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Full text of "The Nautilus"

MBL/WHOl 



THE NAUTILUS 



Volume 102 
1988 



AUTHOR INDEX 



Absalao, R. S. 
Adams, W. F 
Askew, T. A. 

AUFFENBERG, K 
AUFFENBERC, T 
BlELER, R. 

Boss, K. J. 

bouchet, p 

Collins, T. M 

D'AsARO, C. N 

DAtiilio, A 

DuShane, H. 
Emerson, VV. K. 
Gerberich, a. G. 
Gordon, M. E. 
Haimovicl M. 
Harasewych, M. G. 

Hayek. L. C 

Hershleb, R. 
Kabat, a. R. 

KiLGOUR, B V\'. 



36, 



M. 



B. 



99 Lee. H. G. 

125 Mackie. G L 

89, 92 McKiNSTRY. D M 

40. 154 McLean, J. H 

40 \1endl. W. 

1 MlKKELSEN, P 

115 Moore, D. R. 

149 Myers, B. W. 

102 o'sullivan, j. 

1.34 Pawlik. J. R. 

106 Petit, R. E. 

30 Pip, E. 

110 POLACO, O. J 

125 POMPONI, S. .\. 

123 RoMER, N. S. 

82 Sage, W E , III 

, 92 DOS Santos Cruz, R. L. 

56 Smith, DC 

56 Snyder, MA 

164 Thompson, F. G. 

73 Vermeij, G. J 



78, 154 

73 

127 

99 

129 

1 

131 

102 

47 

47 

130, 164 

65 

129 

92 

131 

36, 110 

99 

155 

54 

78 

102 



NEW TAXA PROPOSED IN VOLUME 102 (1988) 



GASTROPODA 

Perotrochus charlestonensis Askew, new species (Pleurotomariidae) 

Macrarcnc difiilala McLean. Absalao and Santos Cruz, new species (Turbinidae) 

Nerita fortidentala Wrmeij and Collins, new species (Neritidae) 

Alvania colombiana Romer and Moore, new species (Rissoidae) 

Faiartia (Murexiella) shaskyi DAttilio and Myers, new species (Muricidae) 

Metula crosnieri Bouchet, new species (Buccinidae) 

Metula africana Bouchet, new species (Buccinidae) 

Latirus martini Sn\der, new species (Fasciolariidae) 

Vasum Stephana Emerson and Sage, new species (Turbinellidae) 
Axi'lella Petit new name (Cancellariidae) 

Terebra imitatrix Auffenberg and Lee, new species (Terebridae) 

Hypselostoma holimanae Thompson and Lee, new species (Pupillidae) 

CEPHALOPODA 

Eledone gaucha Haimovici, new species (Octopodidae) 



89 
100 
102 

131 

106 

150 

151 

54 

36 

130 

154 

78 



82 



rHE NAUTILUS 



Volume 102, Number 1 
February 16, 1988 
ISSN 0028-1344 

A quarterly devoted 
to malacology. 



Marine Biological Laboratory 
LIBRARY 

FEB 2 4 1988 

Woods Hole, Mass. 



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

ASSOCIATE EDITOR 
Dr. R. Tucker Abbott 
American Maiacoiogists, Inc. 
P.O. Bo.x 2255 
Melbourne, FL 32902 

CONSULTING EDITORS 

Dr William K. Emerson 

Department of 

Living Invertebrates 

The American Museum of 

Natural Historv 

New York, NY 10024 

Mr. Samuel L. H. Fuller 
1053 Mapleton .Avenue 
Suffiekl. 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 I'niversity 
Cambridge MA 02138 



Dr. Aurele La Rocque 
Department of Geology 
The Ohio State L'niversitv 
Columbus, OH 43210 

Dr. James H. McLean 
Department of Malacology 
Los Angeles County Musetim of 
Natural History 
900 E.xposition Boulevard 
Los Angeles, CA 90007 

Dr. Arthur S. Merrill 
% Department of Mollusks 
Museum of Comparative Zoology 
Harvard University 
Cambridge, MA 02138 

Dr. Donald R. Moore 

Division of Marine Geologv 

and Geophysics 

Rosenstiel School of Marine and 

Atmospheric Science 

University of Miami 

4600 Rickenbacker Causeway 

Miami, FL 33149 

Mr Richard E Petit 

P.O Box 30 

North Myrtle Beach, SC 29582 

Dr. Edward J. Petuch 
Department of Geology 
P"lorida .Atlantic L'niversitv 
I?6ca Raton,' "Fl' 33431 

Dr. G. Alan Soleiri 
Department of Invertebrates 
Field Museurri o£ Natural Historv 
Chicago, IL 60605 

Dr. David H. Stansber\ 
Museum of Zoology 
The Ohio State Universitv 
Columbus, OH 43210 

Dr. Ruth D Turner 
Department of Mollusks 
Museum of Comparative Zoolog\ 
Harvard Universit\ 
Cambridge, MA 02138 



Dr. Geerat J. V'ermeij 
Department of Biology 
University of Mar\ land 
College Park, MD 20740 

Dr. Gilbert L. Voss 

Division of Biology and Living 

Resources 

Rosenstiel School of Marine and 

Atmospheric Science 

Universit) of Miami 

4600 Rickenbacker Causewav 

Miami, FL 33149 

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THEt^NAUTILUS 



CONTENTS 



Volume 102, Number I 

February 16, 1988 

ISSN 0028-1344 



Rudiger Bieler 
Paula \1. \likkelsen 



Aiiatoni) and reproductive biology of two western Atlantic 
species of N'itrinellidae, with a case of protandrous 
hermaphroditism in the Rissoacea 



Helen DuShane 



Geographical distribution of some Epitoniidae (Mollusca: 
Gastropoda) associated with fungiid corals 



30 



\^ illiam K. Emerson 
Salter E. Sage, HI 



A new species of Vastim (Gastropoda: Turbinellidae) from 
off Somalia 



36 



Kurt Auffenberg 
Troy Auffenberg 



Density, spatial distribution, activity patterns, and biomass 
of the land snail, Geophoriis bothropoma Moellendorff 
(Prosobranchia: Helicinidae) 



40 



Notice 



46 



Marine Biological Laboratory 
LIBRARY 

FEB 24 1988 



Woods Hole, Mass. 



