0.7; 88-109) of those larvae derived 
from 70 ^m eggs (70.8 ^m ± 0.15). 



Prodissoconch I and II Length of 
Mercenaria taxa 

Prodissoconch I lengths from crosses using eggs of M. 
campechiensis and M. campechiensis texana (egg di- 
ameter = 75.4 ixm ± 0.10) were not significantly differ- 
ent (table 1). However, significant differences (P < 0.05) 
in prodissoconch II length were found among the crosses. 
Metamorphosing larvae from crosses involving M. mer- 
cenaria males were significantK longer than from males 
of M. campechiensis texana and M. campechiensis. While 



J. G. Goodsell and A. G. Eversole, 1992 



Page 121 



not statistically significant, it is interesting to note that 
tlie offspring of female M. campechiensis were longer 
than offspring of female A/, campechiensis texana. No 
difference in prodissoconch II lengths of larvae from the 
male parents of the two M. campechiensis taxa was 
detected (Table 1). As expected, the growth (AP) differ- 
ences among crosses were similar to those differences 
observed with the prodissoconch II lengths Crosses in- 
volving male M. mercenaria demonstrated the greatest 
amount of shell growth. 

DISCUSSION 

Differences in prodissoconch I length was detected with 
differences in egg size as small as 10 nm in M. merce- 
naria, however, when similarly sized eggs were used no 
significant differences in prodissoconch I length were 
found among the closely related Mercenaria taxa. Prodis- 
soconch I lengths appear to vary consistently with egg 
size probably from some minimum egg size. Gallagher 
and Mann (1986) suggested that a minimum threshold 
lipid level (= egg size) was necessary to achieve normal 
embryogenesis. Knowledge of the relationship between 
egg size and prodissoconch I length might provide a 
technique for predicting the threshold size of eggs pro- 
duced by a female and the survival potential of its larvae. 

Loosanoff (1959) observed that the shell lengths of 
metamorphosing M. mercenaria and M. campechiensis 
larvae from mass spawnings were similar, suggesting that 
these taxa shared a similar larval endpoint (Pechenik & 
Lima, 1980). In contrast with Loosanoff's (1959) earlier 
study, we found differences in the mean prodissoconch 
II length (= shell length at metamorphosis) among off- 
spring sired by M. mercenaria and M. campechiensis. 
These differences in prodissoconch II length between the 
two species lend support to the placement of the texana 
subspecies within M. campechiensis. Examination of the 
distance between the prodissoconch I and II boundaries 
(AP) demonstrated that differences in prodissoconch II 
lengths were a function of growth differences among the 
taxa after the prodissoconch I boundary had been de- 
posited. The sampling technique in our study was dif- 
ferent from that of Loosanoff (1959). Loosanoff s (1959) 
results were based on mass spawnings with no attempt 
to limit the range of egg diameter. Furthermore, in Loo- 
sanoff's (1959) study, shell length at metamorphosis was 
determined by measuring the length of intact larvae as 
soon as they were observed to have lost their vela. In our 
study, shell length at metamorphosis was determined by 
measuring length at the prodissoconch Il/dissoconch 
boundary on disarticulated values. 

Prodissoconch I length does appear to be a good in- 
dicator of egg diameter. Also, prodissoconch II lengths 
(predetermined endpoint) follow taxonomic lines indi- 
cating genetic input as suggested by Pechenik and Lima 
(1980). Finally, prodissoconch I and II lengths should 
prove valuable in determining the condition of known 
parents and the relative genetic contributions of the par- 
ents in reciprocal crosses. 



Table 1 . Mean, standard error, range and results of least square 
difference test-" for prodissoconch I (P I) and II (P II) lengths 
(^m) and growth (AP), measured as prodissoconch II — pro- 
dissoconch I, by cross. Means not sharing the same letter are 
significantK different 



Prodissoconch I length 


























LSD 


(;:ross'' 


n 


P 1 


+ 


SE 


Range 


Gi 


rouping 


TC 


100 


104.1 


± 


0.34 


91-107 


A 




TT 


50 


103.9 


+ 


0.54 


99-112 


A 




CT 


7,5 


103.7 


+ 


0.47 


91-116 


A 




TM 


150 


102.4 


+ 


0..35 


91-116 


A 




c:m 


225 


101.8 


+ 


0.29 


91-116 


A 




cc 


1.50 


101.4 


-1- 


0.37 


91-116 


A 




Prodissoconch II 1 


ength 


























LSD 


Cross 


11 


P II 


+ 


SE 


Range 


Gi 


rouping 


CM 


225 


260 


± 


0.91 


224-290 


A 




TM 


150 


256.2 


± 


0.72 


22.3-273 


A 


B 


CT 


75 


246.0 


+ 


1.27 


21.5-282 




B C 


TT 


50 


243.1 


+ 


1.27 


224-265 




C 


CC 


150 


243.0 


+ 


0.87 


215-273 




C 


TC 


100 


240 1 


± 


99 


199-261 




C 


Growth 


(AP) 












LSD 


Cross 


n 


AP 


+ 


SE 


Range 


Grouping 


CM 


225 


158.3 


+ 


0.89 


124-191 


A 




TM 


1.50 


1.53.8 


+ 


0.80 


116-169 


A 


B 


CT 


75 


142.4 


± 


1 34 


112-182 




B C 


CC 


1.50 


141.6 


+ 


0.85 


107-174 




C 


TT 


50 


139.2 


+ 


1.24 


124-165 




C 


TC 


100 


1.36.0 


+ 


1.03 


91-162 




C 



^ Alpha = 0.05, df = 8. 

^' Abbreviations; M. campechiensis pure strain (CC), AT cam- 
pechiensis texana pure strain (TT), hybrid from M. campe- 
chiensis female x M. mercenaria male (CM), M. campechien- 
sis texana female x M. mercenaria male (TM), M. 
campechiensis female x M. campechiensis texana male (CT), 
reciprocal hybrid (TC) in descending order of size. 



ACKNOWLEDGMENTS 

The authors wish to thank William Arnold, Robert Bis- 
ker, Thomas Bright, Paul Chanley, James Moore and 
Randal Walker for their aid in the collection of the var- 
ious brood stocks. Staff support and culture facilities were 
provided by the Waddell Mariculture Center, Bluffton, 
South Carolina. Dr. Lawrence Grimes assisted in the 
statistical analyses. The research was supported b\ grants 
from South Carolina Sea Grant Consortium and South 
Carolina Agricultural Experiment Station. 

LITERATURE CITED 

Barber, B. J. and N. j. Blake. 1983. Growth and reproduction 
of the bay scallop Argopcclin irradians irradians (La- 



Page 122 



THE NAUTILUS, Vol. 106, No. 3 



marck) at its southern flislrilnitinnal limits Journal of Ex- 
perimental Marine Bioi(>g\ and Ecologs 66:247-256. 

Bayne, B. L. 1965. Growtii and tleiay of metamorphosis of 
the larvae of Mytilus edulis. Ophelia 2:1-47. 

Bricelj, V. M. 1979. Fecundity and related aspects of hard 
clam (Mercenaria mercenaria) reproduction in Great South 
Bay, New York. M.S. thesis. State L'niversity of New York 
at Stony Brook, 95 p. 

Bricelj, M. V. and R. E. Malouf. 1980. Aspects of reproduction 
in hard clams (Mercenaria mercenaria) in Great South 
Bay, New York. Proceedings of the National Shellfisheries 
Association 70:216-229. 

Crisp, D. J. 1974. Factors influencing the settlement of marine 
invertebrate larvae. In: P. T. Grant and A. M. Mackie 
(eds.). Chemoreception in marine organisms. Academic 
Press, New York, p. 177-265. 

Gallagher, S. M, and R. Maiui. 1986. Growth and survival of 
larvae of Mercenaria mercenaria (L.) and Crassostrea 
virginica (Gmelin) relative to broodstock conditioning and 
lipid content of eggs. Aquaculture 56:105-121. 

Goodsell, J. G. 1991. Egg size and shell morphological features 
of larval and early postlarval Mercenaria. M.S. thesis, 
Clemson University, 78 p. 

Goodsell, J. G , S. C. Fuller, A. G. Eversole, M. Castagna and 
R. A. Lutz. 1992. Larval and early postlarval shell mor- 
phology of several venerid clams. Journal of the Marine 
Biological Association of the Lhiited Kingdom 72:231-255, 

Heffernan, P. B., R. L. Walker and J. L. Carr. 1989. Ga- 
metogeneic cycles of three bivalves in Wassaw Sound, 



Georgia: 1 Mercenaria mercenaria (Linnaeus, 1758). Jour- 
nal of Shellfish Research 8(l):51-60. 

Jablonski, D and R A. Lutz. 198'3 Larval ecology of marine 
benthic invertebrates: paleobiological implications. Bio- 
logical Reviews 58:21-89. 

Lima, G. M. and R. A. Lutz. 1990. The relationship of larval 
shell morphology to mode of development in marine proso- 
branch gastropods. Journal of the Marine Biological As- 
sociation of the LInited Kingdom 70:611-637. 

Loosanoff, V. L. 1959. The size and shape of metamorphosing 
larvae of Veiius (Mercenaria) mercenaria grown at dif- 
ferent temperatures. Biological Bulletin 117:308-318. 

Ockelmann, W. K. 1962. Developmental types in marine 
bivalves and their distribution along the Atlantic coast of 
Europe. Proceedings of the First European Malacological 
Congress 1965:25-35. 

Pechenik, J. A. and G. M. Lima. 1980 Relationship between 
growth, differentiation, and length of larval life for indi- 
vidually reared larvae of the marine prosobranch gastro- 
pod, Crepidiila fornicata. Biological Bulletin 166:.537-549. 

Robertson, R. 1971. Scanning electron microscopy of plank- 
tonic larval marine gastropod shells. Veliger 14:1-12. 

SAS Institute Inc. 1985. SAS User's Guide: Statistics, Version 
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Thorson, G. 1950. Reproductive and larval ecolog\ of marine 
bottom invertebrates. Biological Review 25:1-45. 



THE NAUTILUS 106(3): 123-124, 1992 



Page 123 



Harpa cabriti Fischer, 1860, a replacement name for 
Harpa ventricosa Lamarck, 1816 (Gastropoda: Harpidae) 



Flarald A. Rehder 

Department (it Iinertebrate Zoology 
National Museum of Natural History 
Smithsonian Institution 
Wasiiingtoii, DC; 20560, USA 



ABSTRACT 

Harpa ventricosa Lamarck, 1816, is preoccupied In Harpa 
ventricosa Lamarck, 1801, a name that is here (letermined to 
he an objective junior synonym of Harpa major Hodiiig, 1798 
The name Harpa ventricosa Lamarck, 1816, used for a western 
Indian Ocean species, should be replaced by Harpa cabriti 
Fischer, 1860, a name based on an immature specimen formerly 
considered a junior synonym of Harpa ventricosa Lamarck, 
1816, non Lamarck, 1801. 

Key words: Harpidae, Harpa, nomenclature 



In 1986 Richard E. Petit and I submitted an application 
to the International (,'oininission of Zoological Nomen- 
clature (ICZN) proposing that the specific name Harpa 
articuiaris Lamarck, 1822, be conserved and placed on 
the Official List of Specific Names in Zoology (Rehder 
& Petit, 1987). 

When Miss Ruth A. Cooper of the ICZN was drafting 
the Opinion on this case she encountered a problem with 
our use of the name Harpa ventricosa Lamarck, 1816, 
noting that there was an earlier, validly published, Harpa 
ventricosa Lamarck, 1801 (Lamarck, 1801:79). That por- 
tion of the case involving Harpa ventricosa Lamarck, 




Figure 1. Holot>pe of Harpa cahnti Fischer, 1860. BM(NH) 1899.8.22.126. No locality data. Scale bar = 3 cm. 



Page 124 



THE NAUTILUS, Vol. 106, No. 3 



1816, and Harpa urnifortnis Perry. 1811, was therefore 
withdrawn from the petition, being considered a purely 
la.\onomic question, and Opinion 1518, conserving the 
specific name Harpa articularis Lamarck, 1822, was 
published in 1989. 

Through a now inexplicable oversight I had over- 
looked, in my monograph of the family Harpidae (Reh- 
der, 1973), the problem of homonymy involved in Harpa 
ventricosa Lamarck, 1801, versus Harpa ventricosa La- 
marck, 1816. The present paper was written to clarify 
this matter. 

In 1801 Lamarck published his "Systeme des Animaux 
sans Vertebres", wliich in essence is a treatise on the 
cla.ssification of the invertebrates, listing the genera of 
the various groups, each genus being briefly diagnosed, 
with one or more species cited as examples. The species 
are without diagnoses but have bibliographic references 
to previous descriptions or figures. 

His genus Harpa (Lamarck, 1801:79) is exemplified 
by one taxon, Harpa ventricosa Lamarck, a new name 
for the species illustrated by Lister (1770:pl. 992, fig. 55), 
and by Martini (1777:pl. 119, fig. 1090). Lister's figure 
represents Harpa harpa (Linne, 1758), as noted by La- 
marck (1822:256) and Dodge (1956:196-198). Martini's 
figure is representative of Harpa nmjor Roding, 1798, 
and was the figure on which Roding based his species. I 
therefore propose that the specimen illustrated in Mar- 
tini's figure serve as the type specimen of Harpa ven- 
tricosa Lamarck, 1801. I did this in the case of Roding's 
name (Rehder, 1973:247). This makes Lamarck's name 
an objective junior synonym of Harpa major Roding, 
1798. 

Harpa ventricosa Lamarck, 1816, isa junior homonym 
but not a junior synonym of Harpa ventricosa Lamarck, 
1801. Therefore, the species that had been called Harpa 
ventricosa Lamarck, 1816, by me (Rehder, 1973:251) 
and numerous other authors beginning with Lamarck 
(1822:255), is in need of a replacement name. 

I therefore propose that this species, which is restricted 
to the western part of the Indian Ocean and the Red 
Sea, bear the name of Harpa cabriti Fischer, 1860. The 
type specimen of H. cabriti, which I listed in the syn- 
onymy of Harpa ventricosa Lamarck, 1816 (Rehder, 
1973:252), is a juvenile or immature specimen, and is in 



the collections of The Natural History Museum, London. 
Through the kindness of Ms. Kath> Way I am able to 
illustrate this holotype [BM(NH) 1899.8.22.126] (fig.l). 

Harpa urniformis Perry, 1811, although listed in the 
synonymy of H. ventricosa Lamarck by Deshayes (1843), 
is not identifiable and is not considered to be a senior 
synonym of H. cabriti Fischer. 

LITERATURE CITED 

Deshayes, G. P. 1843. Histoire naturelle des Animaux sans 
Vertebres. Ed. 2. Volume 9. Paris. 725 pp. 

Dodge, H. 1956. A historical review of the mojlusk.s of Lin- 
naeus. Part 4. The genera Bticcinum and Stromlms of the 
class Gastropoda Bulletin of the American Museum of 
Natural History 1 1 1(3): 15:3-312. 

Fischer, P. 1860. Descriptions d'especes nouvelles. Journal de 
Conchyliologie 8(2):208-211, pi. 4, figs. 1-2, 6-8. 

International Commission on Zoological Nomenclature [ICZN] 
1989 Opinion 1518 Harpa articularis Lamarck. 1822 
(Mollusca: Clastropoda): specific name conservation. Bul- 
letin of Zoological Nomenclature 46(1):.53. 

Lamarck, j. B P. A. de M. de 1801. Systeme des Animaux 
sans Vertebres. Deterville, Paris, viii -I- 4.32 p. 

Lamarck, J. B. P A. de M. de 1816. [in 1791-1823]. Tableau 
Encyclopedique et Methodique des trois Regnes de la Na- 
ture. Vingt-Troisieme Partie. Mollusqueset Polypes Aivers. 
Liste des objets representes dans les planches de cette liv- 
rai.son. Panchouche, Paris. 16 pp. pls.391-488. 

Lamarck, J B. P. A de M. de 1822 [in 181.5-1822]. Histoire 
Naturelle des Animaux sans Vertebres. PrivateK pub- 
lished, Paris, vol. 7 [of 7], 711 p. 

Lister, M 1770. Historiae sive Synopsis Methodicae C^onchy- 
liorum. . . Editio Altera. Recensuit et indicibus auxit Gu- 
liehnus Huddesford. Clarendon Press, Oxford, iv + [96] 
pp. 439 pis. [1082 figs.]. 

Martini, F, H. W. 1777. [in 1769-1829]. NeuesSystematisches 
Conchvlien-Cabinet. vol 3 [If 12], vi + 434 p. pis. 66- 
121. 

Perry. G. 181 1. Conchology, or the Natural History of Shells; 
Containing a new arrangement of Genera and Species. 
W. Miller, London, 4 pp. 61 pis. 

Rehder, H. A. 1973. The family Harpidae of the world. Indo- 
Pacific Mollusca .3(16):207-274. 

Rehder, H. A. and R. E. Petit, 1987. Case 2548. Harpa ar- 
ticularis Lamarck, 1822 (Mollusca, Gastropoda): proposed 
conservation of the specific name. Bulletin of Zoological 
Nomenclature 44(l):19-20. 



THE NAUTILUS 106(3):125-126, 1992 



Page 125 



Obituary: 

Rae Baxter, 1929-91 



James H. McLean 

Los Angelt'S Oiunty Museum of 

Natural Histor\ 
900 Exposition Boulf\ard 
Los Angeles, CA 90007, USA 



Alaskan malacology suffered a great loss with the death 
of Rae Baxter, the most knowledgeable expert on Alaskan 
mollusks, on 22 Marcli 1991 at the age of 61. Rae was a 
Field Associate in Malacolog> of the Los Angeles County 
Museum of Natural History. He is survived by his wife 
Sera, daughter Lynx, and son Brant. 

Rae was born on 30 March 1929 in Compton, Cali- 
fornia. He earned a Bachelor of Science degree in fish- 
eries from Humboldt State University, California, in 1955. 
During his days at Humboldt, he became a student and 
collector of Pacific coast mollusks. While there he met 
Sera Miller, a native of Washington State, whom he mar- 
ried in 1955. Finding that northern climates were to their 
liking, Rae and Sera moved to Alaska after graduation. 

From 1956 to 1962, Rae worked for the US Fish and 
Wildlife Service out of Anchorage. Winters during this 
period were spent camping and trapping, during which 
Rae and Sera learned much about cold weather survival 
techniques. In 1960, the Baxters spent five months trav- 
eling in their sport boat from Kachemak Ba\ through 
southeastern Alaska and British C^olumbia to the Stilla- 
quamish River in Washington, camping on shore each 
night. Mollusks were collected at every opportunity and 
mailed back to Alaska. Sediment samples often were 
dried over the same stove used for cooking. 

In 1962 Rae joined the Alaska Department of Fish and 
Game, being stationed first at Cordova. During this pe- 
riod the Baxters built an A-frame dwelling for use as a 
base for commercial salmon fishing at Kasitsna Bay, on 
the southern shore of Kachemak Bay in Lower Cook 
Inlet. Sera was in charge of this venture, which continued 
each summer, with the help of their children. In 1966 
Rae was transferred to the Fish and Game laboratory at 
Bethel, on the Kuskwokwim River in western Alaska. 
The family lived at Bethel during the off season, return- 
ing to Kasitsna Bay each summer, until 1983, when Rae 
retired and returned to Kasitsna Bay year round. In 1989 
Rae finished building a laboratory for his new work as 
a marine biological consultant, and especially for re- 
search on his mollusk collection, for which he had as- 
sembled a good working library. 



Although Rae's work for the state of Alaska concerned 
the commercially important fishes and invertebrates, his 
compelling interest was the systematics and distribution 
of the molluscan fauna of Alaska. Over many years of 
field work throughout Alaska, Rae became familiar with 
the mollusks of the entire state, including the land and 
freshwater species as well as marine species. He es- 
pecialK knew the molluscan fauna of Kachemak Bay 
and would often set out alone in an open skiff for inter- 
tidal collecting and dredging. His spare time that was 
not devoted to collecting and studying mollusks was spent 
examining collections and gathering distributional rec- 
ords on visits to the Los Angeles County Museum of 
Natural History and the US National Museum of Natural 
History. After his retirement, he spent many summers 
aboard research vessels on long cruises in the Bering Sea, 
C^hukchi Sea, and Gulf of Alaska, working as a consultant 
to the National Marine Fisheries Service on Alaskan fish- 
ery assessment surveys. One of these cruises was aboard 
the Soviet ship Novokotovsk in the northwestern Bering 
Sea during the summer of 1990. Mollusks that would 
otherwise have been lost were retrieved by Rae on all 
the cruises. 

At the time of his death he was completing a detailed 
key to the fishes of Alaska for publication by the Amer- 
ican Fisheries Society. Upon completion of that project 
he was looking forward to being free to pursue his study 
of Alaskan mollusks, including the description of a num- 
ber of species that he had recognized as new. He had 
started to construct specimen cases so that his collection, 
until then labeled with station numbers and species code 
numbers, could finally be assembled as a working ref- 
erence collection. Rae's work on mollusks was not to be 
finished, but he has left a legacy both in his publications 
and his collection. 

In 1983 Rae published the first edition of his distri- 
butional checklist. The Mollusks of Alaska, a major proj- 
ect that was greatly facilitated b> the advent of personal 
computers, which enabled him to produce the finished 
text. Two subsequent editions (1985, 1987) were more 
detailed, treating a final total of 1,016 species. Two spe- 



Page 126 



THE NAUTILUS, Vol. 106, No. 3 






m 




1 


MM3k 


1 




H 




H 



Rae Baxter at Kasitsna Bay, Alaska, in August 197.3, screening mud dredged from his skiff (left) and picking salmon from a set net 
(right). 



cies were described or coauthored by Baxter: Macoma 
dexioptera in 1977 and Spiromoelleria kachemakensis 
in 1984. Mollusks named in his honor were Monadenia 
fidelis baxteriana Talmadge, 1954, Anatoma baxteri Mc- 
Lean, 1984, and Cocculina baxteri McLean, 1986. 

Rae was curious about all living things. He was a keen 
observer and was called upon for information by many 
biologists in Alaska. He often supplied live specimens for 
the aquaria at the Pratt Museum in Homer. He was 
always ready to assist the research of many workers around 
the world with specimens and information on Alaskan 
mollusks. Molluscan material from his field work in re- 
cent years was sent to the Los Angeles County Museum 
of Natural History, where it has greatly strengthened the 
Alaskan holdings. In accordance with his wishes, his en- 
tire mollusk collection has been transferred to the Los 
Angeles County Museum of Natural History, where it is 
to be integrated into the research collection and made 
available for study. 

I was privileged to stay with the Baxter family at 
Kasitsna Bay for a week in August of 1973, a wonderful 



experience both for the exposure to the rich molluscan 
fauna and the zeal with which Rae sought to understand 
it. Rae was a kind and generous man who is sorely missed. 

PUBLICATIONS BY RAE BAXTER 
ON MOLLUSKS 

Baxter, R. 1977. A new Alaska Macoma (Mollusca: Bivalvia). 
The Veliger 19(3);272-276. 

Baxter, R. 1983. Mollusks of .\laska, a listing of all mollusks, 
freshwater, land and marine, reported from the State of 
Alaska, with known locations of type specimens, maximum 
sizes, and marine depths inhabited. Privately published, 
xvii -I- 69 p 

Baxter, R. 1985. Mollusks of Alaska, 2nd ed. Privately pub- 
lished, 114 + xxxiii p. 

Baxter, R. 1987. Mollusks of Alaska, 3rd ed. Shells & Sea Life, 
Bayside, California. 163 p. 

Baxter, R. and J. H McLean. 1984. The genera Moelleria 
Jeffreys, 1865, and Spiromoelleria, gen. nov., in the North 
Pacific, with description of a new species of Spiromoelleria 
(Gastropoda: Turbinidae). The Veliger 27(2):219-226. 



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rHE NAUTILUS 



Volume 106, Number 4 
Felnuary 5, 1993 
ISSN 0028-1344 

A quarterly devoted 
to malacology. 



FEB 1 6 133: 




EDITOR-IN-CHIEF 
Dr. M. G. Harasewych 
Division of Mollusks 
National Museum of 
Natural History 
Smithsonian Institution 
Washington, DC 20560 



ASSOCIATE EDITOR 
Dr. R. Tucker Abbott 
American Malacologists, Inc. 
P.O. Box 2255 
Melbourne, FL 32902 



CONSULTING EDITORS 
Dr. Riidiger Bieler 
Department of Invertebrates 
Field Museum of 
Natural History 
Chicago, IL 60605 



Dr. Robert T. Dillon, Jr. 
Department of Biology 
College of Charleston 
Charleston, SC 29424 



Dr. William K. Emerson 

Department of Living Invertebrates 

The American Museum of Natural 

History 

New York, NY 10024 



Dr. Robert Hershler 
Division of Mollusks 
National Museum of 
Natural History 
Smithsonian Institution 
Washington, DC 20560 



Dr. Richard S. Houbrick 
Division of Mollusks 
National Museum of 
Natural History 
Smithsonian Institution 
Washington, DC 20560 



Mr. Richard I. Johnson 
Department of Mollusks 
Museum of (comparative Zoology 
Harvard University 
Cambridge, MA 02138 

Dr. Aurele La Rocque 
Department of Geology 
The Ohio State University 
Columbus, OH 43210 

Dr. James H. McLean 
Department of Malacology 
Los Angeles County Museum of 
Natural History 
900 E.xposition Boulevard 
Los Angeles, CA 90007 

Dr. Arthur S. Merrill 
% Department of Mollusks 
Museum of Comparative Zoology 
Harvard University 
Cambridge, MA 02138 

Ms. Paula M. Mikkelsen 
Harbor Branch Oceanographic 
Institution, Inc. 
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Dr. Donald R. Moore 

Division of Marine Geology 

and Geophysics 

Rosenstiel School of Marine and 

Atmospheric Science 

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Department of Geology 
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Museum of Zoology 
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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 Et7NAUTI LUS 



CONTENTS 



Volume 106, Number 4 

February 5, 1993 

ISSN 0028-1344 



Harald A. Rehder 



A new species of Harpa from the Leeward Islands of ^ ^ m 
Hawaii i ._ . . 127 

Pf'ro^roc'/u/,') niai/r^-ri, a new species of Pleurotomariid \'■^-^•^,l„ M'M' > ■■ ^ - 
from the western Atlantic (Gastropoda: Pleurotomariidae) 150 

The genus Fulgoraria (Gastropoda: Volutidae) of the 

northeastern Kamchatka Peninsula and Sakhalin Island, 

with notes on the paleoecology and distribution of the 

subfamily Fulgorariinae in the Oligocene of the northern 

Pacific ' 137 

A new species of Columbellid Gastropod from the Old 

World tropics 147 

The association between the gastropod Buccinanops 

cochlidium (Dillwyn, 1817) and the sea anemone 

Phlyctenanthus austratis Carlgren, 1949 in Patagonian 

shallow waters 152 

Patterns of diversity and extinction in Transmarian 

Muricacean, Buccinacean, and Conacean Gastropods 155 

Publication Dates of The Nautilus 174 



M. C. Harasewych 
Timothy M. Askew 



Anton E. Oleinik 



William K. Emerson 



Cuido Pastorino 



Edward J. Petuch 



Eugene V. Coan 
M. G. Harasewych 



Notice 



SMITHSONIAN FUNDS FOR MALACOLOGY 
STUDENTS 

The Division of Mollusks, Department of Invertebrate 
Zoology, National Museum of Natural History, Smith- 
sonian Institution announces the availability of the Rose- 
water Fellowship (up to $750) to be awarded to graduate 
students of systematic malacology. This award provides 
support for students conducting systematic studies of 
Mollusca (leading to publication) who require access to 
collections and libraries of the Division of Mollusks, Na- 
tional Museum of Natural History. Funds can be used 
for travel, subsistence, and research costs. Interested stu- 
dents should submit a succinct proposal (1-2 pages), in- 
cluding budget with indication of any matching funds, 
and a supporting letter from faculty advisor(s). Appli- 
cation deadline is March 15, 1993. Award(s) will be an- 
nounced on April 15, 1993. Applications should be sent 
to: 

Dr. M. G. Harasewych 
Division of Mollusks/NHB stop 118 
National Museum of Natural History 
Smithsonian Institution 
Washington, DC 20560 USA 



THE NAUTILUS 106(4):127-129, 1993 



Page 127 



A New Species of Harpa from the Leeward Islands of Hawaii 



Harald A. Rehder 

Department of Invertebrate Zoology 
National Museum of Natural Histor\ 
Smithsonian Institution 
Washington, DC 20560 U.S.A. 



ABSTRACT 

Harpa goodwini, new species, is described from two localities 
in the Leeward Islands of Hawaii, French Frigate Shoals and 
Maro Reef, where it occurs in 82-228 m. This new species is 
most closely related to Harpa cabritii Fischer, 1860, once known 
as H. ventricosa Lamarck, 1816 (not Lamarck, 1801). 

Key words: Gastropoda; Harpidae; Harpa, new species; Ha- 
waiian Islands. 



INTRODUCTION 

On several occasions Mr. Donald Dan has turned over 
to me for study specimens of Harpa received by him 
from correspondents. One of these lots represents a strik- 
ing new species from the Leeward Islands of the Hawaii 
Group. I am grateful to Mr. Dan for the privilege of 
examining this material, and for permitting me to de- 
scribe this new species. 

ABBREVIATIONS FOR INSTITUTIONS 

AMNH — American Museum of Natural History. New 

York 
AMS — Australian Museum, Sydney 
BM — Bernice P. Bishop Museum, Honolulu 
BM(NH) — British Museum (Natural History), London 
LACM — Los Angeles County Museum of Natural His- 
tory, Los Angeles 
MCZ — Museum of Comparative Zoology, Cambridge 
MNHN — Museum National d'Histoire Naturelle, Paris 
MHNG — Museum d'Histoire Naturelle, Geneva 
NSMT — National Science Museum, Tokyo 
USNM — National Museum of Natural History, Smith- 
sonian Institution, Washington, DC 

SYSTEMATICS 

Family Harpidae Bronn, 1849 
Genus Harpa Roding, 1798 
Harpa goodwini, new species 
Figures 1,2; Table 1. 



Description: Adult shell (figs. 1,2) 60-74 mm in length, 
broadly oval, outline of last whorl arcuate. Protoconch 
conical, pale pink, consisting of 2V8 to S'/s conve.x, glossy 
whorls. Postnuclear whorls 3%-4 in number, first l-l'/2 
whorls convex, pale pink in color, showing fine spiral 
cords at shoulder and below that cross fine axial riblets; 
this sculpture increasingly covered in subsequent whorls 
by the upper part of the glaze that covers the ventral 
wall at succeeding apertures and that extends up to the 
sharp spines of the axial ribs where they cross the shoulder 
cord. These spines become increasingly prominent and 
accentuate the flattened channel between suture and 
shoulder. On the upper whorls, the axial riblets in the 
channel are fine cords, but on the body whorl they be- 
come broader and flatter. Body whorl with 13-15 strong 
ribs that are erect with the moderately acute crest slanted 
away from the aperture along the apical half, but more 
rounded and flattened on the lower half of the rib. Ribs 
prominent and of orange color in the subsutural channel. 
At the shoulder, ribs form acute, triangular spines that 
are orange for most of the whorl, but become white near 
the aperture. The intercostal spaces are, as in most species 
of Harpa, axially finely striate. 

Basic color of fresh specimens is pinkish to pinkish 
orange, with the spire whorls appearing yellowish due 
to the glaze covering. Ribs on the body whorl typically 
crossed by a series of eight pairs of dark chestnut lines 
that are generally more pronounced on the last four or 
five ribs and situated as follows: two pairs between the 
spine at the upper end of each rib and Vt, of the distance 
down the length of each rib; another two pairs along the 
center of the w horl; below that a single line (occasionally 
absent), then another pair of lines, and at the base three 
pairs of lines (the last may be obscure or appear as a 
single line), making a total of 17 lines. Between these 
four groups of lines are moderateh broad bands com- 
posed of a central pale pink or orange pink band between 
two white bands. The intercostal spaces are marked with 
wavy, chestnut brown, axial lines, and, occasionally, by 
deep orange-rose splotches in two spiral bands, one in 
the center of the whorl, the other just below the spinose 
shoulder. Aperture oval with top of outer lip meeting 
the parietal wall at a right angle, the outer lip evenly 



Page 128 



THE NAUTILUS, Vol. 106, No. 4 




HA. Rehder, 1993 



Page 129 



Table 1. Ilarpa goaduini. new species. Measurements of shell characters. Linear measurements in mm 



Length 



Width 



No. ribs on 
1)0(I\ whorl 



No. whorls 



No. nuclear 
whorls 



Holoty pe 
Parat\pe 1 
Parat) pe 2 
Parat) pe 3 
Parat) pe 4 
ParatN pe 5 
Paratype 6 
Paratype 7 
Paratype 8 
Paratype 9 
Paratype 10 
Paratype 1 1 
Paratype 12 

Mean 

Range 

SD 



()3.S 
61.0 
73.4 
74.8 
70.2 
70.3 
69.2 
70.4 
70.1 
66.2 
61.4 
60 4 
60.0 

67.0 
60.0-74.8 

.50 



45.1 


42.7 


57.3 


55.2 


51.1 


52.5 


50.5 


48.1 


50.6 


44.7 


43.0 


43.7 


38.4 


47.9 


38.4-57.3 



\0 

13 
14 
14 
15 
15 
13 
15 
14 
15 
14 
15 
14 

14.3 
13-15 

0.7 



7.00 
7.00 
6.00 
6.16 
6.33 
6.25 
6.25 
6.33 
6.25 
6.12 
5.88 
6.00 
6.25 

6.29 

5.88-7.00 

.33 



3 33 


3.25 


2 + 


2.33 


2.50 


2.50 


2.50 


2.50 


2,25 


2.12 


2.12 


2.25 


2..50 


2.51 


2.12-3.33 


0.38 



arcuate. Parietal wall of aperture gently conve.x, colu- 
mella nearly straight, or slightly concave, anal fasciole 
strongly ridged by flattened ends of the ribs, the sinus 
fairly deep. Columella and parietal walls covered by a 
yellowish glaze, the outer lip yellow under the axial ribs, 
with deep chestnut hues on the ribs showing tiirough as 
chestnut spots. The ventral wail has three chestnut spots: 
a strong, elongate central spot that slants apically into 
the aperture above the top of the coiinnella; a weak, oval 
or roughly triangular spot between tlie ribs at the top of 
the parietal wall; and a small spot at the base of the 
columella. 

Type locality: French Frigate Shoals, Hawaiian Islands, 
about 166°10'E, 23°45'N, 137 m. Taken with hermit crabs. 

Material examined: Holotype, USNM 860312, Paratype 
1, USNM 860314, from the type locality; Paratype 2, 
RM; Paratype 3, MCZ; Paratype 4, Goodwin Collection; 
Paratype 5, AMNH 226438; Paratype 6, RM(NH); Para- 
tvpe 7, USNM 860315; Paratype 8, MHNG; Paratype 9, 
LACM; Paratype 10, AMS; Paratype 11, MNHN;'Para- 
type 12, NSMT; all collected between French Frigate 
Shoals and Maro Reef, Hawaiian Islands, in 82-228 m. 

Range: Leeward Islands, Hawaii, from French Frigate 
Shoals, to Maro (Dowsett) Reef. 

Habitat: I am aware of only 13 specimens of this species 
ha\ing been collected, all from lobster traps set in depths 
ranging from 82 m to 228 m. \\\ t\ pe specimens were 
brought into traps by hermit crabs. 



Etymology: This new species is named after Daniel R. 
Goodwin of Honolulu, Hawaii, who collected all of the 
type specimens. 

Comparative remarks: Harpa gooduini is a moderately 
large species characterized by its broad oval shape, its 
outline resembling that of Harpa costata (Linne, 1758) 
but with the outline of the last whorl more evenly ar- 
cuate, and differing by possessing an o\erall pinkish or 
orange-pink coloration when fresh, with dark horizontal 
stripes on the fewer, moderately broad and distant ribs. 
Harpa major Roding, 1798 differs from this new species 
in having a more oval shape, the whorls without the 
broad, flattened canal, and with the ribs fewer in number 
and lacking the dark horizontal lines. 

This new species most closely resembles H. cahritii 
Fischer, 1860 (see Rehder, 1973:251-252; 1992:123), and 
has the same general arrangement of dark chestnut 
blotches on its ventral side. Harpa gooduini has, how- 
ever, a relatively broader shell, w ith the body whorl not 
medially flattened but gently rounded. The axial ribs of 
H. cahritii lack the horizontal dark bands and numerous 
dark lines of H. goodwini. 

LITERATURE CITED 

Rehder, H. A. 1973. The family Harpidae of the world. Indo- 
Pacific Mollusca 3(16):207-274. 

Rehder, H. .\ 1992. A new name for Harpa lentricosa La- 
marck, 1816 (Gastropoda: Harpidae). The Nautilus 106(3); 
12.3-124. 



Figures 1. 2. Harpa guoduini, new species. 1. Holotype, USNM 860312. 2. Paratype 1, USNM 860313, both from French Frigate 
Shoals, Hawaiian Islands, about 166''10'E, 23°45'N, 137 m. Both specimens were brought into traps by hermit crabs. Scale bar = 
2.0 cm. 



THE NAUTILUS 106(4):130-136, 1993 



Page 130 



Perotrochus maureri, a New Species of Pleurotomariid from the 
western Atlantic (Gastropoda: Pleurotomariidae) 



M. G. Harasewych 

Department of Invertebrate Zoology 
National Museum of Natural History 
Smithsonian Institution 
Washington, DC: 20560 USA 



Timothy M. Askew 

Harbor Branch Oceanographic 

Institution, Inc. 

5600 U.S. I North 

Fort Pierce, FL 34946 USA 



ABSTRACT 

Perotrochus maureri. a new species of pleurotomariid, is de- 
scribed from the upper continental slope off northeastern Flor- 
ida and the Carolinas. It can be distinguished from its sister 
species, Perotrochus amabilis (Bayer, 1963), and from the more 
remotely related P. atlanticus (Rios & Mathews, 1968) by its 
smaller, lower, more deeply pigmented shell with fewer spiral 
cords on the selenizone and shell base, as well as on the basis 
of its radular asymmetry and formula. The emergence of pen- 
insular Florida is hypothesized to have separated the ancestral 
population into the Carolinian P. maureri and P. amabilis, 
which appears to be restricted to the Gulf of Mexico. 

Key words: Pleurotomariidae; Perotrochus. new species; bathy- 
al; vicariance; radula. 



INTRODUCTION 

During the course of continuing studies on the ecology 
and systematics of western Atlantic pleurotomariid gas- 
tropods, we had occasion to re-examine a large series of 
specimens that we collected off Charleston, South Car- 
olina and previously referred to Perotrochus amabilis 
(Bayer, 1963) (Askew, 1988:91; Harasewych et al. 1988). 
Comparison of this material, as well as additional spec- 
imens from off Jacksonville, Florida, with the holotype 
and additional specimens of P. amabilis from off north- 
ern Cuba and throughout the Culf of Me.xico revealed 
subtle but consistent differences in size, pigmentation and 
sculpture. We ascribe these differences to allopatric spe- 
ciation, and here describe as new the member of the 
species pair ranging from northern Florida to the Car- 
olinas. 



MATERIALS AND METHODS 

The type material was collected during four dives aboard 
the Research Submersible Nekton Delta, in the area of 
the "Charleston Lumps," a region of rugged topography 
some 90 miles east of Clharleslon, .South (Carolina, on May 
2-4, 1987. Some of the specimens were fixed in 10% sea 



water formalin and stored in 70% ethanol for dissection; 
the remainder were frozen and maintained at — 80°C. 

Additional specimens of the new species, as well as 
comparative material of P. amabilis and P. atlanticus in 
museum collections were examined, and are identified 
in the text by the following institutional acronyms; 

AMNH — American Museum of Natural Historv, New 

York 
AMS — Australian Museum, Sydney 
ANSP — Academy of Natural Sciences, Philadelphia 
BM(NH) — British Museum (Natural History), London 
FM — Fernbank Museum of Natural History, Atlanta 
FMNH — Field Museum of Natural History, Chicago 
FSBC I — Florida Marine Research Institute, St. Peters- 
burg 
HBOM — Harbor Branch Oceanographic Museum, Ft. 

Pierce, Florida 
LACM — Los Angeles County Museum of Natural His- 
tory 
MCZ — Museum of Comparative Zoology, Cambridge 
MNHN — Museum National d'Histoire Naturelle, Paris 
USNM — National Museum of National History, Smith- 
sonian Institution, Washington, DC 

SYSTEMATICS 

Perotrochus maureri new species. 

Synonymy: 

Perotrochus amabilis. .\skew, 1988:91, Harasewych et al. 1988, 
(Non Perotrochus amabilis (Bayer, 1963)). 

Description: Shell (fig. 1) medium-sized (to 52 mm), 
thin, trochoid, non-umbilicate, of up to 9% whorls. Spire 
coeloconoid, straight for first 5 whorls, becoming in- 
creasingly concave thereafter. Protoconch (figs. 2-3), 500 
^m wide, of 1 smooth, glossy, translucent whorl. Tran- 
sition to teleoconch abrupt, delimited by slightly flared 
protoconch lip, onset of spiral and axial sculpture, selen- 
izone. Selenizone initialK nearly abutting suture, de- 
scending to mid-whorl b\ whorl 2, below mid-whorl b\' 
whorl 6. Axial sculpture of pronounced prosoc> rt riblets 



M. G. Harasewych and T. M. Askew, 1993 



Page 131 



Table 1. Measurements of shell characters in Pcrotrochiis niaurcri. P amahilis and P allanliciis. Linear measurements in mm 





Pvwt 


rochiia maurcn 
(n = 10) 




P amahihs 

(n = 8) 




J 


f, allanliciis 

(n = 3) 




Character 


Mean 


Range 


SD 


Mean 


Range 


SD 


Mean 


Range 


SD 


Maximum dianu-tcr 
(MD) 


46.1 


34,4-59.5 


6,6 


72.2 


44.9-87.2 


12.1 


57.0 


56,3-58.2 


0.9 


Shell height 

(SH) 


37.3 


34.7-46.9 


44 


60.58 


41.7-73.3 


6.1 


53.7 


52.7-54.8 


0.9 


SH/MD 


0.822 


.791-832 


001 


0.858 


.840-929 


0.03 


940 


.936-. 951 


00 


No. whorls, 
teleoconch 


8.78 


8.25-9.75 


0,37 


9.76 


9.0-10.75 


0.54 


9,33 


9.0-9.75 


0.31 


No. spiral cords 
on selenizone 


1.80 


1-3 


0.75 


2.88 


2-4 


0.60 


3.33 


3-4 


0.47 


No. spiral cords 
on shell hast- 


UMI 


1 6-22 


1 6 


2S2 


24-31 


2 3 


27 


23-31 


3.3 



(18-20 on whorl 1 ), above and below selenizone, aligned 
in early whorls. Riblets decrease in prominence, being 
reduced to beads on spiral cords by whorl 6. Spiral sculp- 
ture initially of fine threads that cross riblets at angle 
(20-30°) to converge on selenizone from above and be- 
low. Single, continuous spiral cord first appears above 
the selenizone on whorl 2, below the selenizone on whorl 
4. Selenizone with opisthocyrt ribs that are more nu- 
merous than, and unaligned with, prosocyrt ribs. Spiral 
sculpture on selenizone of fine radial threads between 
adjacent ribs on early whorls, single medial spiral cord 
by whorl 4. Number of spiral cords above/on/ below the 
selenizone increasing to 3-4/1/2-3 on whorl 6, 6-8/1- 
3/3-4 on whorl 8. Suture initially grooved, becoming 
flat b\ whorl 5, impressed b\ whorl 7. Aperture hori- 
zontally ovately-rhomboidal. Outer lip smooth, portion 
below slit offset from portion above slit by 30-33° Slit 
narrow, extending posteriorly 84-92° from outer lip. Col- 
umella spiralK coiled, with strong sigmoid flexure near 
adapical margin. Umbilical region excavate but not per- 
forate. Base convex, with 17-22 even spiral cords be- 
tween periphery and umbilical region, which is nacreous 
due to resorption of outer layers of shell. Base color ivory, 
w ith broad axial bands of pale brick red, and narrower 
bands of dark red. Aperture nacreous, iridescent. Oper- 
culum (fig. 4) small, (spanning 0.6 of minor axis of ap- 
erture) multispiral, corneous. 

Anatomy: As the anatomy of P. maureri agrees in most 
regards with that of P. amahilis as described b\' Fretter 
(1964); only supplemental observations on living animals 
are recorded. The foot, head and tentacles are densely 
mottled with dark brick red. When the animal is crawl- 
ing, the posterior portion of the shell is supported on the 
operculum. Tentacles are long, cylindrical, ventrally di- 
rected. Left tentacle bilobed in one of the five specimens 
dissected. The jaws, inner lips and outer lips are all in- 
terconnected, being formed of a single piece of sclero- 
protein. The heavily papillated mantle edges on either 
side of the slit abut, sealing the slit except for a small 
opening along the posterior % of its length. The ctenidia 
do not project beyond the mantle edge in living speci- 



mens. When the animal is disturbed, the hypobranchial 
gland rapidly secretes large volumes of a whitish fluid 
that is immiscible with and denser than seawater. This 
secretion settles around and adheres to the shell. 

Radula: Radulae of 5 specimens (39-52 mm maximum 
diameter) were examined. Radula (figs. 5-10, table 2) 
long (75-82% maximum shell diameter), asymmetrical, 
left-skewed, bifid posteriorly, composed of 92-104 in- 
verted V-shaped rows of teeth. Hickman's (1984) ter- 
minology for the six tooth types is used herein and cor- 
related to other terminologies in table 2. Rachidian Tooth 
(figs. 6, 8) with dorsal surface laterally expanded, forming 
flanges that apparently serve to maintain alignment be- 
tween lateral teeth. Two long. Inner Lateral Teeth, with 
laterally expanded dorsal surfaces and broad, strongly 
curved distal ends (fig. 8, arrows) flank the rachidian 
tooth on each side. Adjacent are 24-26 (number increas- 
ing with shell size) Outer Lateral Teeth (figs. 6-8) that 
are shortest opposite Inner Lateral Teeth of adjacent row 
(fig. 7), and become progressively broader and stouter, 
with the long axis of the basal plates of the outermost 
Outer Lateral Teeth nearly perpendicular (*70°) to that 
of the innermost Outer Lateral Teeth and to the radular 
axis (fig. 8). These in turn are flanked by 21 large, curved 
Sickle Teeth (figs. 5-7). As in P. amahilis, the innermost 
teeth are tricuspid (fig. 7, arrow), but the cusp on the 
concave surface is lost in subsequent teeth (Fretter, 1964: 
181). The transition from Sickle Teeth to Filament-Tipped 
Teeth is gradual, the first Filament-Tipped Tooth dis- 
cerned b\' the presence of two minute bristles on either 
side of the proximal cusp (Woodward, 1901:250). The 
bristles increase in number and become larger, while the 
cusps diminish in size (fig. 9) and ultimateK- are lost in 
the outermost Filament-Tipped teeth. The outermost 8- 
9 teeth (fig. 10), referred to as Paddle-Shaped Teeth, are 
broad, flat, and blunt ended, the preceding 1-2 teeth are 
transitional from Filament-Tipped teeth to Paddle- 
Shaped Teeth and retain vestiges of a filaments along 
the inner distal ends. The Paddle-Shaped Teeth of one 
row overlap the outer Filament-Tipped Teeth of the 
adjacent, more proximal row, forming a telescoping mar- 



Page 132 



THE NAUTILUS, Vol. 106, No. 4 




Figure l.Perotrochus maureri. new species. Apertural. lateral, ventral and dorsal views of holotype (USNM 860320). Off Charleston, 
South Carolina, USA (32°43'57"W, 78°05'41"W) in 195-204 m. Scale bar = 1.0 cm. 



gin that envelops the Filament-Tipped Teeth and fa- 
cilitate,s smooth motion of the radula within its sheath 
on the rasping stroke. The blunt ends of the Paddle- 
Shaped Teeth may also dislodge entangled sponge tissue 
and spicules from the radula and free them to pass into 
the esophagus. 

Type locality: 90 nautical miles E of Charleston, South 
Carolina, USA (.32°43'54"N, 78°06'00"W, to 32°44'02"N, 
78°05'22"W), in 195-213 m. 



Range: Perotrochus maureri occurs on the upper con- 
tinental shelf off Charleston, South Carolina and Jack- 
sonville, Florida, in depths of 193-366 m. 

Type specimens: Holotype, USNM 860320; Paratypes 
1-9, USNM 875218; Paratype 10, AMS C. 169400; Para- 
type 11, BM(NH); Paratype 12, MNHN, Paratype 13, 
FMNH- Paratype 14, FSBC I; all from R/V Nekton 
Delta dive 561, (32°43'57"N, 78°05'4r'W) in 195-204 
m, 4 May 1987. Paratypes 15-17, LACM 2629; Paratype 



M. G. Harasewvch and T. M. Askew, 1993 



Page 133 



Table 2. (loniiiarison cif fornuilae and asviiiinetries iii plcunitiiniariiid radula Sources of data footnoted 
character could not be inferred from cited text or figure. 



indicates that trie 







A: 


Racl 
ian 


lid- 


Later; 


d teeth 




Marginal teetl: 


1 






15: 


Racl 


:,id- 




















ian 




Central 


Lamellate 


Hooked 


Brush 


Flahelliform 






C: 


Racl 


lid- 


Inner 


Outer 




Filament- 


Paddle- 




Taxon 


N 




ian 




laterals 


laterals 


Side 


tipped 


shaped 


Skew* 


Mikadotrochtis heijrichi^ 


■4? 








3 


20-21 


17-22 


63-65 


7-12 


R' 


Perot rochti.s iiuoiianiis- 


1? 








3 


24 


13 


63 


6 


R9.10 


P. aildcdoniciis^ 










1 


22 


14 


53 


7 


R" 


P notialis' 










2 


29 


13 


63 


10 


? 


P. amabilis'^ 










3 


24 


21 


63 


8 


R12 


P. maureri'^ 










2 


24-26 


21 


61-63 


8-9 


L« 


P midas' 










3 


26 


13 


63 


6 


L9.10 


P lucai/a' 










3 


25 


13 


61 


6 


? 


P. africana'* 










4 


25 


12 


50 


6 


? 



A = Standard terminology (e.g.. Fretter & Craham, 1962:169). B = Terminolog\ of Woodward, 1901, and other authors. C = 

Terminolog) of Hickman, 1984. 

* R = Right-skewed asymmetry. L = Left-skewed as\ mmetr> . 

' Tlie radular formula given by Woodward (1901:252) and subsec|uentl\ cited by other authors, is an incorrect summation of the 

data contained in his paper. His report that there are 223 teeth per row (p. 247), and explicit statement that there are 21 lamellate 

teeth (p. 249) indicate that the correct radular formula is (R-3-21-17-63-7), at least for the three specimens that he examined. 

Bouvier and Fischer (1902) report a radular formula of (R-3-20-22-65-12) for this species. 

- Bouvier and Fischer, 1899. 

' Bouchet and Metivier, 1982:310. 

■■ Leme and Penna, 1969:227. 

5 Fretter, 1964:181. 

^ Herein 

"Fretter, 1966:608. 

" Barnard, 1963:157. Listed as approximate counts. 

« Hickman, 1981:190. 

'" Hickman, 198430 

" Bouchet and Metivier, 1982:fig. 2. 

'- Herein, n = 2. 



18-20, USNM 875221; Paratype 21, MCZ, Paratvpe 22, 
ANSP, Paratvpe 23, AMNH, R/V Nekton Delta dive 
557 (32°43'54"N, 78°06'00"W) in 198-213 m, 2 May 
1987. Paratypes 24-26, USNM 846900 (reported in Hara- 
sewvch et al. 1988:94 as Perotrochus amabilis). R/V 
Nekton Delta Dive 560, (32°43'5S"N, 78°05'43"VV) in 
198-210 m, 3 May 1987. Paratypes 27-33, USNM 860321, 
Paratypes 34-35, FM 92.15. 1-2, R/V Nekton Delta 
Dive 562 (32°44'02"N, 78°05'22"W) in 204-213 m, 4 Mav 
1987. Paratype 36, HBOM 65:1988 Johnson-Sea-Link- 
1-1455, about 80 n miles off Charleston, South Carolina, 
USA 32°44.0'N, 78°05.6' W, in heavy rubble zone of rocks 
and sand, 193 m. September 6, 1983. 

Other material examined: HBOM 65:2017 Off Charles- 
ton, SC circa 1982; AMNH 226434, AMNH 226435, 
AMNH 226436, and USNM 869531, ail from Off Jack- 
sonville, Florida, in 366 m. 

Comparative material examined: Perotrochus amabilis 
(F.M. Baver, 1963): Holotype USNM 635625, S.E. of 
Sombrero Light, Florida ('24°29'N, 80°53'W-24°30'N, 
80°50'W), trawled in 220 m, R/V Gerda Cruise 6333, 
haul G-135; USNM 801707, 100 miles NNE of Sagua La 
Grande, Cuba (23°35'N, 79°34'W), in 18"3-238 m, R/V 



Silver Bay sta. 2460; USNM 846648, W of Tampa, Flor- 
ida, rubble bottom, 210 m; USNM 858215, Green Can- 
yon, 100 miles S of Atchafalava Bay, Louisiana 
(27°44'35"N, 91°07'54"W), in 170 m', Johnson-Sea-Link- 
1-2385, 17.6°C; AMNH 183151, WNW of Ft. Mvers, 
Florida, in 220 m; AMNH 244316, Dry Tortugas, Florida 
(no depth); FSBC I 30812, About 97' nautical miles W. 
of Mullet Key, Pinellas County. Florida (27°39'N, 
84°33'W), in 12'6.2-128.0 m. R/V Hernan Cortez; FSBC 
I 33146, About 95 nautical miles W. of Anna Maria 
Island, Manatee County, Florida (27°31'N,84°31'W) in 
135.6-126.5 m. R/V Hernan Cortez; HBOM 65:02190, 
About 100 nautical miles SE of Galveston, Texas, in 268.2 
m. Johnson-Sea-Link-II-933. Perotrochus atlanticus 
(Rios & Mathews, 1968): USNM 846647, Off Rio Grande, 
Rio Grande do Sul, Brazil, in 260 m; AMNH 181294, 
Off Rio Grande, Rio Grande do Sul, Brazil, in 164 m, 
sand & mud; AMNH 244317, Solidao, Brazil. 

Ecology: Specimens were observed in situ at the tvpe 
locality, an area of rugged terrain consisting of steep, 
large (3-30 m) hill crests and \alleNs. Perotrochus maur- 
eri was largely confined to hard substrates, composed of 
slabs or fragments of relithified phosphorite, that lined 



Page 134 



THE NAUTILUS, Vol. 106, No. 4 




Figures 2-4. Perotrochus maureri, new species. 2. Apical and 3. oblique views of protoconch. Scale bars = 250 ^ni. 4. Oviter and 
inner views of operculum. Scale bar = 1.0 mm. 



the tops and sides of the hill crests (see Askew, 1988:90, 
fig. 2). Water temperature was 9.6-9.8°C. Perotrochus 
maureri occurs in considerable densities at the type lo- 
cality, with distances between specimens ranging from 
10 to 30 meters. The diet of P. maureri [as P. amabilis] 
was reported by Harasewych et. al., (1988) to consist on 
sponges of the orders Poecilosclerida, Choristida or Spi- 
rophorida, and the genus Strongylophora . Most speci- 



mens have 7-12 repaired shell breaks, indicating fre- 
quent, unsuccessful predation, probably by crustaceans. 

Etymology: This species is named in honor of Mr. Rich- 
ard S. Maurer in recognition of his long and devoted 
interest in, and support of Malacology. 

Discussion: Perotrochus maureri is most closely related 
to P. amabilis, and more remotely related to P. atlan- 



Figures 5-10. Radiilar ribijon of Perolruchux muurcri. 7i. Dorsal view of critical-point dried buccal mass, showing alternating 
arrangement of booked teeth. Scale bar = .500 ^ni. 6. Anterior view of extended, critical-point dried radula Radula is seen to be 
asymmetrical and left-skewed. Scale bar = 500 ^m. 7. Left side of radula. Scale bar = 500 ^m 8. Rachidian, inner and outer 
lateral teeth Teeth to the left of the rachidian are further anterior than their homologues on the right. Scale bar = 250 ^m 9. 
Filament-tipped teeth from two adjacent rows. Scale bar = 100 ^m. 10. Paddle-shaped teeth. Transition from filament-tipped to 
paddle shaped teeth discernible on teeth 9-10 (arrows). Scale bar = 250 iim. 



M. G. Harasewvch and T. M. Askew, 1993 



Page 135 





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ki IfAA 


kI 


^WK^ 




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s/ ^S^ 


V^Jn^ 


^M 


^jm^i m^^. 


SL '/-' r 1 


■^^ M^^m 


A^H ^UBI^^^Sii.lr/^^iy > 


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I^J^ 


ssfl^ \vHv vlkr^ 


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sfe^^ /. JF^^/zcK 


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l^V iJv^^^Si 


V^^J 


^^mM //^^^ 


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Page 136 



THE NAUTILUS, Vol. 106, No. 4 




Figure 11. Map showing distribution of Perotrochtis amabilis 
(Bayer) (dots) and P. maureri n. sp. (open circles). Solid and 
open stars denote respective type localities. 

ticus. It can be distinguished from both these species by 
its smaller size, lower spire, and its more darkly pig- 
inented shell with fewer spiral cords on the selenizone 
and base (Table 1). The radula of P. maureri differs from 
that of P. amabilis in being left-skewed rather than right- 
skewed, and in having one fewer Inner lateral Teeth, an 
equal or greater number of Outer Lateral Teeth, an equal 
or lesser number of Filament-tipped Teeth and an equal 
or greater number of Paddle-shaped Teeth (Table 2). 
The utility of minor differences in radular formulae for 
distinguishing ta.\a is questionable, as most reports to date 
are based on a single radula per species. Intraspecific 
variation in the numbers of Outer Lateral, Sickle, Fila- 
ment-tipped and Paddle-shaped teeth has been encoun- 
tered when multiple radulae have been examined (Table 
2). As the transitions between the types of teeth are 
gradual (especialK' in the Marginal Teeth), it is also pos- 
sible that different investigators may have assigned one 
or more transitional teeth to different tooth types, thus 
further obscuring the discriminating value of radular 
formulae. Radular asymmetry, however, appears to be 
constant within a species (P. maureri, left-skewed, n = 
5; P. amabilis, right-skewed, n = 2), and is tentatively 
regarded as a u.seful distinguishing criterion. 

Perotroclms maureri is known only from the upper 
continental slope off northeastern Florida and the Car- 
olinas, where it occurs below the 10°C thermocline. Its 
sister species, P. amabilis, is limited to the Gulf of Mexico 
and the Straits of Florida (Fig. 11). A single temperature 
record from Louisiana (17.6°C) indicates that this species 
lives substantially above the 10°C thermocline. We sur- 
mise that the emergence of peninsular Florida separated 
tlie ancestral population during the Neogcne, with /'. 
amabilis evolving in the Gulf of Mexico, while P. maureri 
was isolated on limited areas of hard substrate off the 
Carolinas, and subsequently adapted to colder temper- 
atures. 