THE NAUTILUS 102(l):l-29, 1988 



Page 1 



Anatomy and Reproductive Biology of Two Western Atlantic 
Species of Vitrinellidae, With a Case of Protandrous 
Hermaphroditism in the Rissoacea 



Kiidiger Bieler 

Smithsonian Marine Station 
at Link Port 

5612 Old Dixie Highway 
Ft Pierce, FL 3-1946, USA 



Paula M. Mikkel^en 

Harbor Branch Oceanographic 

Institution 

5600 Old Dixie Highway 

Ft. Pierce, FL 34946, USA 



ABSTRACT 

Two western Atlantic vitrinellids, Cyclostremiscus beauii 
(Fischer, 1857) and Circulus texanus (Moore, 1965) new com- 
bination are redescribed based on a study of live snails from 
the burrows of the stomatopod crustacean Lysiosquilla scabri- 
cauda (Lamarck, 1818). Anatoms and reproductive biology are 
emphasized, with the first recorded description of spawn and 
larval development in the famiK'. Synon\mies are given, and 
a lectotype is selected for Cy. beauii. Literature data concerning 
anatom) are reviewed for the marine near-planispiral risso- 
aceans; Vitrinellidae (including Cyclostremiscus and Circulus, 
with Circulidae as a synonym) is considered distinct from Tor- 
nidae. Unusual morphological aspects of Cy. beauii are dis- 
cussed, including external ciliation patterns, pallial tentacles 
(which are functionalK and morphologically different from 
each other), and stomach morpholog>' (with a large posterior 
chamber). Protandrous sequential hermaphroditism in Cy. beauii 
is inferred (for the first time in the Rissoacea) from morpho- 
logical evidence of sex change correlated with size. 

Key words: Cyclostremiscus; Circulus; Vitrinellidae; Torni- 
dae; Rissoacea; systematics; anatomy; hermaphroditism. 



INTRODUCTION 

Despite the great number of nominal species assigned to 
the poorK defined, cosmopolitan, marine rissoacean fam- 
ily \ itrinellidae Bush, 1897, little is known about the 
biology of its members. Their small size, mostly unknown 
habitats, poor representation in collections, and frequent 
confusion with other small-shelled members of groups 
such as Cyclostrematidae Fischer, 1885, Skeneidae Thiele, 
1929, Turbinidae Rafinesque, 1815, and Tornidae Sacco. 
1896, may account for the lack of attention paid to this 
family, and why authors who have attempted revisions 
(e.g., Trvon, 1888; Bush, 1897; Melvill, 1906; Pilsbry & 
Olsson, 1945; Pilsbry & McGinty, 1945a,b, 1946a,b, 1950; 
Laseron, 1958; Moore, 1964; Adam & Knudsen, 1969) 
relied almost entireK' on shell characters to distinguish 
ta.xa on all taxonomic levels. 

Aside from Pilsbry and McGinty 's (1945a, 1946b) 
sketches of crawling animals of several nominal genera. 



and studies on the gross morphology of Cochliolepis 
parasitica Stimpson, 1858 (by Moore, 1972) and C. 
albicerata Ponder, 1966 (by Ponder, 1966), published 
information on anatomy is available for only one species 
of this family, Circulus striatus (Philippi, 1836) from 
the eastern Atlantic. Data on Ci. striatus, extensively 
presented by Fretter (1956) and later summarized by 
Fretterand Graham (1962, 1978), were based on material 
which Fretter and Graham (1978:228) described as "the 
only live specimens . . . obtained by Fretter (1956) from 
the stomach of the starfish Astropecten . . . dredged on 
sandy bottoms 28-30 m deep in the Gulf of Gascony." 

Thus, the present concept of vitrinellid anatomv' is 
based mainly on Fretter 's description of a single species, 
Circulus striatus, the tvpe species of the name-bearing 
genus of the nominal family Circulidae (see Discussion, 
below). Discussions of phylogenetic relationships (or s\n- 
onymies) between families such as Vitrinellidae and Tor- 
nidae have demonstrated the need for anatomical and 
reproductive data for these groups (e.g., Fretter, 1956; 
Tavlor & Sohl, 1962; Moore, 1965; Golikov & Starobo- 
gatov, 1975; Boss, 1982; Graham, 1982; Ponder, in press). 

Studies on two species of western Atlantic N'itrinellidae 
are reported herein. Populations of Cyclostremiscus 
beauii (Fischer, 1857) and Circulus texanus (Moore, 1965) 
were discovered in Florida in sand-flat burrows of the 
stomatopod crustacean Lysiosquilla scabricauda (La- 
marck, 1818). These burrows are U-shaped and extend 
up to 1.5 m into the sediment, with horizontal distances 
of 6-7 m between the two openings; a pair of stomatopods 
inhabits each burrow system, maintaining it over long 
periods of time (Serene, 1954; R. B. Manning, personal 
communication). Cyclostremiscus beauii, with a shell 
diameter of 6-8 mm, is one of the largest vitrinellids, a 
fact that facilitated detailed studv of its morphology and 
anatomy and allowed a test of Fretter s hypothesis that 
some of the characters found in the much smaller (2 mm) 
Circulus striatus are size-related (Fretter, 1956:380). 

Special emphasis was placed upon characters of the 
reproductive svstem, for which Moore (1964:18) noted 
"Nothing is known of the reproduction of the famiK 



Page 2 



THE NAUTILUS, Vol. 102, No. 1 



except that the animals are dioecious, and that the male 
is provided with a penis." Data presented herein suggest 
that Cy. beauii is a protaiidrous sequential hermaphro- 
dite. Only incomplete data are available for Circulus 
texanus, as animals of this species were collected only 
twice, both times before the actual beginning of this 
study. However, since gross anatomy, spawn, and de- 
veloping eggs were observed for this species, available 
data are presented here. 



MATERIAL AND METHODS 

Samples were taken from stomatopod burrows in shal- 
low-water sand flats in the Indian River lagoon just inside 
the Ft. Pierce Inlet, St. Lucie County, eastern Florida 
(27°28.3'N, 80°17.9'W) using a stainless steel bait pump 
("yabby pump") in conjunction with sieves of 1-2 mm 
mesh. Depths at low tide ranged from less than 0.5 m 
to supratidal, wherein the water level lay several centi- 
meters below the level of the sand. 