ACKNOWLEDGMENTS 

We thank Richard Cooper and Peter Auster of the NOAA 
National Undersea Research Program, University of 
Connecticut, Avery Point, for making submarine time 
available at the Charleston Lumps site. The assistance of 
Susann Braden with the scanning electron microscopy is 
gratefully acknowledged. This is Harbor Branch Ocean- 
ographic Contribution No. 943, and Smithsonian Marine 
Station at Link Port Contribution No. 315. 



LITERATURE CITED 

Askew, T. M 1988. A new species of pleiirotoniariid gastro- 
pod from the western Atlantic. The Nautilus 102(3):89- 
91. 

Barnard, K. H. 1963. Notes on the animals of Gyrinagiganfea 
(Lam.) and Pleurotomaria africana Tomlin. Proceedings 
of the Malacoiogical Societ\ of London .35(4): 155-158. 

Bayer, F. M 1963, .\ new pleurotomariid gastropod trawled 
in the Straits of Florida by R/\' Gerda, Bulletin of Marine 
Science of the Gulf and Caribbean 13(3):488-492. 

Bouchet, P. and B. Metivier. 1982. Living Pleurotomariidae 
(Moiiusca; Gastropoda) from the South Pacific. New Zea- 
land Journal of Zoology 9:309-318. 

Bouvier, E. L. & H. Fischer 1899 Etude monographique 
des Pleurotomaires actuels. Bulletin of the Museum of 
Comparative Zoolog\ , Harvard 32: 193-249, pis. 1-4; also 
published in Journal de (^oncln liologie 47(2):77-151, pis, 
4-7. 

Bouvier, E. L. and H. Fischer. 1902. L'organisation et les 
affinites des Gasteropodes primitifs d'apres I'Etude ana- 
tomique du Pleurotomaria Beyrichi. Journal de Conchy- 
liologie50(2):117-272, pls.2-6. 

Fretter, V 1964. Observations on the anatomy of Mikado- 
trochua amabilis Baser. Bulletin of Marine Science of the 
Gulf and Caribbean 14(1 );172-1S4, 

Fretter, V. 1966, Biological investigations of the deep sea, 16. 
Observations on the anatomv of Perutrochus. Bulletin of 
Marine Science 16(3):603-614. 

Fretter, V. and A. Graham. 1962. British Prosobranch Mol- 
lusks. Ra> Society, London. 755 p. 

Harasewych, M. G., S. A. Pomponi and T M. .\skew. 1988. 
Spongivory in pleurotomariid gastropods. The Nautilus 
102(3):92-98. 

Hickman, C, S. 1981. Evolution and function of asymmetry 
in the .\rcheogastropod Radula. The V'eliger 23(3):189- 
194, figs. 1-22. 

Hickman, C. S. 1984. Form and function of the radulae of 
pleurotomariid gastropods. The X'eliger 27(l):29-36. 

Leme, J. L. M. and L. Penna. 1969. Ocorrencia de Mika- 
dotrochus no Brasil. com descri^ao de uma nova especie 
(Gastropoda: Pleurotomariidae). Papeis Avulsos de Zool- 
ogia 22:225-230. 

Rios, E. de C. and H. R. Mathews. 1968. Nova especie de 
Pleurotomariidae do Brasil (Moiiusca: Gastropoda). -■Krq. 
Estafao de Biologia Marinha I'niversidade Federal do 
Ceara 8(l):65-68. 

Woodward, M. F. 1901. The -dnntomy oi Pleurotomaria beyr- 
ichii Hilg. The QuarterK Journal of Microscopical Science 
44(2):215-268, pis. 13-16, 



THE NAUTILUS 106(4):137-146, 1998 



Page 137 



The Genus Fulgoraria (Gastropoda: Volutidae) of the northeastern 
Kamchatka Peninsula and Sakhahn Island, with Notes on the 
Paleoecology and Distribution of the Subfamily Fulgorariinae in 
the Oligocene of the northern Pacific 



Anton E. Oleinik' 

Hussiaii Acadfiin nt Sciences 
Geological Institute 
Pyzhevsky per. 7 
109017 Moscow, Russia 



ABSTRACT 

Four new species of the genus Fulgoraria Sclnunacher, 1817, 
(subgenus Musashia Ha\ashi. I960) are described from the 
Oligocene Alugian Formation of the llpinsk\ Peninsula, north- 
eastern Kamchatka Peninsula, Russia. Previously described ful- 
gorariine gastropods from the same region, and from the Oli- 
gocene of Sakhalin and Karaginsky Islands, are also figured. 
Oxvgen isotopic analyses of contemporaneous Cyclocardia shells, 
along with a comparison of the ecology of Recent congeneric 
taxa and a paleoecologic anal) sis suggest a bath) al environment 
as most probable for these Oligocene Fulgorariinae. .\ review 
of the Oligocene biogeography of northern fuigorariines along 
the northern Pacific margin, including the western coast of 
North America, indicates that this subfamily had a much broad- 
er distribution during late Paleogene time than today. These 
data point to more favorable climatic conditions (including 
lower water temperatures) for dispersal of fulgorariine volutes 
during the Oligocene 

Key words: Fulgorariinae; systematics; distribution; Oligo- 
cene; Paleogene; Northern Pacific. 



INTRODUCTION 

Volutid gastropods are common as fossils in Cenozoic 
faunas of the northern Pacific region. However, their 
fossil record from some parts of this region, particularly 
the far east of Russia, is still very incompletely known. 
The subfamily Fulgorariinae Pilsbry and Oisson, 1954, 
is the dominant group among North Pacific Cenozoic 
volutes. Recent members of the "northern group" (Shi- 
kama, 1967) of this subfamily are restricted to Japan and 
adjacent seas. Less well-known extinct species occurred 
along both eastern and western margins of the North 
Pacific during both early and late Cenozoic time. 



' Present Address: Department of Earth and Atmospheric Sci- 
ences, Purdue University, West Lafayette, Indiana 47907, USA 



The subfamily Fulgorariinae comprises a group of car- 
nivorous gastropods with a uniserial radula composed of 
tricuspid rachidian teeth (Cooke, 1922; Habe, 1943; Oku- 
tani, 1963; Weaver & du Pont, 1970; Watanabe & Habe, 
1978). The higher systematics of this subfamily is still 
not fully resolved, with the two most recent revisions 
(Shikama, 1967; Weaver & du Pont, 1970) differing pri- 
marily in the ranking of supraspecific taxa. Shikama ( 1967) 
recognizes three genera: Fulgoraria Schumacher, 1817, 
Musashia Hayashi, 1960, and Saotomea Habe, 1943, as 
well as the subgenera Psephaea Crosse, 1871, Nippon- 
ornelon Shikama, 1967, Neopscphaea Takeda, 1953, and 
Miopleiona Dall, 1907. The last two are known only as 
fossils. Shikama's classification is based exclusively on 
shell characters such as the number and shape of colu- 
mellar plaits, the size and form of the protoconch, and 
features of the external shell morphology. All of these 
characters, especially the number of columellar plaits, 
may vary during ontogeny. Interpreting this highly vari- 
able shell morphology is further complicated when work- 
ing with fossil specimens, as they are often incompletely 
preserved. 

Based on shell and radular characters, Weaver and du 
Pont (1970) recognized only a single Recent north Pacific 
genus Fulgoraria, with the subgenera Psephaea, Volu- 
tipisrna Rehder, 1969, Musashia. Kurodina Rehder, 1969 
and Saotomea. These authors regard Nipponornelon as 
a synonsm of Musashia. and do not discuss the taxonomic 
position of the fossil Neopsephaea and Miopleiona. 

The present paper provisionally follows the classifi- 
cation of Weaver and du Pont (1970), but includes Nip- 
ponornelon. Miopleiona . and Neopsephaea as subgenera 
of Fulgoraria. 

Twenty-six species of Fulgorariinae from the latest 
Eocene and Oligocene formations of the north Pacific 
have been figured or described. Of these, eight are known 
only from North America [Poul Creek and Narrow Cape 
(of Sitkinak Island) Formations of Alaska; Blakeley, Twin 
River, and Eugene Formations of Oregon and Washing- 



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THE NAUTILUS, Vol. 106, No. 4 



ton], and have been referred to the subgenera Nippon- 
omelon, Musashia. Miopleiona. and Seopsephaea (Dur- 
ham, 1944; Addicott et ai, 1971; Tegland, 1933; Allison 
& Marincovich, 1981; Moore, 1984). Nine species, re- 
ferred to the subgenera Fulgoraria, Psephaea, Musashia, 
i^'ipponomelon. and Neopsephaea, are restricted to the 
Oligocene of Japan (Ashiya, Kishima. Nishisonogi, Ku- 
mano, and Chikokubo Formations of K% ushu and Hon- 
shu Islands; Momijiyama Formation of Hokkaido) (Oya- 
ma et ai, 1964; Shikama, 1967; Masuda & Noda, 1976). 
An additional seven species, from various localities along 
the northwestern coast of the United States and the Kam- 
chatka Peninsula, have only been identified to the sub- 
generic level (Allison & Marincovich, 1981; Moore, 1984; 
Gladenkov. Sinelnikova & Bratseva, 1987). Only one spe- 
cies endemic to the Kamchatka Peninsula and the Koryak 
Upland, Fitlgoraria {Musashia) olutorskiensis L. Krish- 
tofovich, 1973, has been described to date. A second 
Siberian species, Fulgoraria (Nipponomelon) tohunagai 
(Kanehara, 1937) is more widespread and is also known 
from the Oligocene of Japan. 

MATERIALS AND METHODS 

All specimens in this stud\- were collected during field 
work in eastern Kamchatka and Sakhalin Island between 
1965 and 1986. They were taken from the Alugian For- 
mation of northeastern Kamchatka, the "Laternula" 
sandstones of Karaginsky Island, and the Matchigarian 
Formation of central Sakhalin (fig. 21). The preservation 
of the specimens was often fragmentary, with the figured 
specimens representing the most complete material 

In order to address the questions of the ecolog\ of 
these Oligocene fulgorariines, a paleotemperature anal- 
ysis of shells of Cyclocardia ilpinensis Pronina, 1973, 
which occurs in abundance in the same strata as the 
volutes, was conducted. This standard analysis was based 
on the ratio of the oxygen isotopes ('''O/ '*0). The values 
of '''O ""O (PDB standard, mass spectrograph) were ad- 
justed to the Standard Mean Ocean Water (SMOW) stan- 
dard and corrected for altered isotopic composition at 
higher latitudes. Temperature values were calculated us- 
ing the following formula: 

T = 16.5 - 4.3 (b - A) + 0.14(b - A)^, 

where T is the temperature in °C, b is the instrumentally 
determined difference in the '^'O/ "'O ratio between the 
sample and the standard, and A is the correction for the 
original isotopic composition of sea water. For high lat- 
itudes A = — 1. Prior to analysis, all shells were examined 
using X-ray diffraction analysis to exclude specimens in 
which the aragonitic structure had recr\stallized. 



The following institutional acronyms are used; GI — 
Geological Institute, Russian Academy of Sciences. Mos- 
cow; CMG — Central Museum of Geology, St. Petersburg, 
Russia; USNM National Museum of Natural History, 
Smithsonian Institution, Washington, DC, USA. 

SYSTEMATICS 

Family Volutidae Rafinesque, 1815 
SubfamiK Fulgorariinae Pilsbry and Olsson, 1954 
Genus Fulgoraria Schumacher, 1817 
Subgenus Musashia Hayashi, 1960 

Fulgoraria {Musashia) novoilpinica new species 
Figures 1, 2, 17 

Description: Shell fusiform, slender, with 4-5 postnu- 
clear whorls. Last whorl stout, comprising -3 of shell 
height. Suture moderateK impressed, subsutural band 
absent, .\perture elliptical (Length width — 3). Inner 
lip with one narrow columellar fold and siphonal fold. 
Siphonal canal wide. Axial sculpture of numerous (10- 
12 on penultimate whorl) thin ribs, more pronounced on 
earlier whorls, smoother and wider on bod\ whorl. Spiral 
sculpture of fine, raised threads covering entire surface. 

Material examined: Holot\pe — USXM 468649. length 
(incomplete) 119.5 mm, width 58.3 mm; Paratype — GI 
4072, length (shell strongly decorticated) 124.7 mm. width 
62.2 mm; 1 ju\enile shell. 7 broken shells and fragments, 
2 molds; all from the t\ pe localit\ . 

Type locality: Northwestern part of the Ilpinsky Pen- 
insula, eastern Kamchatka, Russia. Upper part of the 
.Alugian Formation, Oligocene. 

Stratigraphic range: Known onl\ from the Upper part 
of the Alugian Formation of eastern Kamchatka. Oli- 
gocene. 

Comparalive remarks: .\lthough similar to Fulgoraria 
shutoi Shikama, 1967, from the Kishima Formation (Oli- 
gocene, Japan), this new species differs in having a more 
elongated last whorl, narrower axial folds, nearly smooth 
body whorl, and only a single fold rather than several 
strong ones. 

Fulgoraria {Musashia) olutorskiensis (L. Krishtofovich, 

1973) 

Figures 11, 12, 15, 16 

Miopleiona oregonensis Khomenko, 193325, pi 6, fig.20. 
Miopleiona olutorskiensis L. Krishtofovich, 1973:77, pi. 22, 

figs. 8,9, 
Musashia olutorskiensis Dewatiiova & X'oiobueva, 1981:128, 

pi. :32, fig. 2. 



Figures 1 . 2. Fulgoraria {Musashia) novoilpinica, new species. Holotype, I SNM 468649, 1 19.5 mm, .\lugian Formation, Oligocene. 
Figures 3. 4. Fulgoraria (Musashia) genuata, new species. Holotype, USNM 468650, 84.5 mm, .■\lugian Formation. Oligocene. 
Figures 5. 6. Fulgoraria {Seopsephaea) tenuis (Shikama, 1967). GI 1164, 49.2 mm, Matschigarian Formation, Oligocene. Figures 
7, 8. Fulgoraria (Musashia) tilitschikensis, new species. Holotype, USNM 468652, 51.1 mm, Alugian Formation, Oligocene. 



A. E. Oleinik, 1993 



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THE NAUTILUS, Vol. 106, No. 4 



Musashia (Musashia) sp. Gladenkov, Siiielniko\a & Bratseva, 
1987:57, pi. 15, fig.2. 

Description: Shell oblong, fusiform, relatively thin, with 
5-6 postnuclear whorls. Body whorl comprises appro.x- 
imately -3 of shell height. Suture slightU depressed, with 
narrow subsutural band. Aperture elliptical, acute pos- 
teriorly. Inner lip arched, with one narrow columellar 
fold. Callus thin, narrow. Shell surface nearly smooth, 
\\ ith thin growth striae and very fine, raised spiral threads. 

Material examined: Holotype— CMG 21/10285, length 
130.4 mm, width 48 mm, Pachatchi River, Olutorsky 
Region, eastern Kamchatka, Russia, "Hpinian" Forma- 
tion, Oligocene; Paratype— 19/10285, Gulf of Olutorsk, 
Govena Peninsula, eastern Kamchatka, Russia, Oligo- 
cene; GI 1486, Gl 4055 — both from Ilpinsky Peninsula, 
northeastern Kamchatka, Russia, Alugian Formation, 
Oligocene; 9 incomplete shells and fragments and 7 molds. 

Type locality: Olutorsky Region, northeastern Kam- 
chatka, Russia. 

Stratigraphic range: Known from the Oligocene for- 
mations of northeastern Kamchatka; abundant in the 
Alugian Formation of the Ilpinsk\- Peninsula. 

Comparative remarks: With its nearly smooth sculpture, 
this species most closely resembles Fulgoraria [Musashia) 
nagaoi (Shikama, 1967) from the Poronai Formation (Eo- 
cene-Oligocene of Hokkaido), but differs in being more 
elongated, in having fine spiral sculpture, and in having 
a distinct subsutural band. 

Fulgoraria (Musashia) genuata new species 
Figures 3, 4, 19 

Description: Shell oblong, fusiform, slender, w ith three 
stout, preserved whorls, and estimated 5-6 postnuclear 
whorls in intact specimens. Body whorl comprises < % 
total shell length. Suture shallow, slightly impressed. Ap- 
erture elliptical. Columella with one weak columellar 
fold. Sculpture of straight, smooth, widely spaced axial 
ribs (4 per whorl), most prominent on early whorls. Spiral 
sculpture of numerous, fine, slightly raised threads cov- 
ering shell surface. 

Material examined: Holotype — USNM 468650, length 
84.5 mm, width 40.0 mm; Paratype— GI 40721, length 
75.9 mm, width 46.1 mm; both from type locality. 

Type locality: Northwestern part of Ilpinsky Peninsula, 
northeastern Kamchatka, Russia. Alugian Formation. 
Oligocene. 

Stratigraphic range: Known only from the Alugian For- 
mation of the Ilpinsky Peninsula. Oligocene. 



Comparative remarks: Although similar to Fulgoraria 
(Musashia) fujimotoi (Kanno, 1958) from the Hikokubo 
Formation (Oligocene-Miocene, Japan), this new species 
differs in having stouter whorls and a lower spire, in 
having one instead of two columellar folds, and in having 
finer spiral sculpture. 

Fulgoraria (Musashia) cordafa new species 
Figures 13, 14 

Description: Shell elongate, fusiform, slender, with four 
postnuclear whorls, short siphonal canal and narrowly 
channeled suture. Body whorl comprises approximately 
¥4 of shell length. Aperture oblong, with narrow posterior 
angle. Columella with one weak columellar fold, .\xial 
sculpture of low, rounded ribs (— 12 per whorl on pen- 
ultimate whorl). Axial ribs crossed by fine, closely spaced 
spiral lines. 

Material examined: Holotype — USNM 468651, length 
59.1 mm, width 24.3 mm; Paratype 1— GI 4063/2, length 
50.5 mm, width 26.5 mm; Paratype 2 — GI 40531, length 
62.8 mm, width 26.5 mm; two molds, three fragments 
and one impression; all from the type locality. 

Type locality: Northwestern part of Ilpinskv Peninsula, 
northeastern Kamchatka, Russia. Upper part of the .alu- 
gian Formation. Oligocene. 

Stratigraphic range: Known only from the type locality. 

Comparative remarks: This new species is most similar 
to Fulgoraria (Musashia) weaveri (Tegland, 1933) of the 
Blakeley Formation (Oligocene, Washington, USA), but 
has a more elongated shell, more numerous axial ribs, 
and much stronger spiral threads. 



Fulgoraria (Musashia) tilitschikensis new species 

Figures 7, 8, 18 

Description: Shell globoseK fusiform, solid Teleoconch 
of about 5 whorls. Suture narrowly pressed. Body whorl 
comprises % shell length. Aperture semiovate, with thin, 
narrow callus. Inner lip w ith one weak, subvertical col- 
umellar fold. Axial sculpture of rounded arcuate ribs (13 
on body whorl). Spiral sculpture lacking. 

Material examined: Holot\pe — USNM 468652, length 
(incomplete) 51.1 mm, width 29.5 mm, from the type 
locality; Paratype GI 40724, length (incomplete) 65.0 
mm, width 37.6 mm, Ilpinsk> Peninsula, northeastern 
Kamchatka, Russia. .Alugian Formation, Oligocene. 

Type locality: Korf Settlement. Coal River, northeastern 
Kamchatka, Russia. Alugian Formation. Oligocene. 



Figures 9, 10. Fulgoraria (Nipponumclon) cLtokunagai (Kanehara, UW7). GI 3586 1804. 149 mm, " Lalcrnula" Sandstones, 
Karagin.sky Island, Oiigocene-Lower Miocene (?). Figures 1 1, 12, 15, 16. Fulgoraria (Musashia) olutorskiensis (L. Krishtofovich, 
1973). 1 1,12. GI 1486, 61.8 mm, 15, 16. GI 4055, 103.9 mm, Alugian Formation, Oligocene. Figures 13, 14. Fulgoraria {Musashia) 
cordata, new species. Holotype, USNM 468651, 59.1 mm, Alugian Formation, Oligocene. 



A. E. Oleinik. 1993 



Page 141 




cm. 



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THE NAUTILUS, Vol. 106, No. 4 






A. E. Oleinik, 1993 



Page 143 



Recent species of Fulgorarinae: 



DEPTH IN METERS 
250 500 



750 



Fulgoraria (Miisashia) cancellata Kuroda et Habe 
Fulgoraha (Musashia) formosana Azuma 
Fulgoraria (Musashia) clara (Sowerby) 
Fulgoraria (Nipponomelon) kamakurerisis (Otuka) 
Fulgoraria (Nipponomelon) elongata Shikanm 
Fulgoraria (Nipponomelon) prevostiana Crosse 
Fulgoraria (Nipponomelon) angulosa Shikama 




Figure 20. Bath>Tnetric distributions of Recent species of Fulgorariinae of Japan and adjacent seas, based on data in Shikama 
(1967), and Weaver and du Pont (1970). Thickness of hnes indicates relative abundance. 



Slratigraphic range: Known only from the Alugian For- 
mation of northeastern Kamchatka, Russia. Oligocene. 

Comparative remarks: This species resembles Fulgoraria 
(Musashia) shikamai Moore, 1984, from the Poul Creek 
Formation (Alaska, USA) and the Clallam and Twin Riv- 
er Formations (Washington, USA), but this new species 
has more sharpened axial ribs and finer spiral threads. 

Subgenus Nipponomelon Shikama, 1967 

Fulgoraria (Nipponomelon) of. tokiinagai (Kanehara, 

1937) 

Figures 9, 10 

Psephaea tokunagai Kanehara, 1937:16-18, pl.2, figs. 2-,5. 
Fulgoraria (Psephaea) prevostiana Crosse, Shikama, 1954:pi.6, 

fig. 26, 
Fulgoraria tokunagai Kamada, 1962:192, pl21, figs. 5-8. 
Musashia (Nipponomelon) tokunagai Shikama, 1967:100-101, 

pi. 14, fig. 4. 
Musashia tokunagai Devvatilova & Volobueva, 1981:128, pi. 33, 

fig- 1- 
Original Description: "Shell tall and high spired, apical 
angle about 25 degrees. Suture slightK impressed with 
narrow shelf around it. Surface ornamented with a.xial 
folds, which ma\ be somewhat accentuated above and 
tend to disappear on the body whorl. Spirally grooved 
regularly, also over the folds, .Aperture spindle shaped, 
obliquely notched posteriorly, with narrower anterior 
canal. Inner lip provided with two columellar folds and 



covered b\- a thin callus, which adheres to the columella 
and thickens anteriorly," 

Material examined: One specimen, GI 3586/1804, length 
149 mm, width 62.2 mm, from the "Laternula" sand- 
stones, Karaginsky Island, Gnunvayam River, Russia. 
Oligocene-Lower Miocene (?). 

Type locality: Yanagaya bed, \agakura coal mine, Yu- 
moto, Iwaki City, Fukushima Prefecture, Honshu, Japan. 
Lower Miocene. 

Stratigraphic range: "Laternula" sandstones, Karagin- 
sk\ Island, Gnunvayam River, Russia, Oligocene-Lower 
Miocene (?), to Yanagaya bed, Nagakura coal mine, Yu- 
moto, Iwaki Cit\ , Fukushima Prefecture, Honshu, Japan. 
Lower Miocene. 

Comparative remarks: Fulgoraria tokunagai is charac- 
terized by a large size, high spire, large protoconch, and 
numerous axial folds. The spiral sculpture on the single 
specimen from Karaginsky Island is poorly preserved, 
but this specimen retains visible traces of the axial folds 
and thin spiral grooves. The shape and size of this spec- 
imen, especialK the features of the aperture, suture, and 
sculpture, are the same as in typical F. tokunagai. How- 
ever, the Russian specimen differs from the type in hav- 
ing a single columellar fold rather than two. This dif- 
ference, together u ith the poor preserv ation of this shell 
puts the species determination of this specimen in some 
doubt. 



Figure 17. Fulgoraria (Musashia) novoilpinica, new species. Reconstructed drawing of holotype. Figure 18. Fulgoraria (Musashia) 
tilitschikensis. new species. Reconstructed drawing of holotype. Figure 19. Fulgoraria (Musashia) genuata. new species. Recon- 
structed drawing of holotype. Scale bars =10 mm. 



Page 144 



THE NAUTILUS, Vol. 106, No. 4 




Figure 21. Fossil localities of Oligocene Fulgorarinae in the North Pacific referred to in the text. 1 — Honshu Island, 2 — Hokkaido 
Island, 3 — Sakhalin Island, -4 — Kamchatka Peninsula, 5 — Karaginsky Island, 6 — Sitkinak Island. 7 — Kodiak Island, 8 — Oregon, 9 — 
Washington, 10 — California. 



Subgenus Neopsephaea Takeda, 1953 

Fulgoraria {Neopsephaea) tenuis Shikama, 1967 
Figures 5,6 

Musashia (Neopsephaea) /emiis Shikama, 1967:116-117, pi. 13, 
figs. 3,4. 

Original description: "Shell small in size, slender, fusi- 
form and with low spire. Suture shallow and there is no 
subsutural band. Last whorl vaulted at middle and ap- 
erture relatively narrow. Surface of whorl smooth and 
a.xial ribs relatively few and weak, about 11 in penulti- 



mate whorl, gradually becoming obsolete anteriorly. Tip 
of columella straight and narrow. There may be one 
columellar plait". 

Supplemental description: Shell is rather slender, elon- 
gate, fusiform, with about 5 (when restored) whorls sep- 
arated b\ a shallowK impressed suture. Bod> whorl and 
aperture comprising slightl) more than % of total shell 
length. Aperture elliptical, forming acute angles at an- 
terior and posterior margins. Siphonal notch relatively 
narrow. Outer lip thin, simple. Columella with one weak 
columellar fold. Callus verv thin. Shell surface smooth, 




Japan Sakhalin Kamchatka Alaska U.S. 

E 130 140 150 160 170 180 170 160 150 140 130 120 W 




Fulgoraria 

Psephaea 

Musashia 

[Nipponomelon 

Neopsephaea 

Miopleiona 



Cumulative 

Number 
of Species 



Figure 22. Geographic distribution of the subgenera ot Fulgoraria. and (heir species diversity during ihc Oligocene. 



A. E. Oleinik, 1993 



Page 145 



with thin growth Unes. Axial sculpture ot low , rounded 
ribs (11-12 on body whorl) that disappear on anterior 
portion of body whorl. 

Material examined: One specimen, Gl 1 164, length 49.2 
nmi, width 22 mm, near Lake Matschigar, central Sa- 
khalin Island, Russia. Matschigarian Formation. Oligo- 
cene. 

Type locality: Zochuku Sandstone, Kishima Group, Na- 
gasaki Prefecture, K) ushu, Japan. Oligocene. 

Stratigraphic range: Near Lake Matschigar, central Sa- 
khalin Island, Russia, Matschigarian Formation, Oligo- 
cene, to Zochuku Sandstone, Kishima Group, Nagasaki 
Prefecture, K\ushu, Japan, Oligocene. 

Coniparaiive remarks: This species is closeK allied to 
Fulgoraria (Mcopscphaea) antiquior Takeda, but may 
be distinguished b\ its more swollen body whorl, nar- 
rower aperture w ith distinctive posterior margin, smooth 
surface, and a.xial ribs that are fewer in number or en- 
tirely obsolete. 



DISCUSSION 

The lithology of the Alugian Formation of the llpinsky 
Peninsula in eastern Kamchatka consists of gray to dark 
gray claystones and clays. This formation has a total 
thickness of about 950 meters, and is continuous, lacking 
hiatuses or internal unconformities. No shallowing or 
deepening trends are e\ident within the section. The 
strata contain numerous carbonate concretions ranging 
in size from 1 cm to several meters. Some of the claystone 
beds are delicately laminated. These types of sediments 
are indicative of soft bottom environments. 

Molluscan remains are quite common, but are not 
concentrated in particular layers or lenses. Fossils occur 
sporadically throughout the section, both in concretions 
and within the matrix. The molluscan fauna does not 
exhibit mans- changes from the bottom to the top of the 
formation. These data point to a depositional regime 
comprising relatively rapid and uninterrupted sedimen- 
tation (Kidwell, 1988). 

Species diversit\ in the molluscan assemblage of the 
Alugian Formation is low. The most abundant taxa oc- 
curring together with fulgorariine volutes include Trotn- 
inina bicordata (Hatai & Koike, 1957), Trominina ish- 
ihariensis (Ha\asaka & Matsui, 1951), Neptiinea ezoana 
Takeda, 1953, Bathybembix sakhalinensis korjakensis 
Volobueva, 1981, Optotiirris (?)sp., Turritellasp., Cryp- 
tonatica spp., Acila praedivaricata Nagao & Huzioka, 
1941, and Cyclocardia ilpinensis Pronina, 1973. All these 
taxa have relativeK broad bath\ metric ranges, occurring 
from lower neritic to bath)al depths (Hall, 1960; Noda, 
1975; Scarlato, 1981; Moore, 1984). 