Living snails were maintained in Bnger bowls of sea- 
water at room temperature (24 °C). Carmine and fluo- 
rescein sodium particles were used to observe ciliary 
action and currents produced by the animals. For gross 
dissections, shells were cracked and animals subsequently 
relaxed using magnesium sulfate crystals ("epsom salts"). 
Other anaesthetic chemicals (7% magnesium chloride in 
distilled water, menthol crystals) were tried but pro- 
duced little or no effect with gradual addition, or too 
strong an effect resulting in retraction. Methylene-blue/ 
basic-fuchsin and neutral red were used to better delin- 
eate tissues and organs in gross dissections. Structures 
and organs were measured following in part the outline 
given by Davis and Carney (1973: fig. 4A). Terminology 
of the nervous system is after Davis et al. (1976). 

For histological sectioning, animals were relaxed as 
above and fixed in either glutaraldehyde-formalin so- 
lution (4% formalin, 2.5% glutaraldeh\de in 0.1 M phos- 
phate buffer, pH 7.2) or 5% buffered formalin (Humason, 
1962:14). Shells were either broken and removed, or dis- 
solved in a 1% solution of ethylene diamine tetraacetic 
acid (EDTA, adjusted to pH 7.2). Specimens were 
embedded in paraffin, sectioned at 5-7 ^m and stained 
with Alcian Blue/ Periodic- Acid-Schiff (PAS), counter- 
stained with Harris' Hematoxylin/Eosin (Humason, 1962: 
125, 269, 298). Staining reactions described in the text 
refer to this method unless otherwise rioted. Photomi- 
crographs of sections were taken with a Zeiss Photomi- 
croscope-3. 

Radulae and jaws were extracted by dissolving the 
surrouiitling soft tissue in a solution of lO'^ sodium hy- 
droxide. Spermatozoa were prepared for SEM by placing 
a drop of concentrated sperm in seawater onto a coverslip 
placed in a covered petri dish containing droplets of 25% 
glutaraldeliyde, and passing the coverslip through an 
etlianol series ending in acetone, and then critical-point 
drying the sample. Whole animals were fixed, passed 
through an ethanol series, transferred to amyl acetate, 
and critical-point dried. These, together with air-dried 



shells, radulae, jaws, and opercula were coated with gold/ 
palladium, and scanned using a Zeiss Novascan-30. Fig- 
ures 1-3 were photographed using a Hitachi S-570 scan- 
ning electron microscope. Radular terminology is after 
Bandel (1984:3). 

Protoconch and teleoconch diameters were recorded 
as the greatest dimension perpendicular to the columellar 
axis. Teleoconch height was the greatest dimension par- 
allel to the columellar axis, measured from the apex to 
the base of the aperture. Umbilical diameter was the 
greatest distance between the columellar lip and the most 
prominent portion of the umbilical wall, measured in 
ventral \ iew . Teleoconch w horls were counted from the 
protoconch Il-teleoconch line to the farthest extent of 
the periphery (= the point on the outer lip used for 
greatest shell diameter). The number of protoconch whorls 
was determined b\ the method of Ta\ lor (1975:10; sum- 
marized by Jablonski & Lutz, 1980:332, fig. 4). 

Cited repositories are (* indicates location of voucher 
material): 

ANSP — Academy of Natural Sciences of Philadelphia, 
PA. 

CAS — California Academy of Sciences, San Francisco. 
*IRCZM — Indian River Coastal Zone Museum, Harbor 
Branch Oceanographic Institution (HBOI), Ft. Pierce, 
FL. 

MCZ — Museum of Comparative Zoology, Harvard Uni- 
versity, Cambridge, M.A. 

MNHN — Museum National d'Histoire Naturelle, Paris. 
*RSMAS — Rosenstiel School of Marine and Atmospheric 
Sciences, University of Miami, FL. 

UNC-IMS — Institute of Marine Sciences, University of 
North Carolina, Morehead City, NC. 
*USNM — National Museum of Natural History, Smith- 
sonian Institution, Washington, DC. 



RESULTS 
Rissoacea Gray, 1847 

= Truncateilacea Gray, 1840, "submission to be made to ICZN 
to suppress this name" (Ponder, 1985:15). 

Vitrinellidae Rush, 1897 
Cyclostremiscus Pilsbry & Olsson, 1945:266. 

Type species: VitrineUa panamensis C B. Adams, 1852 
(by original designation). 

Cyclostremiscus beauii (Fischer, 1857) 
(figures 1-52; tables 1, 2) 

Adeorbis Beauii Fischer, 1857a:173 [nomen nudum]. 

Adcorlns Reauii Fischer, 1857b:286, pi. 10, fig. 12 [Guade- 
loupe].— Bu.sh, 1897:104. 

Cyclostrema Beam [sic].— A. Adams, 1866:251, pi. 255, fig. 25 
[after Fischer, 1857b]. 

Cyclostrema bicarinata Guppy, 1866:291, pi. 17, figs. 5a, b 
["Miocene" (Lower Pliocene), Jamaica]. 



R. Bieler and P. M. Mikkelsen, 1988 



Page 3 



Skenea sulcata "Bush" Simpson, 1887:61 [nomen nudum, see 
Moore, 1964:131], 

Adeorlris Beaui.—DaW, 1889:150; 1892:345. 

Adeorbis Beam var bicarinata —DaW. 1903:1595 ["Oligo- 
cene," Jamaica]. 

"C.irculus" Incarinatus. — Woodring, 1928:439, pi. 37, figs. 10- 
12 [neotype designation]. 

"Adeorhis" heaui. — Woodring, 1928:440. 

C.irculus stirophorus M. Smith, 1937:67, pi 6, figs. 2a, b [Pho- 
cene, Florida]. 

Cyclostrema angulata. — Hertlein & Strong, 1951:110 [West 
Indies]. 

Cyclostremiscus {Ponocyclus) heaui Incarinatus. — Pilsbry, 1953: 
427, pi. 55, figs. 1-le. 

Vitrinella iSolariorbis) fceaui.— Abbott, 1954:138. 

Vitrinella beaui. — Wells et ai. 1961:267. 

Cyclostremiscus beaui. — Moore, 1964:131-135. — Morris, 1973: 
138, pi. 40, fig. 19.— Porter, 1974:143.— Emerson & Ja- 
cobson, 1976:64, pi. 18, fig. 21 —Abbott & Dance, 1982: 
58, text-fig. 

Cyclostremiscus {Ponocyclus) beaui. — Warmke & Abbott, 1961: 
60, pi. 11, fig. b.— Humfrey. 1975:76, pi. 3, figs. U-lla. 

Cyclostremiscus (Ponocyclus) beauii. — Abbott, 1974:85, text- 
fig. 786.— Rios, 1975:38, pi. 10. fig. 131; 1985:41, pi. 16, 
fig. 177.— Yokes & Yokes, 1983:15, pi. 25, figs. 4-4a. 

Material examined: Lectotype (designated herein): 10.5 
mm, MNHN unnumbered (Guadeloupe). Neotype of 
Cyclostrema bicarinata. 7.4 mm, USNM 115621 (Plio- 
cene, Jamaica); other material: 60 specimens: NORTH 
CAROLINA: 1 specimen with dried animal, BEVER- 
IDGE Sta. I (ex Astropecten). UNC-IMS. FLORIDA: Ft. 
Pierce Inlet: 10 March 1987, 1 male, 3 unsexed; 2-3 May 
1987, 1 male; 24 June 1987, 3 males; 3 August 1987, 6 
males; 31 August 1987, 7 females, 1 1 males; 1 4 September 
1987, I female; 27 September 1987, 3 females, 2 males. 
Peanut Island, Lake Worth Inlet: 11 August 1987, 1 fe- 
male, 1 male. Boynton Beach: 1 shell, ANSP 277740. 
Miami: 1 shell, EOLIS Sta. 311, USNM 449192. Fowey 
Light: 2 shells, EOLIS Sta. 187, USNM 449193; 1 shell', 
EOLIS Sta. 129, USNM 449194; 1 shell, EOLIS Sta. 142, 
USNM 449195; 1 shell, EOLIS Sta. 170, USNM 449196; 

1 shell, EOLIS Sta. 355, USNM 449198. Turtle Harbor: 

2 shells, EOLIS Sta. 61, USNM 449197. Sand Key: 1 shell, 
EOLIS Sta. 162, USNM 449199. Kev West: 1 shell, EO- 
LIS Sta. 63, USNM 449200. Drv Tortugas: 1 shell, USNM 
61114; 1 shell, USNM 271949. Cape San Bias; 1 shell, 
USBF Sta. 2402, USNM 323914. CARIBBEAN: St. Mar- 
tin; 1 shell, ANSP 20621. Jamaica: 1 shell, USNM 426872; 
1 shell, USNM 442372. SOUTH AMERICA: CoveSas, 
Colombia: 1 shell, USNM 364409. 

Description 

Teleoconch (figures 1-3): Shell large for family [usu- 
ally 2-3 whorls, 6-8 mm diameter (x = 6.9, n = 56); 
height: X = 3.8, n = 33; umbilical diameter: x = 1.5, n = 
30; largest specimen (female): diameter 11.5 mm, height 
6.2 mm, umbilicus diameter 2.4 mm, teleoconch whorls 
3%], nearly planispiral. Opaque-whitish, with 5-8 strong, 
concentric ribs on apical side, above peripheral keel. 
Widely-spaced irregular pustules between ribs (figures 



Table 1 . .Anatomical characters and character states, and hab- 
itat types of species in the vitrinellid-toriiid complex 

1) Projecting snout bilobed, lateral extensions: (a) absent; (b) 
present. 

2) Cilia along cephalic tentacles: (a) absent; (b) present 

3) Terminal stiff setae on cephalic tentacles: (a) absent; (b) 
present. 

4) Number of pallial tentacles. 

5) Pallial tentacles: (a) all finger-shaped; {b) upper finger- 
shaped and lower paddle-shaped. 

6) Upper pallial tentacle: (a) naked; (b) with motile cilia; (c) 
with stiff setae; (d) with distinct motile cilia and/or stiff 
setae. 

7) Lower pallial tentacle: (a) naked; (b) with motile cilia; (c) 
with stiff setae; (d) with distinct motile cilia and/or stiff 
setae. 

8) Gill filaments: (a) projecting from aperture in crawling 
position; (b) not projecting from aperture in crawling po- 
sition. 

9) Anterior foot margin: (a) straight or only weakly indented; 
(b) cleft. 

10) Posterior foot margin: (a) simple and rounded or weakly 
indented; (b) cleft. 

11) Operculum nucleus: (a) concentric; (b) subcentral. 

12) Number of whorls on operculum. 

13) Eyes: (a) distinctly developed; (b) lack nerve supply. 

14) Osphradium: (a) small, ciliated groove; (b) distinctly de- 
veloped, paralleling ctenidium. 

15) Penis: (a) simple, without glandular processes, recurved; 
(b) with glandular area, directed straight back; (c) with 
several, finger-like processes. 

16) Habitat: (a) under rocks; (b) under scales of annelid Poly- 
odontes lupina (Stimpson); (c) ? from stomach of starfish; 
(d) in burrows of stomatopod Lysiosquilla scabricauda (La- 
marck); (e) sandy mud bottom; (f) under large boulders 
on well-ox\genated sand\ mud. 



5-8) on first whorl, becoming more dense and regularly 
spaced on second whorl. Fields of pustules on body whorl 
intersected by smaller concentric ridges, 4-6 between 
major ribs, added first between suture and first major 
rib, subsequently between peripheral ribs. Strong cords 
forming sharp peripheral and basal keels, separated by 
wide flat area, inclined ventrally and marked by growth 
lines, spiral cords, and pustules. Base widely umbilicate, 
with 0-5 concentric ribs between basal keel and umbi- 
licus; larger specimens showing decreasing number of 
ribs with increasing size. Umbilical wall often with 2-4 
narrow ribs. Outer lip sinuous, with shallow sutural sinus. 
Microstructure (figure 4) of 3 layers: 2 thick cross-la- 
mellar layers and 1 thin homogenous outer layer thick- 
ened to form spiral ribs. 