Recent species of Fulgorariinae are mainly bathyal 
animals, dredged most often from depths of 250 to 700 
or more meters (fig. 20). Bath\rnetricalK , the bath\al 
zone extends from a mean depth of 200 m to 2000 m, 
and thermally, from the 15°C isotherm in low latitudes 



down to the — 3°C isotherm in high latitudes. Substrates 
are predominantly fine silts, muds, and oozes (Encyclo- 
pedia of Oceanography, 1974; Hickman, 1974). Oxygen 
isotope anaKses of five Cyclocardia ilpinensis shells col- 
lected from the bottom through the top of the Alugian 
Formation allowed the estimation of paleotemperatures 
as ranging from 3°C to 5°C. These data indicate that 
water temperatures remained quite uniform during the 
deposition ot this formation, and support the hypothesis 
of a bathyal environment. When compared to Recent 
bathyal bottom temperatures, these data suggest that the 
climatic conditions during Alugian deposition were sim- 
ilar to those found between 32°N and 45°N, and ap- 
proximate the temperatures that define the distribution 
of Recent species of Fulgoraria. Paleotemperatures es- 
timated for the Alugian Formation exhibit some corre- 
lations with those estimated for Oligocene formations of 
the western coast of North .America. Paleotemperatures 
of the Blakeley Formation (Echinophoria apta zone), 
which contains five species of Fulgorariinae. were esti- 
mated as to be 5-8°C. The remains of Aturia nautiloids 
in great abundance, which may have required a tem- 
perature of at least 16°C could be explained by post- 
mortem transport of empty shells, which is fairly com- 
mon with Recent NautUus. Paleodepth was thought to 
range from 100 m to 350 m (Moore, 1984). Similarly, 
the Narrow Cape Formation of Sitkinak Island (Alaska), 
which contains three species of Fulgorariinae , was de- 
posited in the outer neritic zone of the continental shelf 
(Allison, 1978). A comparison of the Narrow Cape data 
with that of Recent Alaskan mollusks suggests a paleo- 
depth of 100-200 m. and water temperatures of 10-12°C 
in summer months during the Oligocene (Allison & Mar- 
incovich, 1981). According to Hall (1964), this type of 
marine climate can be defined as cool-temperate. The 
Recent species Fulgoraria {Nipponomelon} prevostiana 
(Crosse, 1878) has been dredged from depths of 110- 
732 m off western Hokkaido, Japan, at a bottom tem- 
perature of 11.8°C (Weaver & du Pont, 1970:45). Water 
temperatures for the Alugian Formation thus seem to be 
colder than those in western America. These data more 
probably suggest a deeper depositional environment for 
the Alugian Formation, perhaps no shallower that 400- 
500 m, and therefore colder water. 

Oligocene climatic conditions, therefore, must have 
been quite favorable for a wider distribution of fulgo- 
rariine volutes that is seen today. This is clearK' supported 
by the fossil record (fig. 21). I5ased on the stratigraphic 
distribution and analysis of total species diversity of Ce- 
nozoic volutes, it has been deduced that the Oligocene- 
Early Miocene was the first period of wide distribution 
and high species diversity of northern Pacific Fulgora- 
riinae (Oleinik, 1990). There was, however, an appre- 
ciable amount of endemism, especially at the subgeneric 
level. Fulgoraria s.s. and Psephaea, for example, were 
restricted to the western Pacific, while Miopleiona is 
known onl\ from the northeastern Pacific. These re- 
stricted geographic ranges ma\ have been caused by 
different centers of origin of the North Pacific Fulgo- 



Page 146 



THE NAUTILUS, Vol. 106, No. 4 



rariinae, or were the results of differing dispersal strat- 
egies. Other subgenera, such as Musashia and Nippon- 
omelon, were abundant throughout the north Pacific. 
Total species diversity of Fulgorariinae, however, did 
not vary appreciably during the Oligocene of the north 
Pacific (fig. 22). 

ACKNOWLEDGMENTS 

I thank Dr. Yuri B. Gladenkov and Dr. V'alentina N. 
Sineiniko\a, of the laboratory of Phanaerozoic Stratig- 
raphy of the Geological Institute of the Russian Academ\' 
of Sciences, for providing material; Dr. Sergei I. Kiyash- 
ko, of the Institute of Marine Biology in Vladivostok for 
the oxygen isotope analyses. Photographs of the shells 
were taken by Mr. Andrew A. Okunev of the Geological 
Institute, Moscow. My special thanks to Dr. M. G. Har- 
asewych. National Museum of Natural History, Smith- 
sonian Institution, Washington, D.C., Dr. E. J. Petuch, 
Florida Atlantic University, Boca Raton, Florida, and 
Dr. L. K. Marincovich, U.S.G.S., Menlo Park, California 
for critical review of the manuscript. 



LITERATURE CITED 

Addicott, W. O. 1969. Tertiary Climatic change in the mar- 
ginal northeastern Pacific Ocean. Science 165:583-586. 

Addicott, W. O., S. Kanno, K. Sakamoto, and D. J. Miller, 1971. 
Clark's Tertiar\ molluscan types from the Yakataga dis- 
trict. Gulf of Alaska. U. S. Geological Survey Professional 
Paper 750-C:18-33. 

Allison, R. C. 1978. Late Oligocene through Pleistocene mol- 
luscan faunas in the Gulf of Alaska region. The V'eliger 
21(2);171-188. 

Allison, R. C. and L. Marincovich, Jr 1982. A late Oligocene 
or earliest Miocene molluscan fauna from Sitkinak Island, 
Alaska. U. S. Geological Survev Professional Paper 1233: 
lip. 

Cooke, A. H. 1922. The radula of the Volutidae Proceedings 
of the Malacological Society of London 15:6-11. 

Devyatilova, A. D. and V. I. Volobueva. 1981. Atlas of the 
Paleogene and Neogene fauna of the USSR north-east. 
Moscow, Nedra Publishers, 219 p. [in Russian]. 

Durham, J. W. 1944, Megafaunal zones of the Oligocene of 
northwestern Washington, University of California pub- 
lications, Department of Geological Sciences Bulletin 27(5): 
102-212. 

Encyclopedia of Oceanography. 1974. Leningrad, Hydro- 
meteozdat Publishers. 630 p. [in Russian]. 

Gladenkov, Yu. B., G. M. Bratseva, and V. N, Sinelnikova. 
1987, Marine Cenozoic of the Gulf of Korf, eastern part 
of Kamchatka, [In:] Sketches on the Geology of the north- 
east part of the Pacific tectonic belt. Moscow, Nauka Pub- 
lishers:5-6(), [in Russian], 

Habe, T. 1943. On the radulae of the Japanese marine gas- 
tropods. Venus 13:58-76, 

Hall, C. A., Jr. 1964. Shallow water marine climates and 
molluscan provinces. Ecology 45(2):226-234. 

Hickman, C. S. 1974. Characteristics of bathyal mollusk fau- 
nas in the Pacific coast Tertiary .Xiuuial Report of the 
Western Society of Malacologists 7:41-50. 



Kamada, Y. 1962. Tertiary Marine Mollusca from Joban Coal 
Field, Japan. Palaeontological Society of Japan, Special 
Paper 8:1-187, 

Kanehara, K. 1937, Miocene shells from the Joban Coal Field. 
Bulletin of the Imperial Geological Survev of Japan. 27(1): 
1-21, 

Khomenko, I, P 1933. .\bout the age of Tertiary deposits of 
the Gulf of Korf seashore in Kamchatka, Proceedings of 
the Far East Geological Survey 287:3-26. [in Russian]. 

Kidwell, S. M. 1988. Taphonomic comparison of passive and 
active continetal margins: Neogene shell beds of the At- 
lantic Coastal Plain and northern Gulf of California, Pa- 
leogeography, Paleoclimatology. Paleoecology 63:201-223. 

Krishtofovich, L. V, 1973, Cenozoic Mollusca [In:] New spe- 
cies of ancient plants and animals of the L'SSR. Leningrad, 
Nedra Publishers. Proceedings of the .\ll-Union Oil Re- 
search Intitute 313:77-78, [in Russian], 

Masuda, K, and H, Noda, 1976. Checklist and bibliography 
of the Tertiary and Quaternary mollusca of Japan. Saito 
Ho-on Kai:464 p. 

Moore, E, J. 1984. Molluscan paleontology and biostratigra- 
phy of the Lower Miocene upper part of the Lincoln Creek 
Formation in southwestern Washington Contributions in 
Science, Natural Historv Museum of Los Angeles Countv 
351:1-42. 

Noda, H. 1975. Turciculid Gastropoda of Japan. Sendai Sci- 
ence Reports of the Tohoku University, Second Series (Ge- 
ology) 45(2):51-82. 

Okutani, T. 1963. Report on the archibenthal and abyssal 
gastropoda mainly collected from Sagami Ba> and adja- 
cent waters by the R. V. "Soyo-maru ' during the years 
1955-1963. Journal of the Facult\ of Science. Universitv 
of Tokyo (Biology). 15(3):416-418, 

Oleinik, A. E. 1990, Cenozoic paleobiogeography of gastro- 
pod family Volutidae:view on Tethys-Pacific connections. 
Third International Symposium on Shallow Tethys Sen- 
dai, Japan:57-58. 

Oyama, K., A Mizuno. and T Sakamoto 1964, Illustrated 
handbook of Japanese Paleogene molluscs, Tokyo. Geo- 
logical Survey of Japan. 244 p. 

Pilsbry, H. A, and A, A. Olsson. 1954, Systems of the Volu- 
tidae, Bulletins of .American Paleontology 35(152):271- 
306. 

Scarlato, O. A. 1981. Bivalvesof the western Pacific temperate 
latitudes. Leningrad, Nauka Publishers. Handbooks of the 
USSR fauna edited by the Zoological Institute 126 479 p 
[in Russian], 

Shikama, T, 1954, On the Tertiary formations of Tomikusa 
in South Nagano Prefecture, Science Reports of the Yo- 
kahama National University. Biological and Geological 
Sciences 3:71-108. 

Shikama, T, 1967. System and evolution of Japanese fulgo- 
rarid Gastropoda. Science Reports of the Yokahama Na- 
tional Universitv, Biological and Geological Sciences 13: 
23-132. 

Tegland, N. M. 1933. The fauna of the t\pe Blakeb' Upper 
Oligocene of Washington. Universit\' of California Pub- 
lications, Department of Geological Sciences 23(3):81-174. 

Watanabe, T. and T. Habe. 19768. The largest specimen of 
Kurodina smithi Sowerbv and its radula. Venus 37(2);101- 
102. 

Weaver, C. S, and J E du Pont 1970 Living Volutes. A 
monograph of the living volutes of the world. Delaware 
Museum of Natural History, Greenville, xv + 375 p. 



THE NAUTILUS 106(4): 147-151, 1993 



Page 147 



A New Species of Columbellid Gastropod from the 
Old World Tropics 



William K. Emerson 

Department of Invertebrates 
American Museum of Natural History 
New York, NY 10024-5192, USA 



ABSTRACT 

Cotonopsis monfilsi new species, is described from off Senegal, 
West .Africa in 230-500 m. The generic assignment is made on 
the basis of shell, opercular, and radular characters. The radula 
is typically columbellid, whereas the shell morphology lacks 
some characteristics of the "Strombina" gastropods. Therefore, 
the placement in the S/romfoiia-group genus Cotonopsis is 
tentative, Metula dockeryi new name is proposed to replace 
Metula inflata Dockerv, 1984, not Metula inflata (Houbrick, 
1984). 

Key Words: Prosobranchia, Columbellidae, Cotonopsis, Buc- 
cinidae, Sdetula, west Africa, new species, new name. 



INTRODUCTION 

During the past five years, we have received from several 
shell dealers specimens of an apparently new species of 
columbellid gastropod for study and report. The taxo- 
nomic placement of these shells could not be made with 
certainty until the morphology of the radula was deter- 
mined. .\ radula was eventually found and proved to be 
typically columbellid in form As a result, this gastropod 
can be assigned provisionally to the Strombina-group 
genus Cotonopsis. 

I take pleasure in describing this new species in honor 
of Paul Monfils, who was the first to call tliis perplexing 
species to my attention. 

A review of the literature disclosed that a replacement 
name was required for a fossil species of Metula. A new 
name is provided for Metula inflata Dockery, 1984, not 
Metula inflata Houbrick, 1984. 

ABBREVIATIONS 

AMNH = American Museum of Natural History, New 
York 

LACM = Los Angeles County Museum of Natural His- 
tory, California. 



SYSTEMATICS 

Superfamily; Buccinoidea Rafinesque, 1815 
Family Columbellidae Swainson, 1840 
Genus Cotonopsis Olsson, 1942 

Type species (by original designation): Strombina {Co- 
tonopsis) panacostariceus Olsson, 1942, Pliocene of the 
Burica Peninsula, Costa Rica, Charco Azul Formation 

Cotonopsis monfilsi new species 
(figures 1-10) 

Description: Shell solid, fusiform, whorls inflated with- 
out a sutural ramp; protoconch of 3^2 smooth, convex 
whorls; teleoconch of IVz convex whorls, sculptured with 
numerous fine spiral lirae, numbering about 18 on the 
penultimate whorl and about 32 on the body w horl. Some 
spiral lines bifid; lirae not consistently evenly spaced. 
Thickened varix formed on body whorl above the rim 
of the aperture. Aperture ovate, inner lip terminally thin, 
but with a sublabial ridge with 10 Urate teeth; columella 
w ith 4-5 weak folds; siphonal canal open, short, narrow. 
Fasciole indistinct. Periostracum brownish tan. Back- 
ground color brownish buff-tan with broken whitish bands 
below the suture and with axial blotches interrupting the 
tanish coloration. Aperture white. Operculum thin, cor- 
neous, lenticular with a terminal nucleus at edge. Radular 
ribbon (figs. 9, 10) rachiglossate, typically columbellid 
(cf. Radwin, 1977, fig. 22b). Rachidian tooth a narrow, 
nearly rectangular plate. Lateral tooth shaft-like with 
barbed base, tip of shaft with two hooked cusps. 

Material examined: Holotype (AMNH 232519) and 3 
paratypes (AMNH 232091) from the t\pe localitv; 5 
Paratypes (AMNH 232092) dredged in 300-350 m', on 
muddy bottom, off St. Louis, Senegal, November, 1987, 
ex-Northeast Natural History Imports; 3 Paratypes 
(AMNH 232152) dredged in 450-500 m, on muddy-sand 
bottom, off St. Louis, Senegal, ex-Northeast Natural His- 
tory Imports; 6 referred specimens (5 specimens, AMNH 



Page 148 



THE NAUTILUS, Vol. 106, No. 4 




Figurrs 1-8. Cotonopsis monfilsi n. sp. 1-3, holotype (AMNH 232519) 4-6, paratype (AMNH 232091); figs. 1-6 xl.5. Figs. 
8, paratype (AMNH 232092), spire enlarged to show detail of earl\ whorls; figs. 7, 8, 5.0x. 



232176; 1 specimen, LACM 146956) dredged on silty- 
sand bottom, 230-260 m, off Casamance Province, Sen- 
egal (12''50'N,15°0'W), ex-Northeast Natural History Im- 
ports; 2 referred specimens (AMNH 255056) trawled in 
deep water from off the Dakar harbor entrance, Senegal 
(14°40'N, 17°26'W), ex-Mal de Mer Enterprises. See Ta- 
ble 1 for shell measurements. 

Type locality: dredged in 300 m off St Louis, Senegal, 
West Africa, d6°02'N,16°30'W) by Marcel Pin, 1987, ex- 
Abbey Specimen Shells. 

Distribution: Known only from oU Senegal, West Africa 
in 230 to 500 meters. 



Remarks: The shells of the present specimens resemble 
certain ""S7roni/n>irt"-group gastropods (such as Coton- 
opsis), but the\ are also in some respects reminiscent of 
certain buccinid gastropods (eg. Pisania. Mctula, etc.). 
Before the nature of the radula was known, I asked 
several colleagues for their opinions based on the ex- 
amination of photographs or the study of the specimens. 
Some of them referred the specimens to the buccinid 
genus Pisania Bivona-Bernardi, 1832, in the subfamily 
Pisaniinae Gray, 1857:13, formerly credited to Tryon, 
1881:98 (see Ceriiohorskv, 1971:138; 1975:192, Beu and 
Maxwell, 1987:56; and Bouchet, 1988:149, for comments 
on the subfamilial status of Pisaniinae in Buccinidae, and 



W. K. Emerson, 1993 



Page 149 




Table 1. Slu-ll climensioiis and proportions of the specimens 
of Cotonopsis monfilsi n. sp. n = 20 specimens. Width incfndes 
terminal \arix on k>odv whorl. Measurements in mm 



Figures 9, 10. Cotono|)>.i>. monfiUi n sp 9, holotype u^MNH 
2325f9), radula, about one-thirti of the ribbon 10, detail of the 
rachidian teeth; fig. 9, scale bar = 100 ^ni, fig 10, scale bar = 
200 ixm. courtesy of J. H. McLean 



its rejection without an explanation at tiie subfamilial 
level b\' Ponder and Waren, 1988:305) The resemblance 
of the new species to Recent species of Charitodoron 
(Mitridae) from South Africa was also suggested (see 
Lozouet, 1991:206, figs. 26-30). Other colleagues be- 
lieved the Senegalese specimens to be an undescribed 
columbellid species referable to Cotonopsis (sensu stric- 
to). 

With the recovery of a radula from a dried body of 
the new species, the placement of the new species in the 
family Columbellidae can be confirmed. The radular 
characters are t>pically columbellid in form (figs. 9, 10). 
As in Cotonopsis argentea (Houbrick, 1983:352, fig. 2), 
the radular dentition consists of rachidian teeth that are 
thin, narrow and form a rectangular plate. A wide space 
separates the rachidian teeth from the lateral teeth, which 
are composed of a shaft v\'ith an enlarged base and are 
tipped with two hooked cusps. The shells of the new- 
species possess the thickened varices of Stromhina-grou'p 
gastropods, but the apertural dentition of the inner lip 
differs in having evenly spaced lirate denticles of equal 
size. These are comparable to those of the buccinid genus 
Bartschia Rehder (1943:199, pi. 20, fig. 17; Olsson and 
Bayer, 1972:924, fig. 14). Furthermore, Bartschia sig- 
nificans. the type species, has strongly cancellate sculp- 
ture, whereas the new species possesses fine spiral lirae. 
The radular characters of Bartschia are unknown, but 
on shell features, Beu and Maxwell (1987:62) believe 
Bartschia, together with Metida H. and A. Adams (1853: 
84; Emerson, 1986:27), to be closely related to the buc- 
cinid genus Colubraria Schumacher, 1817. Some of the 
western Atlantic species of Metula [e.g. the West Indian 
M. (Agassitida) agassizi Clench and Aguayo, 1941:179, 
pi. 14, fig. 4; Olsson and Bayer, 1972:917, fig. 11 and the 
east African M. (Kanamarua) rehderi (Kilburn, 1977: 
193, fig. 21)] superficialK- resemble the new species. 
Bouchet (1988:150, fig. 1) illustrated the typically buc- 









Width- 


Type specimens 


Height 


Width 


height 


*AMNH 232519 (holotvpe) 


40.1 


14,9 


0.37 


AMNH 232091 (paratvpe 1) 


41,7 


1,5.2 


0.36 


AMNH 232091 (paratype 2) 


37.2 


13.7 


0.37 


AMNH 232091 (paratype 3) 


37.1 


13.6 


0.37 


AMNH 232092 (paratype 1) 


41.1 


14.4 


0.35 


AMNH 232092 (paratype 2) 


37.9 


13.4 


0.35 


AMNH 232092 (paratype 3) 


.37.1 


13.8 


0.35 


AMNH 232092 (paratype 4) 


37.5 


13.6 


0.36 


AMNH 232092 (paratype 5) 


36.2 


12.8 


0.35 


AMNH 2321.52 (paratvpe 1) 


404 


14.6 


0.36 


AMNH 232152 (paratvpe 2) 


36.9 


12.8 


0.35 


AMNH 232152 (paratype 3) 


34.5 


12.6 


0.37 


Referred specimens 








AMNH 232176, a 


39.7 


14.5 


0.37 


AMNH 232176, b 


36.3 


13.8 


0.38 


AMNH 232176, c 


36,1 


13.9 


0.39 


AMNH 232176, d 


35.0 


14.0 


0.40 


AMNH 2.32176, e 


34.9 


13.7 


0.40 


LACM 146956 


36.2 


13.9 


0.38 


AMNH 255056 a 


39.1 


14.1 


0.36 


AMNH 255056, b 


38.6 


13.4 


0.35 


Mean 


37.1 


13.8 


0.36 


Range 


34.5- 


12.6- 


0.35- 




41.7 


15.2 


40 



* Radula extracted. 



cinid radula of his new species, Metida africana, and 
concluded the radular morphology- is significantly dif- 
ferent from that of the buccinid genus Pisania. 

The placement of the new species in the genus Co- 
tonopsis based on shell characters is tentative. As the 
genus is defined by Jung (1989:158), the teleoconch whorls 
are smooth or axialK sculptured. In the new species, the 
teleoconch sculpture is composed of fine spiral lines and 
no axial sculpture is present. Cotonopsis is represented 
in the tropical western Atlantic by one species (C. ar- 
gentea). The remainder of the 15 species of Cotonopsis 
recognized by Jung (1989), 2 fossil and 12 living species, 
occur in the west American tropical zone. Previously, 
the Strombina-gToup gastropods were believed to be re- 
stricted to the New World tropics, dating from the Mio- 
cene to the present, w ith only 3 of the 35 living species 
occurring in the western Atlantic (Jung, 1989:4). There- 
fore, the presence of a species of Cotonopsis in west 
African waters may reflect a separate lineage of these 
columbellids that may eventually be determined to re- 
quire recognization as a new genus-group taxon. 

REPLACEMENT NAME FOR A HOMONYM 

During my review of the literature for this paper, I noted 
a homonymic species-group name in Metula (Buccini- 
dae). Metula {Metula) inflata Dockery in MacNeil and 



Page 150 



THE NAUTILUS, Vol. 106, No. 4 



Dockery (1984:331, pi. 52, fig. 3; published in "Novem- 
ber, 1984", teste D. T. Dockery III, in Hit.. Dec. 9, 1991) 
is a junior secondary homonym of Acamptochetus [=Me- 
tiila] inflatus Houbrick (1984:421, fig. 1; published July 
6, 1984). I here rename Metula inflata Dockery, 1984 
(not Metula inflata [Houbrick, 1984]), as Metula dock- 
eriji Emerson, NEW NAME. Dockery s ta.xon is a Pa- 
leogene fossil from Mississippi and Houbrick s ta.xon is a 
Recent Philippine species. 

Acamptochetus Cossmann (1901:123; type species by 
original designation: Murex mitraeformis Brocchi, 1814) 
is a junior subjective synonym of Metula H. and A. Ad- 
ams (1853:84; type species by subsequent designation 
Kobelt, 1876: Buccinum clathratum Adams and Reeve, 
1850, [not Kiener, 1834, nor Anton, 1839] = Metula 
atnosi Vanatta, 1913:22), as noted by Emerson (1986:27) 
and Beu and Maxwell (1987:62) and accepted by Bouchet 
(1988:149). 



ACKNOWLEDGEMENTS 

In addition to Paul Monfils of Northeast Natural History 
Imports, Providence, RI, I am grateful to Robert Foster 
of Abbey Specimen Shells of Santa Barbara, CA and 
Robert and Dorothy Janowsky of Mai De Mer Enter- 
prises, West Hempstead, NY for providing specimens for 
study. Silvard P. Kool of the Museum of Comparative 
Zoology, Harvard University, kindly lent specimens of 
Metula for comparisons. The following colleagues co- 
operatively offered comments on the taxonomic place- 
ment of the new species before the nature of the radula 
was known: Walter O. Cernohorsky, M. G. Harasewych, 
Richard S. Houbrick, Peter Jung, R. N. Kilburn, Jorgen 
Knudsen, Paula M. Mikkelsen, Gary Rosenberg, and Gra- 
ham Saunders. James H. McLean generously mounted 
the radula and provided the photographs for the radula. 
I am indebted to my AMNH colleagues, Walter E. Sage, 
III for technical assistance, Andrew S. Modell for pho- 
tographic services and Stephanie Grooms for word-pro- 
cessing the manuscript. 

LITERATURE CITED 

Adams, H. and A. 1853 [-1852]. The Genera of recent Mol- 
lusca, arranged according to their organization, London 
l(13);65-96, pis. 9-12 (June, 1853). 

Anton, H. E. 1839. Verzeichniss der Conchy lien vvelche sich 
in der Sammlung von Hermann Eduard .^nton befinden. 
Halle, xvi -t- 110 p. 

Beu, A. G. and P. A. Maxwell 1987. A revision of the fossil 
and living gastropods, related to Picsiotriton Fischer, 1884 
(Family Gancellariidae, subfamily Plesiotritoninae n, 
subfam.) with an Appendix: Genera of Buccinidae Pisa- 
niinae related to Cotubraria Schumacher, 1817. New Zea- 
land Geological Survey Paleontological Bulletin 54:1-140. 

Bivona-Bernardi, A. 1832, Caratteri dun nuovo genere di 
conchiglie della famiglia delle columellarie del Signer De 
Lamarck Effemeridi Scientifiche e Letterarie per la Sicilia 
2:8-13, 1 pi. 

Bouchet, P. 1988. Two new species of Metula (Gastropoda: 



Buccinidae) with a description of the radula of the genus. 
The Nautilus 102(4):149-153. 

Cernohorsky, W O. 1971. Indo-Pacific Pisaniinae (Mollusca: 
Gastropoda) and related buccinid genera. Records of the 
Auckland Institute and Museum 8:1.37-167. 

Cernohorsky, W. O. 1975. Supplementary notes on the tax- 
onomy of buccinid species of the subfamily Pisaniinae 
(Mollusca: Gastropoda). Records of the Auckland Institute 
and Museum 12:175-211. 

Clench, W. J. and C. G. Aguayo. 1941. Notes and descriptions 
of new deep-water Mollusca obtained by the Harvard- 
Havana Expedition off the coast of Cuba. Memorias de la 
Sociedad Cubana de Historia Natural. Ha\ana 15(2): 177- 
180, 

Cossmann, M 1901 Essais de Paleconchologie Comparee, pt 
4, 293 p, 

Emerson, W. K. 1986. On the type species of Metula H and 
A. Adams, 1853: Buccinum clathratum A. .Adams and 
Reeve, 1850 (Gastropoda: Buccinidae), The Nautilus 100(1): 
27-30, 

Gray, J. E. 1857, Guide to the systematic distribution of 
Mollusca in the British Museum. Part 1, London, 230 p. 

Houbrick, R. S, 1983. A new Strombina species (Gastropoda; 
Prosobranchia) from the tropical western .Atlantic. Pro- 
ceedings of the Biological Society of Washington 96(3): 
349-3.54, 

Houbrick, R, S. 1984. A new "Me/i/Za" species from the Indo- 
West Pacific (Prosobranchia Buccinidae). Proceedings of 
the Biological Society of Washington 97(2):420-424 (pub- 
lished July 6, 1984).' 

Jung, P. 1989 Revision of the S(roni/>i;!a-Group (Gastropoda: 
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1-298. 

Kiener, L. C. 1834 [-1841]. Species general et iconographie 
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Kilburn, R, N 1977, Taxonomic studies on the marine Mol- 
lusca of southern .Africa and Mozambique Part 1, .Annals 
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Kobelt, W. 1876 [-1881]. Illustriertes Conchylienbuch Nu- 
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Lozouet, P. 1991. Mollusca Gastropoda: Eumiira recentes de 
la region neo-caledonienne et Charitodoron fossiles de 
I Oligocene superieur d .AquitainelMitridae) hi. t>osnier, 
A. and P. Bouchet (eds. ) Resultats des Campagnes Ml'- 
SORSTOM, vol 7, no, 8. Memoires de la Museum National 
d'Histoire Naturelle, Ser .A, Zoologie 1.50:205-222 

MacNeil, F. S, and D. T Dockery. Ill 1984 Lower Oligocene 
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(published in November, 1984). 

Olsson, A. A. 1942. Tertiary and Quaternary fossils from 
Burica Peninsula of Panama and Costa Rica. Bulletins of 
American Paleontology 27(106):153-258, 

Olsson, A A. and F M Bayer 1972 .American metulas (Gas- 
tropoda: Buccinidae). Bulletin of Marine Science (Uni- 
versity Miami, FL) 22(4):900-925, 

Ponder, W. F. and A. Waren. 1988. In. Ponder, W. F., D. J. 
Eernisse, and J. H. Waterhouse, eds. Prosobranch Phylog- 
eny, .Appendix, Classification of the Caenogastropoda and 
Heterostropha — .A list of the family-group names and 
higher taxa Malacological Re\ie\\ . Supplement 4:288-326. 

Radwin, G. E. 1977. The family Columbellidae in the western 
Atlantic. The N'eliger 19(4):403-417. 



W. K. Emerson, 1993 Page 151 



Rehder, H. A. 1943. New marine mollusks from the Antilleaii 3:1-310, Academy of Natural Sciences, Pfiiladelpfiia [p. 

region. Proceedings of the United States National Museum 98,1881] 

93(3161):187-203, pis. 19, 20. Vanatta, E. G. 1913. Descriptions of new species of marine 

Tryon, G. W. [1880-] 1881. Manual of conchology; structural shells. Proceedings of the -Academy of Natural Sciences, 

and systematic. Series 1, Tritonidae, Fusidae, Buccinidae, Philadelphia 65(l):22-27, 3 text figs., pi. 2. 



THE NAUTILUS 106(4):152-154, 1993 



Page 152 



The Association between the Gastropod Biiccinaiiops cochlidium 
(Dillwyn, 1817) and the Sea Anemone Phlyctenanthus australis 
Carlgren, 1949 in Patagonian Shallow Waters 



Guido Pastorino 

Division Paleozoologia Invertebrados 

Museo de Ciencias Naturales 

Paseo del Bosque s/ n 

1900 La Plata, Buenos Aires, 

Argentina 



ABSTRACT 

The association between Buccinanops cochlidium (Dillwyn) 
and Phlyctenanthus australis Carlgren is described. Both spe- 
cies inhabit in the sandy infralittoral of Puerto Piramide, Chu- 
but Province, Argentina, The densities and percentages of gas- 
tropods with anemones attached are reported, and the possible 
benefits to the sea anemone and the gastropod are discussed. 