Protoconch (figures 9-11): Diameter 0.40-0.48 mm 
(f = 0.45, n = 32). Protoconch I (prior to hatching) 
smooth, of about 1 whorl (diameter ~ 0.23 mm). Pro- 
toconch II (after hatching, before settling) of an addi- 
tional whorl, sculptured with irregular, more-or-less con- 
centric markings (figure 11). Total protoconch of 2 whorls, 
rather high-spired (spire angle = 50°). 



Page 4 



THE NAUTILUS, Vol, 102, No. 1 



Table 2. Summary of anatomical and habitat data for species in the vitrinellid-tornid complex. Characters and character states 
are listed in table 1. Sources of data listed under each species. ** = type species of genus; * = synonym of type S[)ecics of genus. 
*' = c according to VUmre. 1964:22; d according to Moore. 1964:159 W] biological data from Florida, except for Cochliolepis 
parasitica (also South Carolituil. Co. alhiccrata (-\e« Zfalaiid ' Cirrulus sirialiis (iucrnseN. English Channel 



Characters 



Species 



12 3 4 5 6 



8 9 10 11 12 13 14 15 16 



Croup 1 

**Vitrinella helicoidea C. B .\dunis, 1850 

Pilsbry & McCinty, 194.5a: pi. 2, fig. 5; 1946b: 13. 
*Vitrinella praecox Pil.sbr\ & McGinty, 1946 

Pilsbry & McGinty, 19'45a: pi. 2, fig. 4; 1946b:14. 
Teinostoma carinacallus Pilsbr\ & McGinty, 1946 

Pilsbry & McGinty, 1946b:17, pi. 2, fig. 6b. 
Teinostoma lerema Piisbrv & McGintv, 1945 

Pilsbry & McGintv, 194.5a:6, pi. 2, fig. la. 
Teinostoma parvicallum Piisbrv 6< McGint>, 1945 
Pilsbry & McGinty, 194.5a:4. pi 2, fig. 2.' 
**Pleuromalaxis halcsi (Piisbrv & McGinty, 1945) 

Pilsbry & McGinty, 1945a: 10, pi. 2, fig. 8. 
**Cochliolepis parasitica Stirnpson, 1858 

Stimpson, 1858:307ff.; Moore, 1972:100ff. 
Cochliolepis allncerata Ponder, 1966 
Ponder, 1966.38, pi 5 
*Circultts striatus (Philippi, 1836) 

Fretter, 1956:369ff ; Fretter & Graham, 1978:227£f. 
Circtilus texanus (Moore. 1965) 

This paper 
Cyclostremiscus beaitii (Fischer, 1857) 

This paper 
Cyclostremiscus pentagonus (Gabb, 1873) 
Bush, 1897:127, pi. 22, figs. 6, 12a-g [as Skenea trilix]. 

Group 2 

**Tomura hicaudata (Piisbrv & McGintv, 1946) 

Pilsbry & McGinty, 1945a: pi. 2, fig.' 9; 1946b:15. 
*Parviturhoides interruptus (C. B. .\dams, 1850) 
Moore, 1962:695ff., fig. IB; 1964:21, 156ff.; 
1972:106ff., figs. 5, 6. 

Group 3 

**Tornus subcarinatus (Montagu, 1803) 

Woodward, 1898:140ff., pi. 8, figs. 1-3, 5-7; Fretter & 
Graham. 1978:229ff; Graham, 1982:144ff. 



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External anatomy and organs of the mantle cavity (fig- 
ures 12- 18, 20-25, 27, 34-37): Living animal trans- 
lucent vellovvish-orange, with buccal mass, anteriormost 
gill tilaments, and tip of penis rose-pink; digestive gland 
brownish-orange. Base of ctenidial lamellae on osphra- 
(iial side pigmented white; central area of osphradium 
vvliitf Hanked bv brown lines on each side, resulting in 
pattern of parallel lines on left body side (figure 12). 
Long snout terminating in pair of muscular "lips," sep- 
arated by vertical slit with mouth opening. Lips sepa- 



rated Irom remainder ot snout bv strongiv ciliated groove. 
Serial sections revealed 2 narrow longitudinal bands of 
cilia, situated in grooves on each side, beginning shortly 
behind tip of snout. Two slender, De.xible, cephalic ten- 
tacles reaching appro.ximately twice length of snout when 
fullv extended. Left tentacle fitting into shallow notch 
formed bv ventral shell keel (figure 14). iMack eves on 
outer expanded bases of cephalic tentacles, each equipped 
with spherical, transparent lens. 

Ciephalic tentacles exhibiting elaborate pattern of mo- 



Figures 1-11. Cyclostremiscus heauii. specimens from Ft. Pierce Inlet, Florida (SEM) (figures 1-3, USNM 846323). 1. Shell, 
apical view (8.0 mm diameter). 2. Shell, umbilical view (5.0 mm diameter) 3. Shell, apertural view (4.1 mm diameter) 4. 
Microstructure of lateral body wall; fracture surface parallel to growing edge 5. Teleoconch sculpture, apical view. Circled numbers 
indicate location of sculptural details in figures 6-8. Stars indicate location of sutures. 6. Detail of teleoconch sculpture, first whorl. 
7. Detail of teleoconch sculpture, second whorl. 8. Detail of teleoconch sculpture, third (= body) whorl. 9. Protoconch, apical view. 
,\rro\v indicates sculptural line between protoconchs I and II. 10. Protoconch, lateral view. 11. Sculpture of protoconch I (left, 
smooth) and protoconch II (right, sculptured) Scale bars: 4, 5, 8-10 = 0.1 mm; 6, 7, 11 = 10 fim. 



R. Bieler and P. M. Mikkelsen, 1988 



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THE NAUTILUS, Vol. 102, No. 1 




R, Bieler and P. M. Mikkelsen, 1988 



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lilc cilia and stiff bristles. Ventral tip of each tentacle 
with L-shaped, heaviK -ciliated groove surrounding 
smooth area, forming tactile pad (figure 25, tp). Nu- 
merous stiff bristles (lost during fixation, therefore not 
evident in histological or SEM preparations) distal to and 
just behind pad; additional bristles sparsely distributed 
over distal quarter of tentacle. T\\ o longitudinal ciliated 
tracts, situated in grooves, extending from region of pad 
on ventral side. Near tentacle base, innermost groove 
sloping dorsally toward dorsal midline; outermost groove 
ending proximal to expanded tentacle base (figure 20). 
[•fattened base of tentacle facing snout surface covered 
b\ additional, irregular, parallel tracts of cilia (figure 20). 
Third longitudinal ciliated groove on dorsal side of ten- 
tacle extending from flattened, triangular area just below 
eye to near tentacle tip (figure 18). Ciliated grooves usu- 
alK lined b\ narrow bands of brown pigment. Cilia fulK' 
retractable into grooves. C^ilia in grooves with distinct 
movement pattern (right tentacle: dorsal row — toward 
tentacle base, inner and outer ventral rows — toward tip; 
left tentacle: opposite directions). (These movements are 
easily mistaken for direction of ciliary beating, and there- 
fore current flow. However, further microscopic obser- 
vation with the aid of carmine and fluorescein sodium 
particles revealed lateral beating of the individual cilia 
across the tentacle, away from the snout, indicating that 
the apparent ciliary movement actualK reflects the con- 
duction of nervous impulses along the tentacle.) In cross- 
section, each tentacle usually with 3 nerves, 1 larger 
blood sinus, and several smaller blood spaces in central 
area, especially around nerve cords (figure 24). 

Foot (figures 12-14) elongate (just reaching posterior 
shell margin in crawling animal), flattened, densely and 
finely ciliated, with anterolaterally recurved corners; 
broadh rounded posteriorly and slightK indented at pos- 
terior terminus. Anterior pedal mucous gland (figure 14, 
amg) present, opening at center of transverse slit across 
anterior, leading edge of foot. No posterior mucous gland 
or metapodial tentacles. Locomotion by ciliary action. 

Operculum (figures 17, 34-37) corneous, circular, mul- 
tispiral, with about 7 whorls and small internal knob, 
supported by finely ciliated opercular lobes (figure 17, 
ol) on dorsal side of foot. Lobes simple, unpigmented, 
without tentacles. Lemon-shaped muscle scar on internal 
surface of operculum (figure 35). 

Mantle edge (figures 12, 13) somewhat scalloped, cor- 
responding to spiral ribs of shell. Large monopectinate 



ctenidium, attached along entire length to inner surface 
of mantle, originating on posterior left, curving over 
dorsum, terminating just above right eye where last few 
gill filaments protrude from aperture. Numerous (< 180) 
gill filaments (figures 16, 21) flattened, elongated leaflets, 
almost finger-like when contracted, forming tapered 
blades when extended. Filaments longest in central part 
of gill, decreasing somewhat in size toward both ends. 
"Supporting rods" lacking; filaments well supplied with 
blood spaces and muscles (transverse muscle bands giving 
extended filaments ladder-like appearance; figure 16). 
Both sides of blade carrying wide band of cilia off-center, 
closer to left (osphradial) side. Along right side in same 
relative position, each filament with longitudinal row of 
small embedded crystals (figure 16, cr). Narrow rim clear, 
somewhat thinner on "crystal" side, bearing continuous 
band of cilia. Filaments draining into large efferent bran- 
chial vessel leading to heart (figure 12). Filaments hardly 
reacted to direct physical stimuli, contracting rapidly 
when either cephalic tentacle or finger-shaped pallial 
tentacle (see below) touched. 

Whitish hypobranchial gland (figures 12, 13) paral- 
leling entire length of rectum, most conspicuous poste- 
riorly. Osphradium (figures 12, 22, 23) paralleling almost 
entire left side of gill, comprised of wide central area 
and two strongK developed, heavily ciliated, lateral zones. 
Central area w ith irregular chevron-like pattern of tracts 
of shorter cilia. Osphradial ganglion (figure 12, osg) very 
conspicuous at point about 'A of total length from mantle 
edge. Mantle cavity ending immediately behind poste- 
rior end of ctenidium. 

Two pallial tentacles (figures 13, 27) arising from just 
inside right mantle edge; upper (= most dorsal) tentacle 
finger-shaped, unciliated, somewhat closer to mantle edge, 
curling into shallow sutural sinus, directed dorsally. Low- 
er tentacle paddle-shaped, ciliated at edges (except on 
narrow stalk) and also across its broad surface; narrow 
band at tentacle edge unciliated. Single stiff bristles some- 
times occurring at tip. Lower tentacle originating at point 
on mantle edge ventral and slightly more interior to 
upper finger-shaped tentacle; directed anterolaterally. 
Stimulation with forceps or needle caused immediate 
contraction of upper tentacle (followed by contraction 
of cephalic tentacles and exposed ctenidial filaments). 
Lower tentacle showed little response to touch and can- 
not markedly contract (compare tentacles in figure 27). 
Lower tentacle observed to regulate and enhance water 



Figures 12-17. Cyclostremiscus beauii. 12. Male, left side, in crawling position (shell removed). Penis reflected anteriorly, out 
of mantle cavitv . 13. Female, right side, in crawling position (shell removed). Mantle slightly reflected to show relative insertion 
points of pallial tentacles. 14. Crawling animal, ventral view. 15. Diagrammatic cross-section through mantle cavitv of male, at 
level of osphradial ganglion. 16. Gill filament. 17. Diagrammatic view of closed operculum, as seen when animal is retracted, 
showing position of opercular lobes. Scale bars: 12-14 = 1.0 mm; 15 = 0.5 mm; 16 = 1 mm. 

ag, albumen gland; amg, anterior mucous gland; br, immobile bristles; cit, ciliary tract; com, columellar muscle; cont. connective 
tissue; cr, crvstals; ct, ctenidium; dg, digestive gland; ebv, efferent branchial vessel; es, esophagus; he, heart; hg, hypobranchial 
gland; in, intestine; ki, kidney; Ipt, lower pallial tentacle; mc, mantle cavity; me, mantle edge; ol, opercular lobe; op, operculum; 
OS, osphradium; osg, osphradial ganglion; ov, ovary; pe, penis; pvd, pallial vas deferens; re, rectum; sh, shell; sn, snout; st, stomach, 
ten, cephalic tentacle; upt, upper pallial tentacle. 



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H. Bieler and P. M. Mikkelsen, 1988 



Page 9 



tlow , by paddle positioning and by strong ciliary action, 
it'spccti\el\-, in or out of right side of mantle cavity. 