Key ivords: Symbiosis, Ecology, Anemones, Buccinanops. 



INTRODUCTION 

The association between actinians and gastropod shells 
occupied bv hermit crabs has been frequently described 
(e. g. Faurot, 1910, 1932; Brunelli, 1913; Ross^ 1960; Ross 
& Sutton, 1961a, 1961b; McLean and Mariscal, 1973; 
Balasch et a/., 1977; Brooks and Mariscal, 1986). How- 
ever few papers have been published describing the re- 
lationship between actinians and living gastropods in the 
South Atlantic (Bellisio et ai 1972; Arnaud, 1978). 

This paper reports on the association between Bucci- 
nanops cochlidium (Dillwyn, 1817) and Phhjctenanthiis 
australis Carlgren, 1949, from northern Patagonia. 

MATERIALS AND METHODS 

Specimens of Buccinanops cochlidium and B. globulos- 
um were collected during November 1989 and October 
1990 while SCUBA diving at depths of 5-11 m in the 
sandy infralittoral zone off Puerto Piramide (42°35'S, 
64°17'W), Chubut Province, Argentina. In February 1992, 
18 samples were collected from various tlepths. The in- 
formation obtained from each .sample included depth, 
area, number of individuals of each species of gastropod, 
and number of anemones attached per gastropod. Data 
from samples were combined to give a general pattern 
for Puerto Piramide. DejJth was determined using a cap- 
illary depth gauge (Dacor model lA'.G & MCXi). 

The material was first relaxed at low temperatures and 



then preserved in 70 % formalin. The specimens are 
housed at the Division Zoologia Invertebrados, Museo de 
Ciencias Naturales, La Plata, Buenos Aires, Argentina. 

RESULTS 

The anemone Phlyctenanthus australis commonly cov- 
ers the entire dorsal surface of the mollusk shell with its 
pedal disc, leaving only the aperture free (fig. 4). In some 
cases as many as two specimens of P. australis were 
attached to a single gastropod (fig. 5). Usually only a 
single anemone is attached either to the spire or the body 
whorl. When in situ specimens are viewed from above, 
only the completely extended oral disc of the anemone 
is visible, with the tip of the siphon of B. cochlidium 
extending from beneath its edge. The gastropods usually 
remain completely buried. Occasionally, crawling spec- 
imens with more than one attached anemone were ob- 
served. 




-42-S 



Dib C. Tremouilles 



Figure I. Map of Peninsula Valdes area showing the location 
of the study area. 



G. Pastorino, 1993 



Page 153 




Figures 2-5. Buccinanops cochlidium (DilKvyn) with Phhjctenanthus australis Carlgren attached. 1. Oblique apertural view of 
the gastropod showing egg-capsules attached to the columellar callus. 2. Dorsal view of the same specimen. 3. P. australis covering 
the entire shell of the gastropod. 4. B. cochlidium with two anemones attached (arrowheads). All x 1 

Table I. Halli\ metric distriljuliDii oi liuiiinaniipn glohi»iiint. B cochliiliuiu. ami Flihiitcnantlnis australis at Puerto Piriimide, 
\ri;eiitina 



l)c|)lh range 

(ni) 



5-5.9 



(i-B.9 



8-8.9 



9-9.9 



10-10 9 



No. samples 

Species 

Gastropods nr- 

SD 

% with one anemone 

% with two anemones 

'"c wilhuul ancinuiic^ 









B. 


C0( 


■hlid 


f. 




28.3 


0.60 


0.51 


0.68 


049 


13.8 


0.31 


0.35 








0.16 


10 


— 


82.35 


73.4 








67.74 


66.6 


— 


— 


13.3 








22.58 


20.0 


100 


17 (i5 


1 . ■) . ■) 








9(iS 


13 4 



Page 154 



THE NAUTILUS, Vol. 106, No. 4 



As samples were collected just after the hatching pe- 
riod of B. globiilosum . their densit\ was probably over- 
estimated. However, B. globulosum is alwa\s far more 
abundant than B. cochlidium. Of the total gastropods 
sampled 72.53% had one anemone attached, 13.97% had 
two. Anemone free specimens comprised only 13.50% of 
the sample. No correlation between depth and anemone/ 
gastropod density was found. 

DISCUSSION 

Two species of Buccinanops. B. glolmlosum (Kiener, 
1834) and B. cochlidium (Dillwyn, 1817), inhabit the 
sublittoral zone near Puerto Piramide. Buccinanops glob- 
ulosum ranges from the lower midlittoral into the sub- 
littoral to a depth of 6 m below low tide. At depths greater 
than 6 m it is replaced b> B. cochlidium (see Table 1). 
Buccinanops cochlidium is the only gastropod that par- 
ticipates in the gastropod-anemone association, although 
Phlyctenanthus australis was also found attached to rocks 
in the intertidal zone. Phlyctenanthus australis was not 
previously known from south of Mar del Plata. At this 
locality it may be attached to the decapod Libinia spi- 
nosa Milne-Edwards, 1834 as noted by Zamponi (1977). 
.\mong the other species of gastropods living in the area 
onK Odontocymbiola magellanica (Gmelin, 1791) is large 
enough to be able to carry attached anemones, but P. 
australis is not known to attach itself to the shell of this 
volutid. 

Ross (1974) suggested that one of the possible benefits 
to the anemone was increased mobility, enabling it to 
obtain greater amounts of food and also to avoid adverse 
physiological conditions. The advantages for the gastro- 
pod, on the other hand, are not as clear. McLean and 
Mariscal (1973) and Brooks (1980) suggested that, in the 
case of the hermit crabs, protection provided by the 
nematocysts of the anemone would be a benefit. This 
ma>' also be applicable to B. cochlidium . The gastropod 
apparently is not adversely affected. This may be in- 
ferred from the normal reproductive activity, indicated 
by the egg-capsules attached to the columellar callus of 
the females (figs. 1, 3). 

ACKNOWLEDGMENTS 

I wish to acknowledge M. L. Pastorino who supported 
part of the travel expenses. J. Muzon and T. Simanauskas 
gave valuable comments. I am grateful M. Griffin's help 
with the English version of the manuscript. The manu- 
script benefited from reviews by G. Vermeij and an anon- 
\mous reviser. This work was carried out during the 
tenure of a scholarship granted by the Consejo Nacional 
de Investigaciones Cientificas y Tecnicas (CONICET), 
Argentina. 



LITERATURE CITED 

.\rnaud, P. M. 1978. Observations ecologiques et biologiques 
surle Voliitidae antarctique Harpovohila charcoti (Lamy, 
1910) (Gastropoda prosobranchia). Haliotis 7:44-46 

Balasch, J., J. Cuadras and G, Alonso 1977, Distribution of 
Calliactis parasitica on gastropod shells inhabited by Dar- 
danus arrosor. Marine Behavior and Physiology 5(1):37- 
44. 

Bellisio, N. B., R. B. Lopez and A. Tomo. 1972. Distribucion 
vertical de la fauna bentonica en tres localidades antar- 
ticas; Bahia Esperanza, Isla Peterman y ,\rchipielago Mel- 
chior. Contribucion del Instituto Antartico .'Krgentinol42: 
1-87, 

Brooks, W, R. 1988, The influence of the location and abun- 
dance of the sea anemone Calliactis tricolor (LeSueur) in 
protecting hermit crabs from octopus predators. Journal 
of E.\perimental Marine Biology and Ecology 116:15-21, 

Brooks, W. R. and R. N. Mariscal. 1986. Population variation 
and behavioral changes in two pagurids in association with 
the sea anemone Calliactis tricolor (LeSueur). Journal of 
Experimental Marine Biology and Ecology 103:275-289. 

Brunelli, G 1913, Ricerche etologiche. Osservazioni edes- 
perienze sulla simbiosi dei Paguridi e delle .Attinie Zool- 
ogische Jahrbucher, ,'\bteilung fur allgemeine Zoologieund 
Physiologic der Tiere, Jena 34:1-26. 

Faurot, L. 1910. Etude sur les associations entre les Pagures 
et les Actinies; Eupagurus Prideauxi Heller et Adamsia 
palliata Forbes, Pagurus striatus Latreille et Sagartia par- 
asitica Gosse. .Archives de Zoologie experimentale et ge- 
nerale 5:421-86. 

Faurot, L. 1932. .'Actinies et Pagures. Etude de ps\chologie 
animale. Archives de Zoologie experimentale et generale 
74:139-54. 

McLean, R. B. and R. N. Mariscal. 1973. Protection of a 
hermit crab by its symbiotic sea anemone Calliactis tri- 
color. Experientia 29:128-130. 

Ross, D. M, 1960. The association between the hermit crab 
Eupagurus hernhardus (L) and the sea anemone Cal- 
liactis parasitica (Couch), Proceedings of the Zoological 
Society of London 1.34(1 ):4.3-.57, 

Ross, D. M. 1974, Behavior patterns in associations and in- 
teractions with other animals. In: L. Muscatine and H. M. 
Lenhoff (eds.) Coelenterate Biology, Academic Press, New 
York, p. 163-212. 

Ross, D. M. and L. Sutton, 1961a, The association between 
the hermit crab Dardanus arrosor (Herbst) and the sea 
anemone Calliactis parasitica (Couch). Proceedings of the 
Royal Societ) of London 155:282-291. 

Ross, D. M. and L. Sutton, 1961b The response of the sea 
anemone Calliactis parasitica to shells of the hermit crab 
Pagurus hernhardus. Proceedings of the Roval Society of 
London 155:266-281. 

Zamponi. M O 1977. La anemonofauna de Mar del Plata y 
localidades vecinas. 1. Las anemonas Boloceroidaria y En- 
domvaria Coelenterata, .Actiniaria Neotropica 23(70): 133- 
136.' 



THE NAUTILUS 106{4):155-173. 1993 



Page 155 



Patterns of Diversity and Extinction in Transmarian 
Muricacean, Buccinacean, and Conacean Gastropods 



Edward J. Peluch 

Deparlnieiit of Cifology 
Florida Atlantic I'niversity 
Boca Raton, FL 33-131, USA 



ABSTRACT 

.•\n analysis of patterns of species-richness in ten muricacean, 
buccinacean, and conacean families found in the Maryland 
Miocene formations has uncovered a previously-undetected 
Serravallian Stage extinction event that decimated the fauna 
of the Calvertiaii Subprovince of the Transmarian Province 
Prior to the extinction event, the molluscan fauna of the Langhi- 
an Stage (Calvert Formation) contained man\ tropical elements 
(here referred to as "caloosagenic taxa ) that were derivatives 
of the Miocene Caloosahatchian Province. At the Langhian- 
Serravallian boundary and the initiation of Choptank deposi- 
tion, most of the earlier caloosagenic taxa suffered a regional 
extinction. Breakdown of local ecosystems, probably due to a 
cooling marine climate, increased toward the end of Choptank 
time, culminating in the extinction of prominent endemic thai- 
did and turriil genera During the latest Serravallian Stage and 
the beginning of Tortonian time (St. Mar\ s Formation), the 
marine climate returned to Langhian conditions and there was 
an acconipan\ing return to pre-Choptank levels of diversity 
The post-Choptank molluscan assemblages differed, however, 
in containing new caloosagenic taxa, such as Conus, and had 
an extremely reduced thaidid fauna Several biogeographically 
important new taxa are also proposed and these include three 
new genera, Patuxentrophon n.gen (Muricidae: Trophoninae), 
Calvcrturris n gen. (Turridae: Mangeliinael, and Transma- 
riaturris n gen (Turridae: Mangeliinae), and six new species, 
Ecphorostjcon lindajoijceac n sp. (Thaididae: Ecphorinae), 
Biiccinoftisus paluxentcnsis n.sp. (Fasciolariidae), Busycnty- 
pus choptankciuis n.sp., Turrifulgur miinjiandicus n.sp., and 
Turrtfulgur prunicola n.sp. (all Melongenidae: Busyconinae), 
and Calvcrturris .schniidti (Turridae: Mangeliinae). 

Keywords: Extinction, Maryland, Miocene, Transmarian Prov- 
ince. 



INTRODUCTION 

As a biogeographical entity, the Transmarian Molluscan 
Province of the northern and central Atlantic Coastal 
Plain has been relatively unstudied. This early Neogene 
paratropical province, whose fauna evolved in the Oli- 
gocene and became extinct during the Messinian Mio- 
cene, has recently been shown to contain numerous en- 
demic gastropod taxa (Petuch, I988a,b,c). Primary among 
these are a plethora of distinctive genera in the super- 



families Muricacea, Buccinacea, and Conacea, many of 
which are indicators of the provincial boundaries. 

At its height, the Transmarian province extended from 
the present-day Cape Fear, North Carolina area north- 
ward to at least Nova Scotia (figure 46), and encompassed 
three distinct subprovinces; the southern Pungoian Sub- 
province, the central Calvertian Subprovince, and the 
northern Sankatian Subprovince (Petuch, 1988b). The 
Pungoian Subprovince was centered on the Miocene Al- 
bemarle Embayment of North Carolina, while the Cal- 
vertian Subprovince had its focus on the diverse biotopes 
of the Salisbury Embayment of Chesapeake Bay, south- 
ern New Jersey , Delaware, Maryland, and northern Vir- 
ginia. Since the best-preserved and most accessible Trans- 
marian faunas are located along the Patuxent River, the 
St. Mary's River and the western shore of Chesapeake 
Bay in Maryland, and since these were the principal 
faunas used in my provincial analysis, 1 will deal only 
with Calvertian taxa in this paper. 

In Maryland, the Calvertian Transmarian molluscan 
fauna is contained within several extremely fossiliferous 
members of three classic Miocene formations and a pos- 
sible new formation, the Calvert (contemporaneous with 
the well-known Kirkwood Formation of New Jersey) 
(Langhian Stage), the Choptank (early Serravallian Stage), 
the Little Cove Point Unit (late Serravallian Stage) (the 
status of this unit is still in debate among authorities; I 
here follow the nomenclature of Ward and Blackwelder, 
1980:D4), and the St. Mary's (latest Servallian-early Tor- 
tonian Stages). Within these formations, a molluscan ex- 
tinction event, interspersed between two periods of spe- 
ciation and diversity augmentation, can be recognized 
within the stratigraphic record and these give insight 
into possible middle Miocene catastrophic climatic fluc- 
tuations along the eastern coast of North America. In 
this paper, I also hope to shed some light on the higher 
order diversity patterns of the Transmarian muricacean, 
buccinacean, and conacean gastropod families. Several 
important new taxa are also described here in order to 
fill some nomenciatural gaps in the Calvertian fossil rec- 
ord. These include three new genera, Patuxentrophon 
(Muricidae: Trophoninae), Calverturris (Turridae), and 
Transmariaturris (Turridae), and new species of Ec- 



Page 156 



THE NAUTILUS, Vol. 106, No. 4 



phorosycon (Thaididae: Ecphorinae), Buccinofusus 
(Fasciolariidae). Busycotypus (Melongenidae: Biis\con- 
inae), Turrifulgur (Melongenidae: Busyconinae), and 
Calverturris (Turridae). The three superfamilies studied, 
which include ten families and fort> -one genera (listed 
here in Appendi.v 1 ), are excellent indicators of ecosystem 
collapse during times of e.xtrenie climatological stress. 

THE TRANSMARIAN GASTROPOD FAUNA 

As demonstrated in my earlier work (Petuch, 1988b: 11), 
the Transmarian molluscan fauna represents one of the 
strangest mixtures of gastropods that can be found an\- 
where in the fossil record. Unlike any known Recent 
malacofauna, the Transmarian assemblages contain sym- 
patric suites of both boreal, high arctic gastropod genera 
such as Boreotrophon (Muricidae), Admete (Cancellari- 
idae), Oenopota (Turridae), and Euspira (Natici- 
dae), and eutropical genera such as Conits (Leptoconus) 
(Conidae). Striotercbriim ^Terebridae), Laevityphis 
(Muricidae), and Cymatosyrinx (Turridae). These arctic 
and tropical elements coexisted with a large compliment 
of endemic Transmarian genera. Although the Trans- 
marian Province contained taxa from several tropical 
families, the fauna lacked most of the classic eutropical 
index groups, such as the Strombidae, Turbinellidae, Cy- 
praeidae, Ovulidae, Cerithiidae, Potamididae, and Ly- 
riinae (N'olutidae) (Petuch, 1988b). Based on the absence 
of these eutropical elements, the Transmarian province 
can only be considered to have been a paratropical faunal 
region. 

\s determined by Gibson (1967), using foraminiferal 
assemblages, the marine climate of the Transmarian 
Province ranged from cool-temperate to temperate, with 
only a small seasonal fluctuation in water temperature. 
This climatic stabilit\ allowed the evolution of a physi- 
ologicalK stenothermal temperate molluscan fauna, with 
the endemic genera being unable to live in both the 
colder boreal conditions to the north and in the warmer 
tropical conditions of the Miocene Caloosahatchian Pro\- 
ince to the south (Petuch, 1988b) (figure 46). The Trans- 
marian arctic and tropical elements represent physiolog- 
ically eurythermal "invaders" from the Miocene Boreal 
and Caloosahatchian Provinces. These opportunistic spe- 
cies appear to have evolved physiological tolerations for 
the warmer (for the boreal taxa) or colder (for the tropical 
taxa) water conditions of the Transmarian region, and 
evolved their own endemic Transmarian temperate spe- 
cies complexes. 

Transmarian faunal elements that were derived from 
the tropical Miocene Caloosahatchian Province are here 
referred to as "caloosagenic". The caloosagenic influence 
varied through time in the Calvertian Subprovince, with 
some stages, such as the early Tortonian, containing many 
tropical taxa and other stages, such as the early Serra- 
vallian, containing fewer taxa. Some caloosagens were 
present only during the Langhian Stage and disappeared 
from Mar\ land during the Serravallian Stage, being found 
subsequently in the fossil beds of Virginia and Florida. 



Primary examples of these early Calvertian caloosagens 
include Phyllonolus [P. millvillcnsis (Richards and Har- 
bison, 1942)), Sfurcxiclla (M. shilohensis (Heilprin, 
1887)), Oliva (O. harrisi Martin, 1904), Amaea (A. re- 
ticulata Martin, 1904 and A. prunicola Martin, 1904), 
Niso (N. lineata (Conrad, 1841)), Ficus (F. harrisi (Mar- 
tin, 1904)), and Architectonica (A. trilineatum (Conrad, 
1841)). Other caloosagens make their first appearance in 
the Calvertian Subprovince at the very end of Salisbury 
deposition, in the late Serravallian and early Tortonian 
Stages. Some of these late Calvertian caloosagenic taxa 
include the previously-mentioned conid genus Conus 
(Leptoconus species complex) and the volutacean genus 
Mitra (A/, mariana Martin, 1904). Still other caloosa- 
genic groups, such as the buss conine genera Busycutypus 
and Turrifulgur, the xolutid Volutifusus, and the tur- 
ritellid Torcula, existed within the Calvertian area 
throughout the Langhian, Serra\allian, and Tortonian 
Stages. 

Transmarian faunal elements that were derived from 
the arctic Boreal Province are here referred to as "boreo- 
gens". As in the case of the caloosagens, the boreogenic 
influence also fluctuated through time, being more prev- 
alent in some molluscan assemblages than in others. Un- 
like the caloosagens, however, very few^ boreogens were 
present in the Calvertian Subprovince during the Lan- 
ghian stage. Of these, only a single genus (the naticid 
Euspira) is known to have ranged throughout the entire 
Maryland Miocene. The Tortonian Stage, on the other 
hand, saw the appearance of many new boreogenic taxa 
that previously had not existed within the Salisbury Em- 
bavment. Examples ot these later boreal "invaders' in- 
clude the previously-mentioned genera Admete and 
Oenopota. 



DYNAMICS OF TRANSMARIAN 
DIVERSITY 

By recognizing that three separate groups of faunal el- 
ements coexisted within the Transmarian Province, the 
caloosagens, the boreogens, and the Transmarian endem- 
ics, it is now possible to undertake a fine-resolution anal- 
ysis of the diversit)' patterns of C-alvertian Transmarian 
muricacean, buccinacean, and conacean gastropods. 
These are discussed in the following sections. A listing 
of all known Calvertian species in these three superfam- 
ilies is given in Appendix 2. 

Muricacean diversity patterns: .Although comprising two 
families and fourteen genera, the Transmarian murica- 
cean taxa were not distributed equally over time. Only 
two muricid genera, Laevityphis and Chesatrophon (fig- 
ure 32), and one thaidid genus, Ecphora s.s., are tem- 
poralK distributeil throughout the Langhian, Serraval- 
lian, and Tortonian Stages. During Langhian time, in 
the Calvert and Kirkwood (New Jersey) Formations, only 
two other muricid groups, the caloosagens Murexiella, 
and Phyllonutns. are present. The Langhian ecphorine 
thaidids, on the other hand, underwent a large species 



E. J. Petuch, 1993 



Page 157 



radiation, evolving at least four species of Ecphora s.s., 
at least five species of Trisecphora (figure 38), two species 
of Ecphorosycon. and two species of Chesathais (figure 
34) (Petuch, 1989). (Note: The genus Chesathais, al- 
though possibly arising from a Caloosahatchian stem spe- 
cies (C. biconiciis), was represented in Maryland by the 
C. ecclesiastictis species complex and can be considered 
to be an endemic Transmariaii radiation. ) The large ec- 
phorines apparently dominated the Langhian molluscan 
assemblages, as their shells are abundant in most units 
of the Calvert Formation. The small Calvert muricids of 
the genera Laevitijphis. Chcsatrophon, Miirexiella. and 
Patuxentrophon (figures 1, 2), however, are rare in most 
assemblages and appear to have occupied the ecological 
"fringe" areas. 

During Serravallian time, the Maryland Mtirexiella 
species disappeared, (and presumabK the New Jersey 
Kirkwood Phyllonotus species), while the caloosagen 
Urosalpinx first made an appearance. This ubiquitous 
Caloosahatchian ocenebrine ta.xon survived to become 
an abundant component of the late Serravallian-Tortoni- 
an assemblages of the St. Marys Formation, and is also 
extant in the Recent. Also appearing in the early Ser- 
ravallian was the small muricid Stephanosalpinx (figure 
31). This distinctive muricacean disappeared by the late 
Serravallian and is confined to the Choptank Formation. 

As during the Langhian, the predominant muricacean 
radiation during the early and middle Serravallian time 
was a complex of ecphorine thaidids. During this time, 
the genus Ecphora s.s. was abundantly represented by 
at least four species and four stratigraphic subspecies. 
The subgenus Trisecphora, although not as species-rich 
as during Langhian time, was also abundantly repre- 
sented by two species and at least one stratigraphic sub- 
species. Likewise, the morphologically-conservative ge- 
nus Chesathais was also present in the Choptank 
Formation, in this case as a species and two stratigraphic 
subspecies. Although the primitive genus Ecphorosijcon 
was originally thought to have become extinct at the end 
of Langhian time (Petuch, 1989:43), the recent discovery 
of a rare Choptank species (£. lindajoyceae n.sp., de- 
scribed here) demonstrates that the genus survived into 
earliest Serravallian time and was part of the rich Chop- 
tank ecphorine fauna. 

Although ecphorine species diversity declined at the 
end of Choptank time, muricid diversity increased dra- 
matically at the beginning of Little Cove Point-St. Mary's 
time (late Serravallian-Tortonian). The abundant pres- 
ence of the trophonine genera Chcsatrophon (figure 32), 
Boreotrophon, (figure 29, 30) Scalaspira (figure 4), Li- 
rosoma (figure 3), and the ocenebrine genera Urosalpinx 
and Mariasalpinx (figure 33), particularly underscore 
this shift toward muricid dominance. This predominance 
is emphasized even further by the fact that only a single 
species of Ecphora occurs sympatrically with the rich 
trophonine and ocenebrine faunas found within the var- 
ious members of the St. Mary's Formation. Although only 
a single Ecphora species is found within any bed of the 
St. Mary's Formation, these species are much larger than 



any ancestral species from the Calvert and Choptank 
Formations, and are, in fact, the largest muricaceans 
found in the entire Maryland Miocene {Ecphora gard- 
nerae Wilson, 1987 often exceeds lengths of 110 mm). 
This gigantism gave the post-Choptank ecphorine sur- 
vivors an ecological advantage over their more numer- 
ous, but much smaller, muricid cousins — allowing them 
to utilize large prey items, such as adult specimens of 
the bivalves Mercenaria and Dosinia, which were in- 
accessible to the contemporaneous muricids (Petuch, 1989: 
15). Interestingly enough, of all the late Transmarian 
endemic muricid genera, only Scalaspira, Lirosoma, and 
Boreotrophon (Transmarian complex) survived into the 
early Pliocene (Zanclian Stage). The last species of these 
groups, S. strumosa (Conrad, 1832), L. sulcosa (Conrad, 
1830), and B. tetricus (Conrad, 1832), respectively, can 
be found in the Yorktown Formation. 

Buccinacean diversity patterns: Comprising five fam- 
ilies and twelve genera, the Transmarian buccinacean 
gastropods, unlike the muricaceans, were more equitably 
distributed over time. Eight of the twelve genera are 
found in all three Maryland Miocene formations and are 
represented by continuous species lineages. Only during 
St. Mary's time do additional genera appear, and these 
include the Transmarian endemics Mariafusus (figure 
39), Pseudaptyxis (figure 35), and Bulliopsis. No boreo- 
genic buccinacean taxa are known from the Pungoian, 
Calvertian, or Sankatian Subprovinces. 

Of the five Transmarian buccinacean families, only 
the Fasciolariidae showed any appreciable augmentation 
through time. During Calvert and Choptank time, the 
family Fasciolariidae was represented by only a single 
genus, Buccinoftisus (with two of the species shown here 
in figures 7, 13, 14). By Little Cove Point-St. Mary's time, 
however, the number of fasciolariid genera had jumped 
to three, with the Transmarian endemics Mariafusus and 
Pseudaptyxis possibly representing offshoots that now 
coexisted with their Buccinofusus ancestral stock (Pe- 
tuch, 1988b). The primitive genus Buccinofusus, in par- 
ticular, underwent rapid evolution, producing three spe- 
cies in the Calvert Formation and one species, each, in 
the Choptank and St. Mary's Formation and Little Cove 
Point Unit. LInlike the other Transmarian buccinacean 
families, all the Transmarian fasciolariid genera were 
endemic to the province, with most taxa being confined 
to the Salisbury Embayment of the Calvertian Subprov- 
ince. 

As the complete opposite of the Fasciolariidae, the 
families Buccinidae and Columbellidae were represent- 
ed in the Transmarian Province only by caloosagenic 
genera. The species within these tropically-derived 
groups, however, represented endemic Transmarian spe- 
cies radiations that paralleled those of their congeners to 
the south. The genera Celatoconus, Ptychosalpinx, and 
Solenosteira were all common and prominent compo- 
nents of the Calvert, Choptank, and St. Mary's faunas. 
In the case of Ptychosalpinx, the genus had radiated into 
an endemic Transmarian species complex with at least 



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THE NAUTILUS, Vol. 106, No. 4 




Figure. 1-14. Muricid, thaidid. and fasciolariid gastropods from the Tran.sn.arian Province. 1, 2. Patuxentrophonpatuxentenm 
(Marfn 1904), dorsal and ventral views of 18 tnm specimen. Zone 17, Choptank Formation, Drum Cliff, St. Mary s County, 
Maryland. 3. Lirosoma mariana Petuch, 1988, holotype, length 27 mm. St. Mary's Formation, St. Marys River, Maryland. 4. 
Scalasvira harasewychi Petuch, 1988, holotvpe, length 13 mm, St. Marys Formation, St Mary s River, \ ar>land 5 Ecphorosycon 
lindaiouceae new species, holotvpe, length ,55 mm (incomplete). Zone 16, Choptank Formation, Sandgates, St. Mary s County. 
Maryland UF 21466. 6. Ecphorosycon pamlico (Wilson, 1987). 68 mm specimen from Zone 10, Calvert tormation. Plum 1 omt. 



E. J. Petuch, 1993 



Page 159 



two sympatric species during St. Mary s time. The Col- 
umbellidae, which normally e.xhibits a high degree of 
species-richness in eutropical areas, was represented in 
the paratropical Transmarian Province by only a single 
genus, Mitrella. Although common in all the Maryland 
Miocene exposures, Mitrella never underwent any spe- 
cies radiation and was represented b\ only two sympatric 
species in the Calvert Formation and one species, each, 
in the Choptank and St. Mary's Formations and in the 
Little Cove Point Unit. 

Considering that the overwhelming majority of Trans- 
marian biotopes, as in the Recent Carolinian Province, 
were composed of soft-bottom sand or mud areas, the 
family Nassariidae could be expected to be present in 
the Calvertian region as a large species radiation. Al- 
though comprising only two genera, the caloosagenic 
Ilyanassa and the endemic Bulliopsis, the Nassariidae 
dominated many facies of the Calvert and St. Mary s 
Formations and were the predominant group of small 
buccinaceans in the Transmarian Province. Ilyanassa is 
present in all Maryland Miocene formations as a series 
of species swarms, with at least two species in the Calvert 
Formation, four in the Choptank, and at least three in 
the St. Mary's. This species-richness is comparable to that 
seen in the Pliocene, in such formations as the Duplin 
and Yorktown. The endemic Transmarian genus Bul- 
liopsis, on the other hand, is restricted to the Calvertian 
Subprovince and appears only at the end of Salisbury 
deposition during Little Cove Point and St. Mary's time. 