Alimentary system: Mouth opening between pair of 
muscular lips into large buccal mass (figure 42; length 
1..') mm in specimen 8.5 mm shell diameter). Radula 
protrmling from short radular sac (figure 42, ras) ex- 
tending somewhat behind and to left of buccal mass. 

Paired jaws (figure 41) each crescent-shaped, appro.x- 
imately 0.5 x 0.16 mm (in specimen 7.8 mm shell di- 
ameter), composed of interlocking diamond-shaped ele- 
ments 14 /im in length. \ar\ ing slightK in shape across 
surface of jaw. 

Radula (figures 28-33) taenioglossate, with about 100 
rows (max. 138; n = 7), length == 1.3 mm, width ~ 0.2 
mm. Rachidian tooth (figure 31) wider than long (0.40 
/im w ide. n = 8), with acute posterior corners projecting 
laterally, and concave front. Main cusp narrowly-trian- 
gular, unserrated, with 4-5 flanking cusps on each side 
(number of flanking cusps var\ing within single radula, 
apparently by splitting and fusion), decreasing laterally 
in size; base with 1 strong basal denticle per side, midway 
between posterior corners and central ridge. Lateral tooth 
ifigures 32, 33) with asymmetrical cutting edge, strongly 
indented at front edge, bearing large, narrowK -trian- 
gular, unserrated main cusp and highly variable number 
of flanking cusps (3-6 inner, 7-8 outer) decreasing in 
size laterally. Base of lateral tooth with broad central 
ridge; basal platform long, blade-like. Apex of inner mar- 
ginal (figures 30, 33) with short, stout main cusp, and 
12-14 subequal, inner and outer flanking cusps. Stalk 
long, blade-like, with robust supporting ridge. Apex of 
outer marginal (figures 28, 30) with sharp, undulating 
inner margin (sometimes finely incised into closely-ad- 
hering flanking cusps), and smooth, rounded outer mar- 
gin. 

Esophagus opening widely into buccal cavity, without 
esophageal pouches. Salivary glands (figures 19, 26, 42) 
narrow, tube-like, empt\ ing into buccal mass at its lateral 
mid-point, extending posteriorly along esophagus. Posi- 
tion of salivar\ ducts relative to circumesophageal nerve 
ring highly variable: some or all passing through ring 
(n = 3; figure 49), stopping just anterior to ring (n = 4), 
or extending past without going through ring (n = 2). 
Relati\ e lengths of sali\ ary glands vary. Anterior part of 
esophagus, in cross-section (figures 26, 43), bearing 2 



muscular, longitudinal, dorsal folds, their lower exten- 
sions coiling upwards to form .semi-isolated compart- 
ments, .interior esophagus thus divided into strongly cil- 
iated dorsal food channel (figure 26, dfc), larger ventral 
channel, and two small lateral pockets. Posterior to nerve 
ring (figure 42), ventral projections into main channel 
increasing gradualK in size, but without clear distinction 
between anterior and mid-esophageal regions. Food 
channel remaining dorsal. Dorsal folds and ventral pro- 
jections gradually decreasing in size, with posterior 
esophagus as a simple muscular tube. 

Stomach (figures 12, 38-40) amber in color, slightly 
translucent in li\'ing animal, approximately 4 x 1.5 mm 
(in largest specimen, 11.5 mm shell diameter), lying on 
exterior surface of visceral coil, appressed to the surface 
of digestive gland just posterior to heart and kidney, 
encompassing approximately Vi total length of digestive 
gland. Stomach consisting of two continuous chambers 
(figure 39) differing in function: anterior chamber ('/a of 
total stomach length) containing gastric shield, working 
end of cr\stalline style, and all openings into stomach; 
remaining -/3 forming large sorting and storage chamber. 
Esophagus entering stomach on left side at junction of 
anterior and posterior chambers. From this point, series 
of folds extending transversely across stomach, poste- 
riori) into posterior chamber, and anteriorU toward in- 
testinal opening. Opening to digestive gland lying to 
right of esophageal opening, between it and gastric shield. 
Gastric shield (figures 39, 40), with cup-like lateral wing 
upon which crystalline style rotates, protruding into an- 
terior chamber, and central longitudinal portion with 
flattened lateral expansion that cradles style, positioning 
it against cup-like grinding surface. 

Style sac and intestine, at anterior end of stomach, 
usualK partially obscured b\ connective tissue and kid- 
ney. Style sac (figure 39, ss) narrow, finger-shaped, ap- 
proximately '/3 length of stomach, not communicating 
directly with intestine [Johansson's (1940:1) group 3, re- 
vised after Mackintosh (1925)]. Style completely trans- 
parent, rod-shaped (length 1.8 mm, diameter 0.35 mm), 
rotating within style sac by action of denseK packed cilia 
on style sac walls, protruding into anterior chamber of 
stomach through flesh\ tube-like structure above gastric 
shield. Channel extending betw een digestive gland open- 
ing and intestinal opening at anterior terminus of anterior 
chamber. 



Figures 18-27. Cyclostremiscus beauii, specimens from Ft. Pierce Inlet. Florida (light micrographs or critical-point dried SEM 
preparations). 18. Head-foot (male), with mantle edge reflected posteriorly, dorsal view (SEM). 19. Sagittal section of male through 
buccal mass and esophageal region. 20. .\nterior view of snout and left cephalic tentacle, showing pattern of ciliated tracts on 
ventral surface (SEM). 21. Tip of gill filament ^SEMl. 22. Osphradium, anterior to osphradial ganglion (SEM). 23. Osphradium, 
cross-section through osphradial ganglion 24. Cephalic tentacle, cross-section. 25. X'entral tip of cephalic tentacle, showing tactile 
pad (SEM). 26. Esophagus, cross-section through anterior section, with salivary glands. 27. Pallial tentacles: upper, finger-shaped 
tentacle (retracted), and lower, paddle-shaped tentacle (SEM). Scale bars; 18 = 0.5 mm; 22 = 50 ^m; 19, 20, 23, 24, 26, 27 = 0.1 
mm; 21, 25 = 20 ^m. 

bm, buccal mass; bs, blood space; car, cartilage; cit. ciliar\ tract; cont. connective tissue; df, dorsal folds of esophagus; dfc, dorsal 
food channel of esophagus; es, esophagus; leg, left cerebral ganglion; Ipt, lower pallial tentacle; me, mantle edge; ne, nerve; os, 
osphradium; osg. osphradial ganglion; pe, penis; ra, radula; rs, receptaculum seminis; sag, salivary gland; sn, snout; tp, tactile pad; 
upt, upper pallial tentacle. 