The most extensive macrobuccinacean radiation with- 
in the Transmarian Province is seen in the busyconine 
melongenids. These large and abundant predatory gas- 
tropods were represented in the Calvertian Subprovince 
by species swarms in three genera; Busycotypus (ex- 
amples shown here in figures 15-20), Sycopsis (figures 
36, 37), and Tiirrifulgur (examples shown here in figures 
21-23, 26, 27). These groups are present in all the Mary- 
land Miocene formations and exhibit only gradual mor- 
phological shifts through time. Sycopsis. the least com- 
monly encountered busyconine, was present in the 
Transmarian Province as only a single species per for- 
mation. The caloosagenic genera Tiirrifulgur and Bu- 
sycotypus, on the other hand, were often present as sym- 
patric species pairs, and in some cases, such as Busycotypus 
during St. Mary's time, was present as a complex of four 
species. The species radiations of both of these genera 
will be discussed in greater detail under the description 
of new Busycotypus and Turrifulgur species in the sys- 
tematic section at the end of this paper. 



Conacean diversity patterns: Of the three Transmarian 
conacean families, only two, the Turridae and the Tereb- 
ridae, are temporally distributed throughout the Mary- 
land Miocene and are found in all three formations. The 
third family, the Conidae, is only known from the St. 
Mary s Formation and makes its appearance in the Cal- 
vertian Subprovince only at the end of Salisburv depo- 
sition. Although an abundant and diverse family in the 
Miocene Caloosahatchian Province, the Conidae was 
represented in the Transmarian Province by only a single 
subgenus of Conus. This group, Leptoconus (figures 40, 
41), had evolved into a complex of three endemic sym- 
patric species, C. diluvianus Green, 1830, C. sanctae- 
mariae Petuch, 1988, and C. asheri Petuch, 1989, by the 
end of St. Mary's time. 

The Terebridae, although a common component of all 
Transmarian molluscan assemblages, was present as only 
two genera, the caloosagenic Strioterebrum and the en- 
demic genus Laevihastula. The absence of core tropical 
terebrids such as Myurella, Hastula. and Paraterebra, 
which are all found in the Miocene Caloosahatchian and 
Gatunian Provinces to the south, points to the marginal 
paratropical nature of the Transmarian Province, and 
the Calvertian Subprovince in particular. Strioterebrum 
shows an interesting bimodal evolutionary pattern, with 
two spurts of speciation, one during Calvert time and 
one at the end of Salisbury deposition, during St. Mary's 
time. The decline in Strioterebrum species-richness dur- 
ing Choptank time may be linked to lower water tem- 
peratures during the early Serravallian stage. L'nlike Stri- 
oterebrum. Laevihastula is absent in the Calvert 
Formation but underwent a large species radiation dur- 
ing St. Mary's time, evolving at least four sympatric 
species. 

The third Transmarian conacean family, the Turridae, 
is present in the Calvertian Subprovince as four subfam- 
ilies, twelve genera, and over thirty-five species, making 
it the single largest toxoglossate radiation in the Maryland 
Miocene. Of the twelve turrid genera, nine were endemic 
to the Transmarian Province, two were caloosagenic, and 
one was boreogenic. The most species-rich subfamily in 
the Calvertian Subprovince, the Clavinae, comprised 
three genera; the Transmarian endemics Chesaclava (fig- 
ure 42) and Sedilopsis. and the caloosagen Cymatosy- 
rinx. The largest number of species of these genera, par- 
ticularly Cymatosyrinx, is found in the St. Mary's 
Formation, and this species-richness probably reflects a 
response to the warmer marine climate during the early 
Tortonian. 



Calvert County, Marviand. 7. Buccinojusus devexus (Conrad. 1843). 60 mm specimen from Zone 10, C^aivert Formation. Plum 
Point, Calvert County, Maryland 8. Ecphorosycon kalyx (Petucli, 1988). 23 mm specimen from Zone 12, Calvert Formation, 
Scientist s Cliffs, Calvert County, Maryland 9. Ecphorosycon lindajoyccae new species, reconstructed paratvpe, length 69 mm. 
Zone 16, Choptank Formation. Sandgates, St. Mar\ 's Count), Maryland 10. Ecphorosycon pandico (Wilson, 1987), juvenile 
specimen, length 32 mm. Zone 10, Calvert Formation. Plum Point, Calvert Count), Marvland 11. Buccinofusus parilis (Conrad, 
1832), 106 mm specimen from St. Mary s Formation, St. Mary's River, Maryland. 12. Buccinofusus chesapeakensis Petuch, 1988, 
holotype, length 93 mm. Little Cove Point Unit, Little Cove Point, Calvert County, Maryland. 13, 14. Buccinofusus patuxentensis 
new species, dorsal and ventral views of holotvpe, length 76 mm. Zone 17, Choptank Formation. Drum Cliff, St. Marv's County, 
Maryland, UF 21499 



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THE NAUTILUS, Vol. 106, No. 4 




Figures 15-28. Busvconine meloiigeiml and ti.rrid gastropods from the Transmarian Province. 15, 16. Busycotypuschoptankensis 
new species, dorsal and ventral views of h.ilotNpe, length 192 mm. Zone 19, Choptank Formation, Drum C:liff, St. Mary s County, 
Maryland UF 2.'3798. 17. Rusycotypu.s ralvcrtcnsis I'etuch, 1988, dorsal view of holot>pe, 44 nun (mcomplete). Zone 10, C,aivert 
Formation, Plum Point, Calvert County. Maryland 18. Unsyrolypus clwsapcahcnsis Petuch, 1988, dorsal view of holotype length 
83 mm Little Cove Point Unit, Little Cove Point, Calvert County, Maryland i9. Biisycotypm coronatum (Conrad, 1840), dorsa 
view of 59 mm specimen, St. Mary's Formation, St. Marys River, Maryland. 20. Busycotypus rugosum (Conrad, 1843) ventra 
view of 64 mm specimen, St. Mary's Formation, St. Mary's River. 21, 26. Turrifulgur marylandicus new species, dorsal and ventral 



E. J. Petuch, 1993 



Page 161 



In the Maryland Miocene, the subtamily Mangeliinae 
was represented by the most number of genera, with the 
Transmarian endetnics Calvert urris, MariadriUia. and 
Transnmriatnrris, tlie caloosagen Ghjphustuniti. and the 
boreogen Oenopota. L'nlike the C^lavinae, the mangeliine 
lurrids were not equally distributed through time, but 
were divided into two groups: one that was confined to 
the CaKert and C^lioptank Formations and the other that 
appeared onl> during St. Mar\ s time. The former group 
contains Calverturris (figures 24, 25), Transmariat urris 
(figure 28), and Clyphostoma. ail of which disappeared 
during the mid-Serravallian. The latter group contains 
only MariadriUia, which is common in the St. Mary's 
Formation. 

The subfamily Turriculinae contained the largest- 
known Calvertian turrid species radiation, that of the 
endemic genus Mariatiirricida (figure 45). This group of 
large, fusiform conaceans had evolved over seven species 
during the Maryland Miocene, and these were probably 
the primary predators on large infaunal polychaetes. The 
last-known species, Mariatiirricida hiscatenaria (Con- 
rad, 1834), may have competed with the caloosagenic 
Leptoconus species complex of the St. Mary's Formation. 
Another endemic Transmarian turriculine group, Ches- 
asyrinx (figure 43), is rare in the Calvert and Choptank 
Formations but is abundant in the St. Mary's Formation, 
where it evolved into at least two sympatric species. This 
surge of Chesasyrinx evolution may have been in re- 
sponse to the warmer marine climate during St. Mary's 
time. Coinciding with the early Tortonian speciation acme 
of Chesasyrinx is the appearance of a new turrid group, 
Nodisurculina (figure 44), which is restricted to the St. 
Mary's Formation. 

The subfamily Turrinae, which is characterized by 
large species radiations and complexes of genera in most 
Neogene tropical provinces, was poorly represented in 
the Transmarian Province. Only a single genus, Herni- 
pleurotoma, was present, but was equally distributed 
throughout the entire Maryland Miocene and is found 
in all three formations. This endemic Transmarian genus 
is morphologically very conservative, with all known spe- 
cies closely resembling each other. Throughout its tem- 
poral range, Hemipleurotoma probably occupied the 
ecological "fringe " area, having had to compete with 
the extensive vermivorous radiations of the subfamilies 
Clavinae, Mangeliinae, and Turriculinae. I'nderscoring 
this possible competitive exclusion is the fact that onl\ 
a single Hemipleurotoma species is found in each for- 
mation, as opposed to the contemporaneous sympatric 



species swarms of such successful vermivores as ^edi- 
liopsis, Cymatosyrinx. and Mariaturricula. 

DYNAMICS OF TRANSMARIAN EXTINCTION 

Based on both the presence and absence, through time, 
of caloosagenic and endemic taxa in the Calvertian area, 
an early Serravallian extinction event can be delineated 
within the temporal sequence of the Maryland Miocene. 
This extinction interval began at the initiation of Chop- 
tank deposition, at the boundary of Planktonic Fora- 
minifer Zones NlO-Nll and N12'(Gibson, 1983:38), and 
reached a peak at the end of Choptank deposition, at 
the boundary of Planktonic Foraminifer Zones N12 and 
N13. During this time, which ranges from 13 million 
years B.P. to approximately 12.3 million years B.P. (Gib- 
son, 1983: fig. 2), several dominant gastropod groups 
disappeared from the Transmarian ecosystems, resulting 
in post-Choptank molluscan assemblages that differed in 
appearance from those of Langhian age. 

Since eutropical caloosagenic taxa such as Clyphos- 
toma, Oliva, Niso, Ficus. and Architectonica disap- 
peared at the end of Calvert time, but boreogenic taxa 
such as Oenopota and Euspira persisted throughout 
Choptank time, it can be assumed that a climatic cooling 
event was the primary culprit in the extinction of the 
dominant Langhian Transmarian gastropod taxa. As 
pointed out by Stanley (1986), a "refrigeration " event, 
such as the Serravallian cooling episode, is most probably 
the cause of both regional and mass extinctions of neritic 
faunas throughout the Phanerozoic. Since many of the 
typical Transmarian endemic genera, such as Chesatro- 
phon, Buccinofusus, Sycopsis, Chesaclava, and Maria- 
turricula. survived the Serravallian extinction and are 
present in the subsequent St. Mary's Formation, the 
Choptank-aged refrigeration event was not as severe as 
that of the mass extinction episode during the late Tor- 
tonian and Messinian Stages (the "Transmarian Extinc- 
tion" of Petuch, 1988b: 12). During that late Miocene 
time, over 95% of the Transmarian gastropod species 
lineages disappeared. The Serravallian extinction epi- 
sode, then, may have been a precursor to the catastrophic 
Transmarian Extinction of two million years later. The 
reduction of muricacean, buccinacean, and conacean 
species-richness during Choptank time is shown, graph- 
ically, in figure 47. 

Although not totally devastating to Transmarian gas- 
tropod assemblages, the Serravallian extinction was ap- 
parently severe enough to cause the extinction of tem- 



views of holotype, length (incomplete) 60 mm. Zone 10, Calvert Formation, Plum Point, Calvert C;oiinty, Maryland, I'F 21467. 
22. Turrifidfiur lurriculus Petuch, 1988, holot> pe, length 32 mm, .St. Mary's Formation. .St, Mar\ 's Hivt-r, 2.'$. Turrifidgur fusiforme 
(Conrad, 1840). (i9 mm specimen, St. Mary's Formation, St. Mary's River. 24. Calverturris hcllacrcnata (Conrad, 1841), 28 mm 
specimen. Zone 10, Calvert Formation, Plum Point, (Calvert County, Maryland 25. Calverturris schmidti new species, dorsal view 
of incomplete holotype (outline reconstructed), length 2.3 mm. Zone 17, Choptank Formation at Drum Cliff, St. Mary's County, 
Maryland, VF 21.500. 27. Turrifulgur prunicola new species, dorsal view of holotype, length 3.5 mm. Zone 10, Calvert Formation, 
Plum Point, Calvert County, Maryland, UF 21468 28. Transmariatiirris calvertensis (Martin, 1904), dorsal view of 40 mm specimen. 
Zone 10, Calvert Formation, Plum Point, Calvert County, Maryland. 



Page 162 



THE NAUTILUS, Vol. 106, No. 4 




Figures 29-45. Representative species of endemic Traiismarian gastropod genera and endemic species complexes. 29. Boreotro- 
phon lindae Petucli, 1988, length 13 mm, St Mary's Formation (Boreotrophon Fischer, 1884, Transmarian species complex). 30. 
Boreotrophon harasewychi Petuch, 1988, length 16 mm, St Mar> 's Formation {Boreotruphcm Fischer, 1884, Transmarian species 
complex) .31. St ephanusalpinx candelabra Petuch, 1988, length 18 mm, Choptank Formation, lateral view showing small labial 
tooth (Stephanosalpinx Petuch, 1988). 32. Chesatwpliun chesapeaheanus (Martin, 1904), length 11 mm, St. Marys Formation 
(Chesatrophon Petuch, 1988). 33. Mariasalpinx emilyae Petuch, 1988, length 28 mm, St Mary's Formation {Mariasalpinx Petuch, 



E. J. Petuch, 1993 



Page 163 




Figure 46. Miocene eastern North America, showing conti- 
nental configuration (stippled area) and molluscan faunal prov- 
inces, superimposed upon the outline of Recent North America 
(dotted line) C = Miocene Caloosahatchian Molluscan Prov- 
ince, T = Transmarian Province, with its three subprovinces. 
the Pungoian U), the Calvertian (2), and the Sankatian (3); 
thick fence-line represents the boundarv between the Trans- 
marian and Miocene Caloosahatchian Provinces; S = Salisbury 
Embayment; A = .Albemarle Embayment 

perature-sensitive organisms such as the Transmarian 
Nautilus species (Martin. 1904:130). the Transmarian 
archeocetian whales of the genera Priscodelphinus Cope, 
1868, Ixacanthus Cope, 1890, Cetophis Cope, 1868, and 
Delphinodon Leidy . 1869. and the porpoise genus Oryc- 



30t 



§25 

CO 

m 

3) 

2201 



w 

TJ 

m 
15 

m 



10 




C CH L S 

FORMATION 
Figure 47. Species-richness of Calvertian Transmarian muri- 
cacean, buccinacean. and conacean gastropods during the mid- 
dle Miocene, show ing drop in diversity during Choptank-Little 
Cover Point time. Dotted line = number of known murica- 
ceans; dashed line = number of know n buccinaceans; solid line 
= number of know n conaceans Calvertian formations include 
the Calvert (C), Choptank (CH), Little Cove Point Unit (L), 
and St Mary's (S). Numbers of species per superfamily are 
taken from the data listed in .Appendi.\ 2. 



terocetus Cope. 1867 (Case. 1904). These last mentioned 
cetacean genera disappeared abruptly at the end of Cal- 
vert deposition, contemporaneously with the disappear- 
ance of the caloosagenic gastropod genera Glyphostoma, 
Niso, Oliva, and Ficus. Besides the endemic cetaeans, a 
large fauna of other Transmarian vertebrates died out 
at the Langhian-Serrav allian boundary . Included are the 
giant endemic stingray Raja dux Cope. 1867. the Myl- 
iobatus gigas Cope, 1867 — M. pachydon Cope, 1867 
comple.x of rays, the giant carchariid sharks Carcharias 
collata Case. 1904. C. laevissimiis (Cope, 1867), C. mag- 



1988). 34. Chesathais lindae Petuch. 1988, length 50 mm, Choptank Formation [Chesathais Petuch, 1988 1. 35. Pseuduptyxis 
sanctaemarine Petuch, 1988, length 26 mm, St Mary s Formation [Pseudaptyxis Petuch, 1988). 36. Sycopsis lindae Petuch, 1988, 
length 65 mm, Choptank Formation ySycopsis Conrad, 1867). 37. Sycopsis tuberculatum (Conrad, 1840), length 52 mm, St. Mary's 
Formation (Sycopsis Conrad, 1867) 38. Ecphora [Trisecphora) eccentrica Petuch, 1989. length 58 mm, Calvert Formation [Tri- 
secphora Petuch, 1988, subgenus of Ecphora Conrad, 1843). 39. Mariafusus marylandicus (Martin. 1904), length 60 mm, St. Mary's 
Formation (Mariafusus Petuch, 1988). 40. Conus {Leptoconus) asheri Petuch, 1988, length 42 mm. St. Mary's Formation (Lep- 
toconus Swainson. 1840, subgenus of Conus Linnaeus. 1758. Transmarian species comple.x). 41. Conus (Leptoconnus) diluvianus 
Green, 1830, length 63 mm. St Mary s Formation {Leptoconus Swainson, 1840, subgenus of Conus Linnaeus, 1758, Transmarian 
species complex) 42. Chesaclava quarlesi Petuch, 1988, length 14 mm, St. Mary's Formation (Chesaclava Petuch, 1988). 43. 
Chesasyrinx rotijera (Conrad, 1830), length 24 mm, St Mar\ s Formation (Conrad, 1862), length 14 mm, St. Mary's Formation 
(Nodisurculina Petuch, 1988). 45. Mariaturricula biscatenaria (Conrad, 1834), length :35 mm, St. Mars s Formation (Mariaturricula 
Petuch, 1988). 



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THE NAUTILUS, Vol. 106, No. 4 



na (Cope, 1867), and C. incidens Case, 1904, and the 
giant manatee Trichcchus giganteus (DeKay, l^i42) 
(Case, 1904). It seems noteworthy that the Calvert-Chop- 
tank molluscan extinction correlates with the impressive 
mammalian and elasmobranch extinctions, indicating that 
there was a province-wide catastrophic event that af- 
fected many animal groups. 

After the initial Langhian-Serravallian boundary re- 
frigeration event, there was a gradual and further cli- 
matic deterioration during Choptank time. This gradu- 
alistic pattern is demonstrated by the sequential extinction 
of several taxa. Victims of the initial Langhian-Serra- 
vallian boundary event were the caloosagenic, eutropicai 
muricid genera Murexiella and PhijUonotus, which dis- 
appeared at the end of Calvert-Kirkwood time. These 
thermophilic genera, which are widespread in the Recent 
eutropicai Caribbean Province (Petuch, 1987), would have 
been particularly sensitive to lowered water temperature. 
Another caloosagenic muricid genus, Laevityphis, was 
extremely rare in the Choptank Formation, possibly in- 
dicating a sensitivity to cooler temperatures and a mar- 
ginal survival ability. Likewise, the ecphorine thaidid 
subgenus Trisecphora, which had previously undergone 
a large species radiation in the Calvert Formation, was 
present during Serravallian time only as a single species 
per stratigraphic unit. This drop in Trisecphora species 
richness may also have been in response to lower water 
temperatures. 

Of particular interest is the thaidid genus Ecphuro- 
sycon, which appeared to have its center of speciation 
in the warmer southern Pungoian Subprovince (Petuch, 
1989). This distinctive Transmarian endemic survived 
into the very beginning of Choptank deposition (Zone 
16) as a single Calvertian species, but became extinct 
before the deposition of the upper bed of the Drum Cliff 
Member (Zone 17). This ephemeral existence of the ther- 
mophilic genus Ecphorosijcon also points to a cooling 
event during earliest Serravallian time. The possibly more 
cool-tolerant genus Ecphora s.s., however, proliferated 
into several species complexes during Choptank time and 
this accounts for the lag in muricacean extinction seen 
in figure 47. Climatic conditions apparently deteriorated 
so greatly by the end of Serravallian time, however, that 
even less thermophilic genera such as Ecphora s.s., Ches- 
athais, and Trisecphora declined in species richness. At 
the end of (>hoptank deposition, the latter two thaidid 
genera and all but one complex of Ecphora s.s. became 
extinct, underscoring the ecological impact of the cooling 
event. In the overlying Little Cove Point Unit (late Ser- 
ravallian Stage), only one, or possibly two, species of 
Ecphora s.s. are known to exist; a feeble "ghost" of the 
large Calvert and Choptank ecphorine radiations. 

The turrid genera Calvcrturris and Transmariaturris 
and the trophonine genus Patuxcntrophon also disap- 
peared during late C;hoptaiik time, along with Ecpho- 
rosycon, Chesathais and Trisecphora. The simultaneous 
loss of these stenothermal temperate endemics could only 
have been produced by a refrigeration catastrophe. In- 
terestingly enough, the large C^alvertiaii trophonine ra- 



diation, which appears in the subprovince only at the 
end of the Serravallian refrigeration event, may have 
been a left-over of a boreal invasion during the coldest 
time, at the Choptank-Little Cove Point Unit boundary. 
This strong post-Choptank boreogenic influence, which 
includes Boreotrophon and Scalaspira, may have rep- 
resented physiological adaptations to warmer water con- 
ditions during Little Cove Point-St. Mary's time. 

SYSTEMATICS 

The type material of the following new species is de- 
posited in the invertebrate paleontology collection of the 
Florida Museum of Natural History, University of Flor- 
ida, Gainesville, Florida, and bears UF numbers. 

Order Caenogastropoda 

Superfamily Muricacea 

Family Thaididae 

Subfamily Ecphorinae Petuch, 1988 

Genus Ecphorosijcon Petuch, 1988 

Ecphorosycon lindajoyceae new species 
(figures 5, 9) 

Material examined: HOLOTYPE — length 55 mm (in- 
complete), in basal bed (Zone 16) of Choptank Forma- 
tion, along waterline of Patuxent River at Sandgates, St. 
Mary's County, Maryland, UF21466; PARATYPE— 
length 69 mm (reconstructed with plasticine clay), same 
locality as holotype, collection of author. 

Description: Shell inflated, w ith globose body whorl and 
distinctly pyriform outline; spire protracted, scalariform; 
shoulder sloping; subsutural area flattened, producing 
stepped spire whorls; body whorl ornamented v\ith 3 
rounded, thin, low, adherent cords; cord along shoulder 
of body whorl projecting upward (posteriorward), be- 
coming progressivel) more bladelike on earlier whorls; 
entire body whorl, spire whorls, and siphonal canal sculp- 
tured with spiral threads; spiral threads give entire shell 
silky texture; spire whorls ornamented w ith 2 thin cords, 
with cord along shoulder being bladelike in form; si- 
phonal canal proportionally short and broad for genus; 
umbilicus wide, flaring, well developed. 

Etymology: Named for my wife, Linda Joyce Petuch, 
who assisted me with my collecting along the St. Mary's 
River and C-hesapeake Ba\ . 

Discussion: Previously (Petuch, 1989:43) I had stated 
that the genus Ecphorosycon had become extinct at the 
end of Langhian time, in the upper beds of the Calvert 
Formation. The discovery of an Ecphorosycon species in 
the lowest bed of the Choptank Formation (Zone 16), 
however, demonstrates that the genus survived into early 
Serraxallian time. Within Zone 16, the last surviving 
Ecphorosycon, E. lindajoyceae, is rare, with only a few 
fragmentary specimens having ever been collected. The 
new species has never, to my knowledge, been collected 
in the upper C^hoptank beds (Zones 17, 18, and 19), whose 
faunas ha\e been relativelv well-studied. The absence of 



E. J. Petuch, 1993 



Page 165 



this large and distinctive ecphorine from Zone 17 indi- 
cates that Ecphowsycon became extinct in earliest Ser- 
ravalliaii time, making it an excellent index fossil for 
Zone 16. 

Ecphowsycon lindajoyceae is most similar to £. pam- 
Uco (Wilson, 1987) from Zone 10 of the Plum Point 
Member of the Calvert Formation and from the Bon- 
nerton Member of the Pungo River Formation of North 
Carolina. The new species differs from £. pamlico (figure 
6, 10) in being a much wider, much more inflated shell 
with a wider, stumpier, less elongated siphonal canal. 
The spire of £. pamlico is higher and much more pro- 
tracted than the spire of E. lindajoyceae, which is dis- 
tinctly lower and depressed. The spiral threads on the 
spire, body whorl, and siphonal canal of the new species 
are also much finer than those of the coarsely-sculptured 
£. pamlico. The new species is also quite different from 
E. kalyx (Petuch, 1988) from Zones 12 and 14 of the 
Plum Point Member of the Calvert Formation (figure 
8), and differs in being a much more inflated, globose 
shell, in having reduced, less developed cords, and in 
having a sculpture pattern of fine spiral threads. 

The preservation of shells within Zone 16 is poor, with 
almost every specimen being collected in a fragmentary 
state. The paratype of £. lindajoyceae (figure 9) was 
actually flattened by the pressure of sedimentary com- 
paction and was collected, piecemeal, as a handful of 
fragments. These were later reassembled over a template 
of plasticine clay. The holotype is the best, most com- 
plete, specimen found to date. Although fragmentary, 
the holotype exhibits enough salient characteristics to 
allow for easy separation from the older £. pamlico and 
£. kalyx. At the type locality, Sandgates on the Patuxent 
River in Zone 16 of the Choptank Formation, Ecpho- 
wsycon lindajoyceae co-occurs with a large ecphorine 
fauna, including Erp/iora (Ecphora) me ganae sand gate - 
sensis Petuch, 1989, £. (Ecphora) riheri harasewychi 
Petuch, 1989, £. (Ecphora) choptankensis vokesi Petuch, 
1989, Ecphora (Trisecphora) smithac Petuch, 1988, and 
Chesathais lindae donaldasheri Petuch, 1989. 

Family Muricidae 

Subfamily Trophoninae Cossmann, 1903 

Genus Patuxent rophon new genus 

Diagnosis: Shell vase-shaped, with sharply angled, car- 
inated shoulder and high, distinctly scalariform spire; 
body whorl wide and inflated, tapering rapidly anteriorly 
into siphonal canal; siphonal canal long, slender, straight, 
equal in length to body whorl; bod\ whorl ornamented 
with 10 large, rounded cords; smaller, secondary cords 
often present between larger, primary cords; shoulder 
cord largest; spire whorls ornamented with 5 cords; si- 
phonal canal ornamented with 15-20 large, rounded 
cords; aperture oval. 

Type species: Chrysodomus patuxentensis Martin, 1904, 
C;hoptank Formation, Serravallian Miocene of Maryland 
(figures 1, 2). 



Other species in Patuxentrophon: Patuxentrophon 
unnamed species. Zone 10 ot the Plum Point Member 
of the (Jalvert Formation. 

Etymology: A combination of "Patuxent", for the Pa- 
tuxent River of Maryland, and the muricid genus Tro- 
phon. 

Discussion: Martin (1904:184) referred this small muri- 
cid group to the archaic neptuniid genus Chrysodomus 
Swainson, 1852, primarily on the basis of a similarity in 
ribbed sculpture patterns. Patuxentrophon, however, dif- 
fers from neptuniids in being a much tinier shell with a 
proportionally much longer and much better developed 
siphonal canal. The new genus is distinctly trophonine, 
having the same general size, siphonal canal develop- 
ment, and spiral sculpture pattern as living Boreotrophon 
species such as the Oregonian B. disparilis (Dall, 1891) 
and living Trophonopsis species such as the Alaskan T. 
kamchatkanus (Dall, 1902). Patuxentrophon also shares 
many characteristics with the late Serravallian, Tortoni- 
an, and Messinian trophonine genera Lirosoma Conrad, 
1862 (figure 3), and Scalaspira Conrad, 1862 (figure 4), 
having the sculpture of the former and the shape of the 
latter. It is possible that the Langhian-early Serravallian 
Patuxentrophon represents the stem-stock from which 
the later Miocene Lirosoma and Scalaspira evolved. 

Superfamily Buccinacea 

Family Fasciolariidae 

Subfamily Fasciolariinae Gray, 1853 

Genus Buccinofusus Conrad, 1868 

Buccinofusus patuxentensis new species 

(figures 13, 14) 

Material examined: HOLOTYPE — Length 77 mm, 
from Zone 17, Drum Cliff Member of Choptank For- 
mation, at Drum Cliff, Jones Wharf, St. Mary's County. 
Maryland, Patuxent River, UF 21499; PARATYPE— 
length 54 mm, same locality as holotype, collection of 
author. 

Description: Shell elongately fusiform, with very ele- 
vated, protracted spire; subsutural area sloping, shoulder 
rounded; shoulder and middle section of body whorl 
ornamented with 10-12 low, axially-arranged riblike 
knobs; spire whorls with 8-12 riblike knobs bordering 
suture; entire shell, in turn, sculptured with extremely 
numerous fine spiral cords and threads; cords finest along 
subsutural area, becoming coarser on midbody, and 
coarsest on siphonal canal; siphonal canal well developed, 
short in proportion to length of spire. 

Etymology: Named for the Patuxent River, which bor- 
ders the type locality. 

Discussion: Buccinofusus patuxentensis occupies the 
evolutionary midpoint between the primitive B. devexus 
(Conrad, 1843) of the Calvert Formation (figure 7) and 
the more advanced B. chesapeakensis Petuch, 1988 of 
the Little Cove Point Unit (figure 12) and B. parilis 



Page 166 



THE NAUTILUS, Vol. 106, No. 4 



(Conrad, 1832) of the St. Mary's Formation (figure 11). 
Morphologically, B. patuxentensis resembles the slender 
and proportionally higher-spired B. devexus in shape, 
but shares the coarser spiral sculpture of B. chesapeak- 
ensis. Of the known Buccinofusus species, B. patuxen- 
tensis is closest to the Little Cove Point B. chesapeak- 
ensis, but differs in having finer and more numerous 
spiral cords on the body whorl and spire, and in having 
stronger and more numerous knobs on the spire whorls. 
Through time, from the Langhian to the Tortonian, 
the genus Buccinofusus underwent parallel shifts in two 
morphological characters; shell length-width ratio and 
degree of sculpture coarseness. The oldest species, B. 
devexus, is the most slender and is only faintly sculpted 
with fine spiral threads. The Choptank B. patuxentensis 
retains the slender shape of B. devexus but presages the 
heavier sculptural pattern of the later forms. The Little 
Cove Point B. chesapeakensis retains the sloping shoulder 
and fine sculpturing on the subsutural area, as seen in 
the earlier species, but has the inflated body whorl and 
coarse, widely-spaced cords around the midbody and 
siphonal canal that typify the advanced forms. This char- 
acteristic Transmarian lineage culminates in the St. Mary's 
B. parilis, which has the most inflated body whorl, lowest 
spire, and uniformly coarse sculpture over the entire 
shell. 

Family Melongenidae 

Subfamily Busyconinae Finlay and Marwick, 1937 

Genus Busycotypus Wenz, 1943 

Busycotypus choptankensis new species 
(figures 15, 16) 

Material examined: HOLOTYPE— Length 192 mm 
(incomplete), in Zone 19 of Choptank Formation, at Drum 
Cliff, Jones Wharf, St. Mary's County, Maryland, along 
Patu.xent River, UF 21498. 

Description: Shell vase-shaped, very inflated, with 
ovately-cylindrical body whorl; shoulder very broad, 
wider than length of body whorl; shoulder sharply an- 
gled, carinated, ornamented with 16-20 low, undulating 
knobs per whorl; spire low, stepped; sutural area with 
wide, deep, flat-bottomed sulcus, producing widely can- 
aliculate spire whorls; periphery of sutural sulcus bor- 
dered with large, rounded, undulating cord; body whorl 
ornamented with longitudinal growth lines, devoid of 
spiral sculpture; spire whorls ornamented with 6-8 faint 
spiral threads; siphonal canal broad, ornamented with 
10-12 low, nearly obsolete spiral cords; aperture e.\- 
tremely wide, open, flaring, with smooth, unornamented 
interior; parietal area glazed. 

Etymology: Named for the Choptank Formation, for 
which this new species is an index fossil. 

Discussion: As in the case of Buccinofusus patuxen- 
tensis. Busycotypus choptankensis fills in a gap in the 
evolutionary record of the Maryland Miocene Busyco- 
typus lineage. The new species is closest to B. chesa- 



peakensis Petuch, 1988 (figure 18) of the younger Little 
Cove Point Unit, especially in being a smooth, unorna- 
mented shell with low, undulating shoulder coronations. 
The Choptank ancestor differs from its Little Cove Point 
descendant, however, in being a much more inflated 
shell, by being proportionally thinner and more fragile, 
and in having less developed, lower coronations on the 
shoulder and spire whorls. Of the two related St. Mary's 
species, B. coronatum (Conrad, 1840) (figure 19) and B. 
rugosum (Conrad, 1843) (figure 20), B. choptankensis is 
most similar to the latter, but differs in having lower, 
less developed coronations and in lacking the heavy spiral 
corded sculpture that characterizes B. rugosum. From 
the small, ancestral B. calvertensis Petuch, 1988 (figure 
17), B. choptankensis differs in being a smooth, unsculp- 
tured shell with a much better developed, wider cana- 
liculate suture. 

Through time, the Transmarian coronated Busyco- 
typus species exhibited three morphological trends; be- 
coming proportionally less inflated, developing wider and 
deeper sutural sulci, and developing larger and more 
prominent shoulder and spire coronations. The primitive 
B. calvertensis is a very wide, inflated shell with small 
coronations and a tiny, almost nonexistent sutural canal. 
Busycotypus choptankensis retains the inflated shell shape 
of the ancestral B. calvertensis, but has large, well de- 
fined shoulder coronations, and has a very well devel- 
oped, wide sutural canal. The next youngest in the se- 
quence, B. chesapeakensis, has a wide sutural canal like 
B. choptankensis but has even larger, better defined 
shoulder coronations. Unlike the primitive species B. cal- 
vertensis and B. choptankensis, however, B. chesapeak- 
ensis has the slender, more attenuated body form of the 
advanced species. The two St. Mary's species, B. corona- 
tum and B. rugosum, both appear to have evolved from 
the Little Cove Point B. chesapeakensis, and share the 
same attenuated body form. Busycotypus coronatum re- 
tains the smooth, unsculptured bod\ whorl of 6. chop- 
tankensis and B. chesapeakensis, but has the largest, most 
pronounced shoulder coronations of the whole complex. 
The contemporaneous and sympatric B. rugosum. on the 
other hand, has a heavily sculptured, corded bod> whorl 
and large, rounded, knoblike shoulder coronations. The 
noncoronated Bustjcotypus species from the St. Mary's 
Formation, B. asheri Petuch, 1988, and B. alveatum 
(Conrad, 1863), appear to belong to a separate species 
complex and are not closely related to the coronated 
forms. 



Genus Turrifulgtir Petuch, 1988 

Turrifulgur marylandicus new species 

(figures 21, 26) 

Material examined: HOLOTYPE — Length (incom- 
plete) 60 mm, in Zone 10, Plum Point Member of Calvert 
Formation, at Plum Point, Calvert County, Maryland, 
UF21467. 

Description: Shell elongated, cylindrical in form, with 



E. J. Petuch, 1993 



Page 167 



high, protracted spire; shoulder sharply angled, low on 
body whorl, below wide, ver\ sloping subsutural area; 
shoulder ornamented with 12 evenly-spaced, small, 
sharply-pointed knobs; spire whorl ornamented with 12 
knobs per whorl; siphonal canal (missing on holotype) 
elongated and slender (extrapolated from holotype); body 
whorl sculptured with numerous fine spiral threads, which 
become nearl> obsolete around mid-bod\; sloping sub- 
sutural area sculptured with 12 large spiral threads; si- 
phonal canal sculptured with numerous large, evenly- 
spaced spiral cords; aperture proportionally narrow. 

Etymology: Named for the State of Maryland, 

Discussion: The elongate, high-spired Turrifulgur 
marijlandicus stands out as unique among the known 
Transmarian Turrifulgur species. The extremely atten- 
uated bodv form is reminiscent of T. atraktoides (Gard- 
ner, 1944) from the late Burdigalian Oak Grove For- 
mation of northern Florida, and the Langhian T. 
marylandicus appears to belong to the same species com- 
plex. The new Maryland busyconine, however, differs 
from the older Oak Grove species in being an even more- 
elongated shell with a more drawn-out body whorl, in 
having a more protracted, scalariform spire, and in hav- 
ing a much more sloping shoulder and subsutural area. 
Turrifulgur nmnjlandicus also has fewer knobs per whorl 
on the spire than does its more finely-ornamented Ca- 
loosahatchian relative. 

This new protracted Transmarian Turrifulgur is sym- 
patric with T. prunicola n.sp. (described in the next 
section) at Plum Point. Turrifulgur marylandicus differs 
from this congener in being a larger, more elongated 
shell with a much higher spire. The presence of two 
sympatric species of Turrifulgur in the Calvert For- 
mation reflects a similar pattern seen in the early Langhi- 
an Shoal River Formation of northern Florida, where 
two species, T. aldrichi (Gardner, 1944) and T. dasum 
(Gardner, 1944), also co-occur. 



Turrifulgur prunicola new species 

(figure 27) 

Material examined: HOLOTYPE— Length 35 mm, in 
Zone 10, Plum Point Member of Calvert Formation, at 
Plum Point Calvert County, Maryland, UF21468. 

Description: Shell vase-shaped, fusiform, with sharply- 
angled shoulder; spire slightly stepped, comparatively 
low and flattened for genus; subsutural area only slightly 
flattened, subplanar; shoulder of body whorl ornamented 
with 14 small, low, evenly-spaced knobs; spire whorls 
ornamented with 14 small knobs per whorl; body whorl 
pinching-in abruptly to base of siphonal canal; siphonal 
canal slender, narrow; body whorl, siphonal canal, spire 
whorls and subsutural area sculptured with numerous 
very fine spiral threads, giving shell silky appearance; 
aperture oval. 

Etymology: Named for Plum Point. Calvert County, 
Maryland, the type locality. 



Discussion: Although sympatric with Turrifulgur 
marylandicus in Zone 10, T. prunicola belongs to a sep- 
arate species lineage. The narrow T. marylandicus ap- 
pears to be a member of the T. atraktoides (Gardner, 
1944)-r. dasum (Gardner, 1944) species complex, while 
T. prunicola probably belongs to the T. aldrichi (Gard- 
ner, 1944) species complex. Both Calvert species are ca- 
loosagenic offshoots of these contemporaneous complex- 
es. Although the T. prunicola lineage survived until the 
Tortonian Stage (as T. fusiforme (Conrad, 1840) and T. 
turriculus Petuch, 1988), the T. marylandicus lineage 
died off during the Langhian Stage, and is not found in 
the upper beds of the Calvert Formation nor in the 
Choptank Formation. 

Turrifulgur prunicola is closest in general shell mor- 
phology to T. turriculus Petuch, 1988 (figure 22) from 
the Windmill Point Member of the St. Mary's Formation. 
The new species differs from its St. Mary's descendant 
in having a much lower spire, in having fewer and small- 
er knobs on the shoulder and spire whorls, and in having 
finer and more delicate spiral threaded sculpture. Tur- 
rifulgur prunicola is also similar to T. fusiforme (Con- 
rad, 1840) (figure 23) from Zone 24 of the St. Mary's 
Formation, but differs in being a smaller shell with a 
more sharply angled shoulder and proportionally lower 
spire. 

Superfamily Conacea 

Family Turridae 

Subfamily Mangeliinae Fischer, 1887 

Genus Calverturris new genus 

Diagnosis: Shells elongately fusiform, with high, pro- 
tracted spires and extended, slender siphonal canals; 
shoulders subcarinated, ornamented with nodulose cord 
below greatly sloping subsutural area; nodulose cord bor- 
ders suture on spire whorls; body whorls ornamented 
with 6-8 large, evenly-spaced spiral cords; fine spiral 
threads present between cords of some species; siphonal 
canals ornamented with numerous spiral threads; anal 
notch shallow, with greatest indentation corresponding 
to nodulose shoulder cord; protoconchs proportionally 
small, rounded, domelike, composed of one and one-half 
whorls; apertures narrow. 

Type species: Pleurotoma bellacrenata Conrad, 1841, 
Calvert Formation, Langhian Miocene of Marvland (fig- 
ure 24). 

Other species in Calverturris: Calverturris schmidti 
n.sp., Choptank Formation, Serravallian Miocene of 
Maryland, described here. 

Etymology: A composite of "Calvert", for the Calvert 
Cliffs of Maryland, and "turris", "tower". 

Discussion: This small group of Transmarian turrids is 
one of the most morphologically-distinctive of the known 
Chesapeake Miocene conacean gastropods. The promi- 
nent nodulose carina around the shoulders of Calver- 
turris species, along with the large, evenly-spaced cords 



Page 168 



THE NAUTILUS, Vol. 106, No. 4 



on the body whorls, sets this genus aside from all other 
Transmarian mangeliine taxa. In general body form and 
sculpture pattern, Calverturris is closest to Sediliopsis 
Petuch, 1988, but differs in having larger and more in- 
flated shells and in having the characteristic nodulose 
shoulder carina. 

As Calverturris is only known from the Calvert and 
Choptank Formations, the genus appears to have become 
extinct during mid-Serravallian time. 

Calverturris schniidti new species 

(figure 25) 

Material examined: HOLOTYPE — Length (fragmen- 
tary) 23 mm, in Zone 17, Drum Cliff Member of Chop- 
tank Formation, at Drum Cliff, Jones Wharf, St. Mary's 
County, Maryland, along Patuxent River, UF 21500. 

Description: Shell shape and form as outlined in genus 
description; shoulder carina with evenly-spaced, large 
nodules; single large cord present on sloping subsutural 
area, just posterior to nodulose carina; body whorl or- 
namented with 67 large spiral cords (extrapolated from 
fragmentary holotype); fine threads present between large 
spiral cords. 

Etymology: Named for Mr. Robert Schmidt of Calvert 
Countv, Maryland, who collected the holotvpe at Jones 
Wharf. 

Discussion: The Choptank Calverturris schmidti is sim- 
ilar to the Calvert C. bellacrenata, the type of the genus, 
but differs in having larger and less numerous shoulder 
knobs and in having fine spiral threads between the large 
cords on the body whorl. The steeply-sloping subsutural 
area of C. bellacrenata is also smooth and devoid of spiral 
sculpture, while that of C. schmidti is sculptured with a 
single large cord and several faint spiral threads. 

While Calverturris bellacrenata is a common species 
in Zone 10 at Plum Point, C. schmidti is a much rarer 
shell, with only a few fragmentary specimens having 
ever been collected along the Patuxent River. Appar- 
ently, the genus was already dying out by Choptank time. 

Genus Transmariaturris new genus 

Diagnosis: Shells elongately terebriform, with rounded 
shoulders and greatly protracted, elevated spires; si- 
phonal canals proportionally short, stubby; subsutural 
area of all species slightly depressed, producing faintly 
indented sutural band; last whorls with smooth, unor- 
namented shoulders; spire whorls of some species orna- 
mented with rows of low, riblike knobs along suture 
margin; spire whorls of some species smooth, unorna- 
mented; body and spire whorls sculptured with only very 
faint, delicate spiral thread, giving shells silky texture; 
siphonal canals ornamented with 10-12 thin spiral cords; 
apertures proportionalK small, oval; protoconchs un- 
known; anal notch shallow, with indentation correspond- 
ing to depressed subsutural band. 



Type species: Pleurotoma (Hemipleurotoma) calver- 
tensis Martin, 1904, Calvert Formation, Langhian Mio- 
cene of Maryland (figure 28). 

Other species in Transmariaturris: "Pleurotoma 
{Hemipleurotoma)" choptankensis Martin, 1904, Chop- 
tank Formation, Serravallian Miocene of Maryland. 

Etymology: A combination of "Transmaria" , for the 
Transmarian Province, and "turris". 

Discussion: Based on general shell shape, I had origi- 
nally placed this compact group of elongated, terebri- 
form turrids in the clavine genus Chesaclava Petuch, 
1988 (Petuch, 1988:35). Further study, however has shown 
that these two groups of turrids differ in a number of 
ways and actually belong in separate subfamilies. Trans- 
mariaturris, although convergent on the protracted spire 
torm of Chesaclava, differs from that genus in having 
larger, more inflated shells with proportionally larger 
body whorls, by having longer and better developed 
siphonal canals, in having more indented anal notches 
that produce solenozones, and in having spriral sculp- 
turing. The small, smooth Chesaclava species closely re- 
semble miniature Cymatosyrinx Dall, 1889 species and 
develop a varix-like adult lip and stromboid notch. These 
two characters are not seen in Transmariaturris. 

Although the diagnostic protoconchs were missing on 
all Transmariaturris specimens examined, I have placed 
this new genus in the subfamily Mangeliinae, primarily 
on its similarity to the genus Calverturris. Both genera 
are of the same general shape and size, and share the 
same type of anal notch and outer lip structure. Trans- 
mariaturris differs from Calverturris, however, in lack- 
ing the large, prominent spiral cords on the body whorl 
and also in lacking the distinctive nodulose shoulder ca- 
rina. The subsutural area of Calverturris is planar, while 
that of Transmariaturris is slightly depressed. 



ACKNOWLEDGEMENTS 

For help in collecting research material along the St. 
Mary's and Patuxent Rivers and along the Calvert Cliffs, 
I thank Mr. Donald F. Asher, Mechanicsville, Maryland, 
Mr. Robert Schmidt. Calvert Countv', Maryland, and my 
wife, Linda J. Petuch. 



LITERATURE CITED 

Case, E. C 1904. Systematic paleontology, Miocene, Mam- 
malia, Aves, and Reptilia. In: Clark, W. B., G. B. Shattuck, 
and W. H. Dall (eds). The Miocene deposits of Maryland. 
Maryland Geological Survey, Miocene, p. 3-93. 

Gibson, T G. 1967 Stratigraphy and paleoenvironment of 
the Phosphatic Miocene Strata of North C^arolina. Geolog- 
ical Society ol ,\nierica Bulletin 78:631-650 

Gibson, T. G. 1983 Straligra|)li\ of Miocene through I.ower 
Pleistocene Strata of the L^nited States Central .Atlantic 
Goastal I'lain In. Ray, C. E. (ed). Geology and paleon- 
tology of the l^ee Creek Mine, North Carolina I. Smith- 
sonian Contributions to Paleobiology 53:35-80. 



E. J. Petuch, 1993 



Page 169 



Martin, G. C. 1904. Systematic Paleontology, Miocene, Mol- 
lusca. Gastropoda. In: Clark, W. B., G. B. Shattuck, and 
W. H. Dall (eds ) The Miocene deposits of Maryland 
Maryland Geological Survey, Miocene, p. 131-270 

Petuch, E. J. 1987. New Caribbean molluscan faunas. The 
C'oastal Education and Research Foundation, ('harlottes- 
\ille, Virginia, 154 p 

Petuch, E. J. 1988a. New species of Ecphora and Ecphorine 
Thaidids from the Miocene of Chesapeake Bay, Maryland. 
U.S.A. Bulletin of Paleomalacology 1(1):1-16. 

Petuch, E. J. 1988b. Neogene history of tropical American 
mollusks. The Coastal Education and Research Founda- 
tion, Charlottesville, Virginia, 217 p. 

Petuch, E. J. 1988c. New gastropods from the Maryland Mio- 
cene. Bulletin of Paleomalacology l(4):69-80. 

Petuch, E J 1989 Field guide to the Ecphoras The Coastal 
Education and Research Foundation, Charlottesville, Vir- 
ginia, 140 p. 

Stanley, S. M. 1986. Anatomy of a regional mass extinction: 
Plio-Pleistocene decimation of the Western Atlantic Bi- 
valve fauna. Palaios 1:17-36. 

Ward, L. W. and B. W. Blackvvelder 1980 Stratigraphic 
re\ision of I'pper Miocene and Lower Pliocene Beds of 
the Chesapeake Group, Middle Atlantic Coastal Plain. U.S. 
Geological Survey Bulletin 1482-D 61 pp .5 pis 



Appendix i. (Inntiiiued 



T C B 



Appendix 1. Muricacean, Buccinacean, and Conacean Higher 
Taxa from the Calvertian Subprovince of the Transmarian 
Province. T = Transmarian endemic genus, C = C'aloosagenic 
genus, B = Boreogenic genus 

T C B 

Superfamily Muricacea 

Family Muricidae 

Subfamily Muricinae da Costa, 1776 
Phyllonotus Swainson, 1833 * 

SubfamiK Muricopsinae Radwin and 
D'Attilio, 1971 
Miircxiclla Clench and Farfante, 1945 * 

SubfamiK Ocenebrinae Gray, 1847 
Mariawlpinx Petuch, 1988 * 

Sephanosalpinx Petuch, 1988 * 

Vrosalpinx Stimpson, 1865 * 

Subfamily Trophoninae Cossmann, 1903 

Boreotrophon Fischer, 1884 * 

Chesafrophon Petuch, 1988 * 

Lirosoma Conrad, 1862 * 

PatuxerUropbon Petuch, 1992 * 

Scalaspira C'onrad, 1862 * 

SubfamiK Txphinae Cossmann, 1903 
Laevitijphis Cossmann, 1903 * 

Eamil) Thaididae 

Subfamily Ecphorinae Petuch, 1988 

Chesathais Petuch, 1988 * 

Ecphora Conrad, 1843 * 

Ecphorosycon Petuch, 1988 * 

Trisecphora Petuch, 1988 * 

SuperfamiK Buccinacea 

FamiK Fasciolariidae 



Subfamily Fasciolariinae Gray, 1853 

Buccinofnstis Conrad, 1868 

Marinfusus Petuch, 1988 
SubfamiK Fusininae Swainson, 1840 

Pseudaptyxis Petuch, 1988 
FamiK Melongenidae 
Subfamily Busyconinae Finlay and 
Mar wick 1937 

Busycotypus Wenz, 1943 

Sycopsis Conrad, 1867 

Turrifiilgur Petuch, 1988 
Fatnilv Buccinidae 

CU'latocontts Conrad, 1862 

Ptychosalpinx Gill, 1867 

Solenosteira Dall, 1890 
Familv Nassariidae 

Bulliopsis C^onrad, 1862 

Ilyanassa Stimpson, 1865 
Eamil) C^olutnbellidae 

Mitrella Risso, 1826 
Superfamily Conacea 
Eamil) Conidae 

Leptoconus Swainson, 1840 
Eamil) Terebridae 

Lacvihastiila Petuch, 1988 

Striotercbrum Sacco, 1891 
Family Turridae 
Subfamily Clavinae Powell, 1942 

Chcsaclava Petuch, 1988 

Cymatosyrinx Dall, 1889 

Sediliopsis Petuch, 1988 
Subfamil) Mangeliinae P'ischer, 1887 

Calvcrturm Petuch, 1992 T 

Glyphostonia Gabb, 1872 

Ocnopota Morch, 1852 

Mariadrillia Petuch, 1988 

Transmariaturris Petuch, 1992 
Subfamily Turriculinae Powell, 1942 

Chcsasyrinx Petuch, 1988 

Mariaturricula Petuch, 1988 

Nodisurculina Petuch, 1988 
Subfamil) Turrinae Swainson, 1840 

Uemipleurotorua Co.ssman, 1903 



Page 170 



THE NAUTILUS, Vol. 106, No. 4 



Appendix 2. Muricacean, Buccinacean, and Conacean Species from the Calvertian Subprovince of the Transmarian Province. C 
= Calvert Formation, Ch = Choptank Formation, L = Little Cove Point Unit, S = St. Mary s Formation 



C 



CH 



L 



S 



Muricidae 

Muricinae 

'Phi/llonotus millvillensis (Richards and Harbison, 1942) 

Muric-opsinae 
Murexiella shiluhcnsis (Heilprin, 1887) 

Ocenebrinae 

Mariasalpinx emilyae Petuch, 1988 
Mariasalpinx new species 
Stephanosalpinx candelabra Petuch, 1988 
Urosalpinx riisticus (Conrad, 1830) 

Trophoninae 

Borcotrophon hara.sewychi Petuch, 1988 
Boreotrophon laevis (Martin, 1904) 
Borcotrophon lindae Petuch, 1988 
Che.satrophon chesapeakeanus (Martin, 1904) 
Chcsatrophon new species a. 
Chesatrophon new species b. 
Lirosoma mariana Petuch, 1988 
Patuxentrophon patuxenicnsis (Martin, 1904) 
Patuxcntrophon new species 
Scalaspira harasewychi Petuch, 1988 
Scalaspira vohesae Petuch, 1988 

Typinae 
Laevityphis acuticosta (Conrad, 1830) 
Laevityphis new species a. 
Laevityphis new species b. 

Thaididae 

Ecphorinae 
Chesathais ecclesiasticus (Dall, 1915) 
Chcsathai.s lindae lindae Petuch, 1988 
Chcsathaix lindae dunaldasheri Petuch, 1989 
Chesathais lindae drumcliffensis Petuch. 1989 
Chcsathais whitfieldi Petuch, 1989 
Eephora asheri Petuch, 1988 
Eephora ealvertensis Petuch, 1988 
Eephora ealvertensis subspecies 
Eephora chesapeakensis Petuch, 1992 
Eephora ehoptankensis ehoptankensis Petuch, 1988 
Eephora ehoptankensis delieata Petuch, 1989 
Eephora choptaiikensis vokesi Petuch, 1989 
Eephora gardnerae gardnerae Wilson, 1987 
Eephora gardnerae angusticostata Petuch, 1989 
Eephora germonae Ward and Gilinsky, 1988 
Eephora mcganae meganae Ward and Gilinsky, 1988 
Eephora meganae sandgatescnsis Petuch, 1989 
Eephora rikeri riken Petuch, 1988 
Eephora rikeri harasewychi Petuch, 1988 
Eephora lurneri Petuch, 1992 
Eephora wardi Petuch, 1989 
Eephora tvardi subspecies 
Eephora leilliamsi Ward and Gilinsky, 1988 
Eephora (Trisecphora) eeeentriea Petuch, 1988 
Eephora (Triseephoru) martini I'ctuch, 1988 
Eephora (Trisecphora) patiixentia patuxentia Petuch, 1988 
Eephora (Trisec])}iora) patuxentia shaltucki Pctuth, 1989 
Eephora (Trisecphora) ])runieolu Pelucli, 1988 
Eephora (Trisecphora) scientistensis Petuch, 1992 
Eephora (Trisecphora) schmidti Petuch, 1989 
Eephora (Trisecphora) srtiithae Petuch, 1988 
Eephora (Trisecphora) smithae subspecies 



E. J. Petuch, 1993 



Page 171 



Appendix 2. Continued 



CH 



Ecphora (Trisecphora) tricostata Martin, 1904 
Ecphorosycon kahjx (Petuch, 1988) 
Ecphowsycun lindajoijcvae Petuch, 1992 
Ecphorosycon paiulicu (Wilson, 1987) 
Ecphorosycon new species 

Fasciolariidae 
Buccinofusus (?) calvertanus (Martin, 1904) 
Buccinoftisus chcsapeakensis Petuch, 1988 
Buccinofusus devexus (Conrad, 1843) 
Buccinofusus migrans (Conrad, 1843) 
Buccinofusus parilis (Conrad, 1832) 
Buccinofusus patuxentcnsis Petuch, 1992 
Muriafusus marylandicus (Martin, 1904) 
Pscudaplyxis sanclacrnuruic Petuch, 1988 

Meiorjgenidae 

Busyconinae 
Busycotypus alveatum (Conrad, 1863) 
Busycotypus asheri Petuch, 1988 
Busycotypus calvertensis Petuch, 1988 
Busycotypus chcsapeakensis Petuch, 1988 
Busycotypus choptankensis Petuch, 1992 
Busycotypus coronatum (Conrad, 1840) 
Busycotypus rugosum (Conrad, 1843) 
Sycopsis lindac Petuch, 1988 
Sycopsis tuberculatum (Conrad, 1840) 
Syncopsis new species a 
Sycopsis new species b. 
Turrifulgur fusiforme (Conrad, 1840) 
Turrifulgur marylandicus Petuch, 1992 
Turrifulgur prunicola Petuch, 1992 
Turrifulgur turriculus Petuch, 1988 
Turrifulgur new species a. 
Turrifulgur nev\ species b. 

Buccinidae 
Celatoconus asheri Petuch, 1988 
Celatoconus protractus (Conrad, 1843) 
Ptychosalpinx lienosa (Conrad, 1843) 
Ptychosalpinx lindae Petuch, 1988 
Ptychosalpinx pustulosus Petuch, 1988 
Ptychosalpinx new species 
Ptychosalpinx new species 
Solenostcira cundierlandiana (Gabb, 1860) 
Solenosteira new species 

Nassariidae 
Bulliopsis integra (Conrad, 1842) 
Bulliopsis marylandica (Conrad, 1862) 
Bulliopsis ovata (Conrad, 1862) 
Bulliopsis quadrata (Conrad, 1830) 
Bulliopsis subcylindrica (Conrad, 1862) 
Ilyanassa calvertensis (Martin, 1904) 
Ilyanassa clongata (Whitfield. 1894) 
Ilyanassa greensboroensis (Martin, 1904) 
Ilyanassa gubernatoria (Martin, 1904) 
Ilyanassa marylandica (Martin, 1904) 
Ilyanassa pcralta (Conrad, 1868) 
Ilyanassa peraltoides (Martin, 1904) 
Ilyanassa trivittatoides (Whitfield, 1894) 
Ilyanassa new species 

Columbellidae 
Mitrclla calvertensis (Martin, 1904) 
Mitrclla communis (Conrad, 1862) 



* 
* 

* 
* 
* 
* 

* * 
« * * 

* * 

* 
* 

* 

* 

* 
* 

* * 

* * 
* 

* 



Page 172 



THE NAUTILUS, Vol. 106, No. 4 



.Appendix 2. C^oiitinucd 



Cll 



Mitrella new species a. 
Milrella new species b. 
Mitrella new species c. 
('onidae 
Contis (Leptorimtis) asheri Petiicli, 1988 
Contis (Leptoconus) diluvianiis Green, 1830 
Contis (Lcptoco7itts) sanctaemariae Petucli, 1988 
Terebridae 
Laevihasliila inornata (Whitfield, 1894) 
Laevihastula marylandica Petucli, 1988 
Lacvihastiila paluxentia (Martin, 1904) 
Laevihastula simplex (Conrad, 18.30) 
Laevihastula sublirata (C^onrad, 1863) 
Strioterebrum ealvertensis (Martin, 1904) 
Strioterchrum eurvilineata (Whitfield, 1894) 
Strioterebrum eurvilirata (Conrad, 1843) 
Strioterebrum dalli (Martin, 1904) 
Strioterebrum sineera (Dall, 1895) 
Strioterebrum ichitficldi (Martin, 1904) 
Tiirridae 
(^la\ inae 
Chesaelava dissimilis (Conrad, 1830) 
Chesaclava pseudeburnea (Whitheld, 1894) 
Chesaelava quarlesi Petuch, 1988 
Chesaclava whitfieldi (Martin, 1904) 
Chesaclava new^ species 
Cymatosijrinx limatula (Clonrad, 1830) 
Cynuitosyrinx mariana Petuch, 1988 
Cymatosyrinx pyramidalis (Martin, 1904) 
Cymntosyrinx new species a, 

Cymatosyrinx new species li. 

Scdiliopsis angulata (Martin, 1904) 

Sediliupsis ealvertensis (Martin, 1904) 

Sediliopsis distans (Conrad, 1862) 

Sediliopsis gracilis (Conrad, 1830) 

Sediliopsis incilifera (Conrad, 1830) 

Sediliopsis paluxentia (Martin, 1904) 
Mangehinae 

Calverturris bellacrenata (C^onrad, 1841) 

Calverturris schmidti Petuch, 1992 

Clyphosloma obtusa (Martin, 1904) 

Oenopota cornelliana (Martin, 1904) 

Ocnopota marylandica (Petuch, 1988) 

Oenopota parva (Conrad, 18.30) 

MariadriUia parvoidea (Martin, 1904) 

Transmariaturris ealvertensis (Martin, 1904) 

Trausmariaturris choplankrnsis (Martin, 1904) 
Turricuhnae 

Chesasyrinx mariana (Martin, 1904) 

Chesasyrinx rotifera (Conrad, 18.30) 

Chesasyrinx new species 

Mariaturricula marylandica (Conrad, 1834) 

Mariaturricula biscalenaria (Conrad, 1841) 

Mariaturricula rugata (Conrad, 1862) 

Mariaturricula new species a, 

Mariaturricula new species b. 

Mariaturricula new species c. 

Mariaturricula new species d. 

Nodisurculina engonata (Conrad, 1862) 
Turrinae 

Ilemipleurotoma communis (Conrad, 1830) 



E. J. Petuch, 1993 Page 173 



Appendix 2. Continued 

c: CM 1. 

Hcmiplcurotoma protocommiinis (Martin, 1904) * 

Ilcmiplcurcitoma new species * 

' Known from the Cal\ert-equi\alent kirkwdod Formation of New Jersey; possibi\ occurring in the Calvert Formation. 



THE NAUTILUS 106(4):174-180, 1993 



Page 174 



Publication Dates of The Nautilus 



Eugene V. Coan 

Department of Invertebrate Zoology 
California Academy of Sciences 
Golden Gate Park 
San Francisco, CA 94118 USA 



M. G. Harasewych 

Department of Invertebrate Zoology 
National Museum of Natural History 
Smithsonian Institution 
Washington, DC 20560 USA 



As the publication dates of earlier issues of The Nautihis 
are nowhere readily accessible, the purpose of this note 
is to provide this information in a single comprehensive 
listing. 

Although six indices have been published for The Nau- 
tilus and for its predecessor. The Conchologists Ex- 
change (Pilsbry & Johnson, 1895; Henderson, 1927; La 
Rocque, 1951, 1963, 1972; Abbott, 1979), none provide 
a listing of publication dates. The dates listed below are 
compiled from a variety of sources. 

Dates for issues published during Henry A. Pilsbry s 
tenure as Editor (volume 3, number 1, to volume 71, 
number 3) were determined largely from his published 
bibliographies (American Malacological Union, 1940; 
Baker^ 1958; Clench & Turner, 1962), and are identified 
by an asterisk (•) following the date. As noted in the first 
of these bibliographies (American Malacological Union, 
1940;4), the dates for volumes 13-45 inclusive were de- 
rived from ledgers maintained by Charles W. Johnson, 
then Managing Editor. 

Publication dates of issues prior to volume 13, as well 
as of volume 46, number 1, had been extrapolated based 
on the dates of receipt of these issues by several libraries 
(American Malacological Union, 1940:4). These dates, 
identified by a dagger (t), are estimates and may in some 
cases prove to be one or more days later than the actual 
date of publication. 

During H. Burrington Baker's association with The 
Nautilus, first as Business Manager (volume 46, number 
2, to volume 71, number 3) then as Editor and Editor 
Emeritus (volume 71, number 4, to volume 84, number 
1), publication dates for a volume were published in the 
"Notes and News" section of the first or second issue of 
the subsequent volume. This practice was continued by 
the succeeding editor R. Tucker Abbott (volume 83, 
number 1, to volume 100, number 4) for volume 85. 
Dates determined from these listings are marked by a 
double dagger (|) following the date. 

Exact dates for volumes 86-89 have not previously 
been published. Mailing dates for these issues were de- 
termined from records on file in the editorial offices of 
the journal (PS Forms 3541 and 3542), are identified by 
a section mark (§). 

Beginning with volume 90, dates of publication were 
printed on the cover and/or title page of every issue, 
and are not included here. 

Dates derived from the bibliography of W. H. Dall 
(Bartsch et a!.. 1946), identified by a number sign (#), 



supplement and/or corroborate the dating of the earliest 
issues. For issues that could not be dated in any other 
manner, the date of the library stamp of the Library of 
Congress (identified as LC), the National Museum of 
Natural History (identified as USNM) the Museum of 
Comparative Zoology, Harvard Ihiiversity (identified as 
MCZ), or the Boston Society of Natural Historv (iden- 
tified as BSNH, now in the library of R. Tucker Abbott) 
is provided. 



The Conchologists' Exchange 

1(1); July 1886 

1(2); August 1886 

1(;3); 27 September 1886 LC 

1(4); October 1886 

1(5); 6 December 1886 t 

1(6); after 28 December 1886 f 

1(7); January 1887 [February 25 LC] 

1(8); Februarv 1887 [March 9 LC] 

1(9/10); 30 April 1887 USNM 

1(11); May 1887 

1(12); 28 June 1887 LC 

2(1); 20 July 1887 MCZ [10 August #, 12 August LC 

2(2); August 1887 [15 September LC:] 

2(;3); September 1887 [13 October LC] 

2(4); October 1887 [5 November LC, USNM] 

2(5); November 1887 [15 December LC] 

2(6): December 1887 [23 January 1888 LC] 

2(7): January 1888 [20 February LC] 

2(8): Februarv 1888 [26 March LC] 

2(9); 2 May, 1888 t [7 May LC] 

[Publicalion suspended.] 



The Nautilus 



3(1); 5 Mav 1889 f, # 
3(2): 9 June 1889 t 
3(3); 14 July 1889 f, # 
3(4): 11 August 1889 t 
3(5): 1 October 1889 t 
3(6): 18 October 1889 t 
3(7); 13 November 1889 t 
3(8); 7 Januarv 1890 f 
3(9); 11 Februarv 1890 f. # 
3(10); 12 March 1890 t 
3(11); 15 April 1890 t 
3(12): 7 May 1890 f 

4(1): 9 June 1890 t 
4(2): 27 June 1890 t 



E. V. Coan and M G. Harasewvch, 1993 



Page 175 



4(3): 6 July 1890 t 
4(4): 5 August 1890 t 
4(5): 1 October 1890 t 
4(6): 14 October 1890 f 
4(7): 7 November 1890 t 
4(8): 22 December 1890 f. # 
4(9): 11 January 1891 t 
4(10): 1 February 1891 f 
4(11): 4 March 1891 f 
4(12): 5 April 1891 t 

5(1): 19 Mav 1891 f, # 
5(2): 17 June 1891 f. # 
5(3): 10 July 1891 t, # 
5(4): 19 August 1891 t. # 
5(5): 17 September 1891 f 
5(6): 18 October 1891 t 
5(7): 18 November 1891 f 
5(8): 9 December 1891 f 
5(9): 14 Januarv 1892 f. # 
5(10): 5 Februarv 1892 t 
5(11): 25 March 1892 f. # 
5(12): 17 April 1892 f 

6(1): 15 Mav 1892 t 

6(2): 1 June 1892 f 

6(3): 6 July 1892 f 

6(4): [?] August 1892 t 

6(5): 11 September 1892 t 

6(6): 4 October 1892 t 

6(7): 8 November 1892 f 

6(8): 12 December 1892 t 

6(9): 12 Januarv 1893 t 

6(10): 10 February 1893 # [12 Februarv f] 

6(11): [?] March 1893 t 

6(12): 9? April 1893 t [10 April MCZ] 

7(1): 7 June 1893 t 
7(2): 19 June 1893 f 
7(3): 17 July 1893 f 
7(4): 15 August 1893 t 
7(5): 3 September 1893 f. # 
7(6): 13 October 1983 t 
7(7): 5 November 1893 t 
7(8): 13 December 1893 f, # 
7(9): 5 January 1894 t 
7(10): 30 Januarv 1894 f 
7(11): 1 March 1894 f 
7(12): 2 April 1894 t. # 

8(1): 2 Mav 1894 t. # 
8(2): 4 June 1894 f 
8(3): 8 July 1894 t. # 
8(4): 3 August 1894 f. # 
8(5): 7 September 1894 f 
8(6): 5 October 1894 t 
8(7): 1 November 1894 f. # 
8(8): 3 December 1894 f, # 
8(9): 2 January 1895 t 
8(10): 3 Februarv 1895 t, # 
8(11): 4 March 1895 f, # 
8(12): 1 April 1895 f 

9(1): 2 Mav 1895 t. # 
9(2): 2 June 1895 t 
9(3): 8 Julv 1895 f. # 



9(4): 7 August 1895 t 
9(5): 3 September 1895 f. # 
9(6): 1 October 1895 t 
9(7); 4 November 1895 t, # 
9(8): 5 December 1895 t 
9(9): 2 Januarv 1896 t. # 
9(10): 3 Februarv 1896 t. # 
9(11): 10 March 1896 t 
9(12): 12 April 1896 t 

10(1): 3 May 1896 t. # 
10(2): 2 June 1896 f, # 
10(3): 2 Julv 1896 t, # 
10(4): 9 August 1896 f 
10(5): 1 September 1896 t. # 
10(6): 9 October 1896 t. # 
10(7): 3 November 1896 f 
10(8): 3 December 1896 f 
10(9): 31 December 1896 t 
10(10): 2 Februarv 1897 f 
10(11): 7 March 1897 t, # 
10(12): 2 April 1897 t 

11(1): 6 Mav 1897 t, # 
11(2): 1 June 1897 t 
11(3): 29 June 1897 f. # 
11(4): 5 August 1897 f, # 
11(5): 1 September 1897 f 
11(6): 4 October 1897 f. # 
11(7): 1 November 1897 f. # 
11(8): 6 December 1897 t, # 
11(9): 3 January 1898 f. # 
11(10): 1 Februarv 1898 t 
11(11): 4 March 1898 f 
11(12): 3 April 1898 t. # 

12(1): 1 May 1898 f. # 
12(2): 9 June 1898 t 
12(3): 30 June 1898 t. # 
12(4): 4 August 1898 t, # 
12(5); 1 September 1898 t 
12(6): 3 October 1898 t 
12(7); 7 November 1898 f. # 
12(8): 2 December 1898 f 
12(9): 9 January 1899 t 
12(10); 1 February 1899 t 
12(11): 5 March 1899 t. # 
12(12): 3 April 1899 f, # 

13(1): 6 May 1899 ■ 
13(2): 3 June 1899 • 
13(3): 29 June 1899 • 
13(4): 31 Julv 1899 • 
13(5): 31 August 1899 • 
13(6); 2 October 1899 • 
13(7): 3 November 1899 •, # 
13(8); 7 December 1899 •, # 
13(9): 2 Januarv 1900 v # 
13(10): 8 Februarv 1900 • 
13(11): 1 March 1900 •, # 
13(12); 2 April 1900 • 

14(1): 2 May 1900 • 
14(2): 2 June 1900 •, # 
14(3): 2 July 1900 - 
14(4): 1 August 1900 v # 



Page 176 



THE NAUTILUS, Vol. 106, No. 4 



14(5): 4 September 1900 • 
14(6): 2 October 1900 -, # 
14(7): 3 November 1900 • 
14(8): 6 December 1900 •, # 
14(9): 30 December 1900 • 
14(10): 1 Febniarv 1901 v # 
14(11): 1 March 1901 v # 
14(12): 6 .\pril 1901 •, # 

15(1): 3 Mav 1901 •. # 
15(2): 4 June 1901 • 
15(3): 1 July 1901 • 
15(4): 30 July 1901 •, # 
15(5): 3 September 1901 •. # 
15(6): 3 October 1901 • 
15(7): 4 November 1901 > 
15(8): 7 December 1901 v # 
15(9): 8 January 1902 v # 
15(10): 5 February 1902 • 
15(11): 6 March 1902 •, # 
15(12): 5 April 1902 • 

16(1): 5 May 1902 • 
16(2): 2 June 1902 •. # 
16(3); 5 July 1902 • 
16(4): 2 August 1902 •, # 
16(5): 8 September 1902 • 
16(6): 6 October 1902 • 
16(7): 3 November 1902 •, # 
16(8): 3 December 1902 •, # 
16(9): 5 January 1903 •, # 
16(10): 10 February 1903 • 
16(11): 28 February 1903 • 
16(12): 11 April 1903 •, # 

17(1): 20 May 1903 • 
17(2): 11 June 1903 • 
17(3): 3 July 1903 • 
17(4): 12 August 1903 •, # 
17(5): 4 September 1903 •, # 
17(6): 7 October 1903 •, # 
17(7): 6 November 1903 •, # 
17(8): 7 December 1903 • 
17(9): 8 January 1904 •, # 
17(10): 6 February 1904 •, # 
17(11): 5 March 1904 •. # 
17(12): 2 April 1904 • 

18(1): 5 May 1904 v # 
18(2): 6 June 1904 • 
18(3): 8 July 1904 • 
18(4): 4 August 1904 • 
18(5): 6 September 1904 • 
18(6): 5 October 1904 • 
18(7): 7 November 1904 •, # 
18(8); 17 December 1904 • 
18(9); 12 January 1905 • 
18(10): 11 February 1905 •, * 
18(11): 6 March 1905 •, # 
18(12): 10 April 1905 •, # 

19(1): 9 May 1905 • 
19(2): 5 June 1905 •, # 
19(3): 10 July 1905 •, # 
19(4): 5 August 1905 • 
19(5): 9 September 1905 • 



19(6); 13 October 1905 • 
19(7): 4 November 1905 • 
19(8): 8 December 1905 v # 
19(9): 6 January 1906 •, # 
19(10); 8 February 1906 • 
19(11): 9 March 1906 -. # 
19(12); 5 April 1906 -. # 

20(1); 14 May 1906 • 
20(2); 11 June 1906 • 
20(3); 2 July 1906 • 
20(4): 18 August 1906 •, # 
20(5): 22 September 1906 • 
20(6); 13 October 1906 • 
20(7); 5 November 1906 • 
20(8): 10 December 1906 • 
20(9): 11 January 1907 • 
20(10); 12 February 1907 v # 
20(11); 4 March 1907 •, # 
20(12); 12 April 1907 v # 

21(1); 13 May 1907 • 

21(2); 12 June 1907 v # 

21(3); 6 July 1907 • 

21(4): 16 August 1907 • 

21(5); 18 September 1907 • 

21(6): 10 October 1907 • 

21(7): 7 November 1907 • 

21(8): 9 December 1907 •, # [issue mislabeled as vol. 22] 

21(9): 3 January 1908 •. # [issue mislabeled as vol. 22] 

21(10): 3 February 1908 • [issue mislabeled as vol. 22] 

21(11): 7 March 1908 v # 

21(12): 4 April 1908 •, # 

22(1); 9 May 1908 •, # 
22(2): 12 June 1908 • 
22(3): 16 July 1908 v # 
22(4/5); 5 September 1908 • 
22(6); 12 October 1908 • 
22(7): 14 November 1908 •. # 
22(8); 11 December 1908 •. # 
22(9); 11 January 1909 • 
22(10); 13 February 1909 • 
22(11); 11 March 1909 •, # 
22(12); 14 April 1909 •, # 

23(1): 17 Mav 1909 • 
23(2): 4 June 1909 • 
23(3): 9 July 1909 • 
23(4): 4 September 1909 • 
23(5); 2 October 1909 v # 
23(6): 13 November 1909 • 
23(7); 9 December 1909 • 
23(8); 17 January 1910 • 
23(9); 17 February 1910 • 
23(10): 8 March 1910 • 
23(11); 15 April 1910 •. # 
23(12): - never issued 

24(1); 19 Mav 1910 •, # 
24(2); 17 June 1910 • 
24(3): 6 Jul\ 1910 •, # 
24(4): 2 August 1910 •, # 
24(5): 2 September 1910 • 
24(6): 1 October 1910 • 
24(7); 14 November 1910 • 



E. V. Coan and M. G. Harasewvch, 1993 



Page 177 



24(8): 12 December 1910 •, # 
24(9); 16 January 1911 • 
24(10): 4 February 1911 •, # 
24(11): 6 March 1911 •, # 
24(12): 10 April 1911 • 

25(1): 19Mav 1911 • 
25(2): 16 June 1911 •. # 
25(3): 5 July 1911 •, # 
25(4): 18 August 1911 • 
25(5): 11 September 1911 • 
25(6): 19 October 1911 •, # 
25(7): 11 November 1911 • 
25(8): 13 December 1911 •, # 
25(9): 15 January 1912 • 
25(10): 15 February 1912 • 
25(11): 8 March 1912 v # 
25(12): 13 April 1912 • 



26(1): 11 May 1912 ' 
26(2): 1 June 1912 • 
26(3): 12 July 1912 • 
26(4): 5 August 1912 • 
26(5): 6 September 1912 
26(6): 10 October 1912 • 
26(7): 11 November 1912 • 
26(8): 12 December 1912 • 
26(9): 4 Januarv 1913 •, # 
26(10); 5 February 1913 • 
26(11); 4 March 1913 •. # 
26(12): 2 April 1913 v # 

27(1): 8Mav 1913 • 
27(2): 21 June 1913 • 
27(3); 18 July 1913 • 
27(4); 7 August 1913 • 
27(5); 9 September 1913 ' 
27(6); 9 October 1913 • 
27(7); 6 November 1913 • 
27(8); 10 December 1913 •. 
27(9); 2 Januarv 1914 •, # 
27(10): 9 February 1914 • 
27(11); 19 March 1914 -, # 
27(12); 16 April 1914 • 



# [issue mislabeled as no, 4] 



28(1): 16 Mav 1914 -. # 
28(2): 13 June 1914 •, # 
28(3); 10 July 1914 - 
28(4); 17 August 1914 • 
28(5); 22 September 1914 
28(6); 15 October 1914 v 
28(7): 20 November 1914 
28(8); 21 December 1914 • 
28(9): 20 Januarv 1915 • 
28(10); 18 February 1915 • 
28(11): 10 March 1915 •, # 
28(12): 16 April 1915 •, # 

29(1); 19 Mav 1915 • 
29(2); 3 June 1915 • 
29(3); 6 July 1915 • 
29(4): 4 August 1915 • 
29(5); 4 September 1915 •. 
29(6); 11 October 1915 • 
29(7): 10 November 1915 • 
29(8); 17 December 1915 • 



# [issue mislabeled as no. 5] 



29(9); 7 January 1916 • 
29(10): 1 February 1916 v 
29(11): 6 March 1916 • 
29(12); 8 April 1916 • 

30(1): 19 May 1916 v # 
30(2); 5 June 1916 • 
30(3): 14 July 1916 v # 
30(4): 19 August 1916 • 
30(5); 28 September 1916 ■ 
30(6); 6 November 1916 • 
30(7): 4 December 1916 • 
30(8): 30 December 1917 • 
30(9); 20 January 1917 • 
30(10); 6 March 1917 • 
30(11); 29 March 1917 • 
30(12): 7 May 1917 • 



[THE NAUTILUS changes to a quarterly publication 
schedule.] 



31(1) 
31(2) 
31(3) 


14 July 1917 •, # 
18 October 1917 • 
14 January 1918 • 




31(4) 


5 May 1918 • 




32(1) 


20 July 1918 V # 




32(2) 


22 October 1918 • 




32(3) 


17 January 1919 •, 


# 


32(4) 


5 May 1919 • 




33(1) 


16 July 1919 •, # 




33(2) 


6 November 1919 




33(3) 


22 January 1920 •, 


# 


33(4) 


20 April 1920 • 





34(1); 19 July 1920 •, # 
34(2): 6 November 1920 • 
34(3); 11 January 1921 •, # 
34(4): 5 May 1921 -, # 



35(1) 
35(2) 
35(3) 
35(4) 

36(1) 
36(2) 
36(3) 

36(4) 



26 July 1921 •, # 

5 December 1921 •. # 

23 January 1922 -, # 

24 April 1922 • 

24 July 1922 •, # [issue mislabeled as vol. 35] 
7 October 1922 •, # 
22 January 1923 • 
14 April 1923 • 



37(1): 23 July 1923 •, # 
37(2): 11 October 1923 v # 
37(3); 15 Januarv 1924 •, # 
37(4): 24 April 1924 •, # 



38(1) 


14 July 1924 •, # 


38(2); 22 October 1924 • 


38(3); 19 January 1925 •. # 


38(4); 2 May 1925 v # 


39(1); 8 July 1925 •, # 


39(2) 


31 October 1925 • 


39(3) 


11 January 1926 •, # 


39(4) 


9 April 1926 ■ 



Page 178 



THE NAUTILUS, Vol. 106, No. 4 



40(1): 12 July 1926 • 
40(2): 22 October 1926 •, # 
40(3): 1 February 1927 • 
40(4): 29 .April 1927 •, # 

41(1): 23 July 1927 • 
41(2): 27 October 1927 • 
41(3): 17 January 1928 • 
41(4): 25 .\pril 1928 • 

42(1): 14 Julv 1928 • 
42(2): 25 October 1928 • 
42(3): 15 January 1929 • 
42(4): 11 April 1929 • 

43(1): 11 July 1929 • 
43(2): 17 October 1929 • 
43(3): 15 January 1930 • 
43(4): 24 April 1930 • 

44(1): 17 July 1930 • 
44(2): 21 October 1930 • 
44(3): 27 January 1931 • 
44(4): 27 April 1931 • 

45(1): 13 July 1931 • 
45(2): 14 October 1931 • 
45(3): 9 January 1932 • 
45(4): 9 April 1932 • 

46(1): 20 July 1932 BSNH [23 July 
46(2): 22 October 1932 t 
46(3): 25 January 1933 i 
46(4): 25 April 1933 i 

47(1): 16 June 1933 t 
47(2): 1 November 1933 t 
47(3): 26 January 1934 t 
47(4): 4 May 1934 t 

48(1): 10 July 1934 t 
48(2): 15 October 1934 t 
48(3): 19 January 1935 i 
48(4): 24 April 1935 i 

49(1); 22 July 1935 t 
49(2): 8 November 1935 :j: 
49(3): 30 January 1936 t 
49(4): 1 May 1936 t 

50(1): 14 July 1936$ 
50(2): 29 October 1936 t 
50(3): 29 January 1937 i 
50(4): 4 May 1937 i 

51(1): 3 July 1937 t 
51(2): 22 October 1937 i: 
51(3): 18 January 1938 i 
51(4): 21 April 1938 i: 

52(1): 22 July 1938 t 
52(2): 28 October 1938 t 
52(3): 23 January 1939 i 
52(4): 26 April 1939 t 

53(1): 21 July 1939:): 
53(2): 20 October 1939 t 



t] 



53(3): 28 January 1940 t 
53(4): 29 April 1940 t 

54(1): 23 July 1940 t 
54(2): 2 November 1940 i 
54(3): 4 February 1941 $ 
54(4): 5 May 1941 :j: 

55(1): 11 July 1941 t 
55(2): 24 October 1941 t 
55(3): 12 January 1942 i 
55(4): 7 May 1942 t 

56(1): 23 July 1942 1: 
56(2): 14 October 1942 t 
56(3): 15 February 1943 t 
56(4): 19 April 1943 t 

57(1): 23 July 1943 :j: 
57(2): 30 October 1943 t 
57(3): 9 February 1944 i 
57(4): 15 May 1944 t 

58(1): 17 August 1944 t 
58(2): 24 November 1944 % 
58(3): 19 February 1945 t 
58(4): 20 June 1945 t 

59(1): 6 September 1945 t 
59(2): 27 December 1945 t 
59(3): 9 February 1946 t 
59(4): 27 June 1946 t 

60(1): 30 August 1946 t 
60(2): 18 December 1946 t 
60(3): 11 March 1947 t 
60(4): 2 June 1947 t 

61(1): 14 Julv 1947 t 
61(2): 18 December 1947 $ 
61(3): 2 March 1948 $ 
61(4): 24 May 1948 t 

62(1): 22 July 1948 t 
62(2): 8 December 1948 t 
62(3): 18 March 1949 i 
62(4): 8 June 1949 :t: 

63(1): 19 September 1949 t 
63(2): 1 November 1949 t 
63(3): 13 February 1950 t 
63(4): 4 April 1950 t 

64(1): 5 July 1950 t 
64(2): 27 October 1950 t 
64(3): 15 February 1951 t 
64(4): 7 May 1951 t 

65(1): 27 August 1951 t 
65(2): 9 November 1951 t 
65(3): 25 February 1952 t 
65(4): 22 May 1952 i 

66(1): 25 July 1952 t 
66(2); 17 November 1952 t 
66(3): 2 February 1953 :|: 
66(4): 8 June 1953 t 



E. V. Coaii and M. G. Harasewych. 1993 



Page 179 



67(1) 
67(2) 

67(3): 
67(4): 



24 July 1953 t 
11 November 1953 t 
18 February 1954 $ 
17 May 1954 t 



80(3): 24 January 1967 t 
80(4): 24 April 1967 :j: 



68(1), 24 July 1954 t 



68(2) 
68(3) 
68(4) 



15 November 1954 t 
11 February 1955 t 
28 April 1955 t 



69(1): 1 August 1955 t 
69(2): 5 November 1955 $ 
69(3): 11 February 1956 •, t 
69(4): 10 May 1956 •, ij: 

70(1): 13 August 1956 t 
70(2): 12 November 1956 t 
70(3): 11 February 1957 •. t 
70(4): 29 April 1957 •. i 



71(1) 
71(2) 
71(3) 
71(4) 



16 August 1957 i 
4 November 1957 
4 March 1958 ij: 
24 April 1958 i 



72(1); 21 July 1958 t 
72(2): 1 October 1958 if. 
72(3): 15 January 1959 t 
72(4): 2 April 1959 t 

73(1): 20 July 1959:1: 
73(2): 3 October 1959 t 
73(3); 25 January 1960 ^ 
73(4): 4 April 1960 t 



74(1) 
74(2) 
74(3) 
74(4) 

75(1) 
75(2) 
75(3) 
75(4) 

76(1) 
76(2) 
76(3) 

76(4) 

77(1) 
77(2): 
77(3): 

77(4): 

78(1) 
78(2) 
78(3) 

78(4) 



I July 1960 t 

5 October 1960 $ 

II Januar\ 1961 t 

6 April 1961 $ 

3 July 1961 t 

4 October 1961 ^r 
9 January 1962 i 
9 April 1962 $ 

5 July 1962 $ 

14 November 1962 t 
17 January 1963 t 

19 April 1963 t 

6 July 1963 t 

5 October 1963 i 

7 January 1964 i 
14 April 1964 t 

6 July 1964 t 

11 October 1964 t 
25 January 1965 i 

20 April 1965 t 



79(1); 9 July 1965 t 
79(2); 15 October 1965 t 
79(3); 25 January 1966 i 
79(4); 25 April 1966 t 

80(1); 6Julv 1966 t 
80(2); 11 October 1966 t 



81(1) 
81(2): 
81(3): 

81(4): 

82(1): 
82(2): 
82(3): 
82(4): 

83(1): 
83(2): 
83(3): 
83(4): 

84(1) 

84(2): 
84(3): 

84(4) 

85(1) 

85(2) 
85(3) 
85(4) 



6 July 1967 t 
16 October 1967 t 
25 January 1968 i 
18 April 1968 :|: 

24 July 1968 t- 

28 October 1968 t 
31 January 1969 i 
28 April 1969 t 

28 July 1969 t 
31 October 1969 t 
23 January 1970 i 
30 April 1970 t 

16 July 1970 t 
5 October 1970 i 

25 January 1971 i: 

26 April 1971 t 

20 July 1971 t 
4 October 1911 t 

27 January 1972 ij: 

28 April 1972 t 



86(1): 27 July 1972 § 
86(2-4): 1 December 1972 § 



[Publication changes to one volume per calendar year] 



87(1) 
87(2) 
87(3) 
87(4) 

88(1) 
88(2) 
88(3) 
88(4) 

89(1) 
89(2) 
89(3) 
89(4) 



24 January 1973 § 
27 April 1973 § 
30 July 1973 § 

4 October 1973 § 

29 January 1974 § 

30 April 1974 § 
22 Julv 1974 § 

25 October 1974 § 

31 January 1975 § 
30 April 1975 § 
18 July 1975 § 

29 October 1975 § 



[Subsequent issues have dates of publication printed on 
the title page and generally also on the cover.] 

LITERATURE CITED 

Abbott, R. T. (compiled by L, B, Hastings and M. C. Teskey). 
1979, Indexes to The Nautilus: Geographical (volumes 
1-90) and scientific names (volumes 61-90). Melbourne, 
Florida (American Malacologists) iv + 238 pp. 

American Malacological Union. 1940. Scientific contributions 
made from 1882 to 1939 b\ Henry A. Pilsbry, Sc. D. The 
American Malacological Union. 63 pp 

Baker, H B. 1958. Scientific contributions made from 1940 
to 1957 by Henry A. Pilsbry. The Nautilus 71(3);104-112. 

Bartsch. P., h' A. Rehder, and B. E. Shields. 1946. A bibli- 
ography and short biographical sketch of William Healey 
Dal] Smithsonian Miscellaneous Collections 104(15):l-96, 
Ipl. 



Page 180 



THE NAUTILUS, Vol. 106, No. 4 



Clench, V\'. J. and R D Turner. 1962 New names introduced 
by H .\ Pilsbry in the Mollusca and Crustacea -Acadenn 
of Natural Sciences of Philadelphia, Special Publication 4: 
218 pp. 

Henderson, J. B., Jr. (assisted by VI U'. Poole; edited by W 
H. Dall). 1927. Index to The Sautilus volumes 3-34, 
1889-1921. Michigan (Clapp it Walker) 434 pp 

La Rocque, A. (assisted by G. Smithe & H. W. Harry). 1951. 
Index to The Sautilus volumes 35-60, 1921-1947 Ann 
Arbor, Michigan (University of Michigan). 332 pp 



La Rocque, A. 1963. .Author index to The Nautilus volumes 
3-75 and its predecessor The Conchologists' Exchange, 
volumes 1 and 2. Columbus, Ohio, 279 pp 

La Rocque, .\. 1972 .Author index to The Saitliluf^ \olumes 
76-85. Sterkiana 47:21-38. 

Pilsbry, H A. and C. VV. Johnson 1895. Index to the "Con- 
chologists Exchange." The Nautilus 9(4):43-48; (5):57-60. 



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