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THE NAUTILUS, Vol. 102, No. 1 




R. Bieler and P. M. Mikkelsen, 1988 



Page 11 



Posterior chamber (figures 39, 47, pch) with irregular 
longitudinal folds along its left side leading from esoph- 
ageal opening, and well-defined longitudinal groove on 
its right side leading toward gastric shield. Central area 
\\ ith series of ciliated transverse folds. 

Proximal portion of intestine (figure 39, in) consisting 
of 3 histologicalK separate sections: (1) slightly bulging 
section immediately adjacent to stomach, containing 
large typlilosole, leading into second section via small 
loop; (2) very muscular, ball-shaped section passing un- 
der style sac to join third section; and (3) moderately 
ridged intestine proper, initially quite narrow, dilating 
slightly as it passes, without further loops or undulations, 
toward rectum in mantle cavity. Anus set back from 
mantle edge. No special ciliated tracts from anus to man- 
tle edge and exterior. 

Observed flow of particles {figure 39, smalt arrows): 
Large and small food particles enter stomach through 
esophagus. Counter-clockwise whirlpool at esophageal 
opening preliminariK' sorts particles according to size. 
Smaller particles pass laterally to right in groove toward 
gastric shield and style. Large particles move into pos- 
terior chamber along left series of longitudinal folds, 
passing to far posterior terminus of stomach. From there, 
particles pass into central area where peristaltic action 
manipulates and returns large particles to anterior cham- 
ber. ConcurrentK . smaller particles separate and/or break 
oft, and follow transverse folds toward right longitudinal 
groove, and then anteriorly within groove toward gastric 
shield. Large particles in central area of sorting chamber 
continue moving anteriorly to region of esophageal open- 
ing, where they pass rapidly by ciliary action directly to 
intestinal opening. Small particles entering area of gastric 
shield are manipulated by clockwise-rotating style against 
cup-shaped flange of gastric shield. Resultant particles 
move directly left into opening of digestive gland, guided 
by cuticularized lateral folds near style sac opening. Un- 
acceptable particles and material returning from diges- 
tive gland are shunted via longitudinal groove toward 
intestinal opening. Ball-shaped proximal section of in- 
testine probably serves as a pellet compressor. 

Live specimens fed on single-celled algae and detritus 
scraped from laboratory- aquaria walls. Fecal pellets oval, 
0.33 X 0.19 mm (n = 5), round in cross-section, with 
rounded ends. 

Renopericardial system: Two-chambered heart and 
surrounding kidney visible on left surface of visceral coil, 
posterior to ctenidium and hypobranchial gland (figure 
12). Kidney large, with nephridial gland on its outer wall. 



Kidney opening at posteriormost end of mantle cavity, 
without conspicuous ciliated tract associated with open- 
ing. No gonopericardial or renogonadial ducts observed. 

Nervous system: (iircumesophageal ganglia (figures 42, 
43, 49) moderately concentrated. RPG ratio [Davis et 
ai, 1976:263; defined as length of pleurosupraesophageal 
connective/(length of connective + length of right pleu- 
ral ganglion + length of supraesophageal ganglion)] av- 
eraged 0.49 (n = 7). Cerebral ganglia connected by nar- 
row commissure, each separated from pleural ganglia by 
constriction. Tentacular nerves with distinct swellings at 
their bases. Pedal ganglia each with paired connectives, 
connecting anteriorly to cerebral ganglia, posteriorly to 
cerebropleural junctions. Pedal commissure very short, 
narrow. Subesophageal ganglion somewhat smaller than 
supraesophageal ganglion; connective to its pleural gan- 
glion much shorter. Statocysts (figure 44, stc) about 110 
)xm diameter. Buccal ganglia small, conspicuous, joined 
by commissure passing beneath esophagus at posterior 
end of buccal mass. 

Highly-vacuoled connective tissue surrounding nerve 
ring, as well as other organs and areas throughout body. 

Reproductive system: .Animals in male phase (for dis- 
cussion of sex change, see below) distinguished from 
functional females by smaller size and by dark-orange, 
rather than creamy-orange or beige, gonadial coloration. 
Penis large (size at rest: length 2.5-3.0 mm, width at 
midlength 0.4 mm), muscular, somewhat flattened (fig- 
ures 12, 18, 42, 44) arising just behind and slightly right 
of bases of cephalic tentacles, coiling counter-clockwise 
back into mantle cavity. Fully extended penis may be 
longer than cephalic tentacles {e.g., 4.4 mm vs. 3.0 mm). 
Subcentral penial duct terminating in opening on slightly 
hooded tip, which, unlike remainder of penis, is uncil- 
iated. Long, closed pallial vas deferens extending from 
penis, along right side of body, to prostate in posterior 
part of mantle cavity (figure 44). Vas deferens relatively 
wide (diameter ~ 0.2 mm), tubular, forming prominent, 
somewhat undulating ridge that differs from surrounding 
tissue by its shiny, unciliated surface and by its white 
appearance caused by heavy internal ciliary action. Pros- 
tate orange, egg-shaped, 0.65-0.90 x 0.35-0.48 mm (n = 
5), lying ventral to rectum, connected to right pallial 
vvall, its lumen communicating with mantle cavity by 
slit (~ 0.35 mm length) at its base (figures 44, 48). Vis- 
ceral vas deferens much narrower, passing through pos- 
terior mantle wall, leading along inner coil of visceral 
mass, where widened, extensively-coiled portion func- 
tions as vesicular seminalis (figure 47, vs) before reaching 



Figures 28-37. Cyclostremiscus beauii. radula and operculum, specimens from Ft. Pierce Inlet, Florida (SEM). 28. Outer marginal 
teeth. 29. Radula, whole mount 30. Inner marginal teeth. 31. Rachidian teeth. 32. Lateral teeth. 33. Tips of lateral (left) and 
inner marginal (right) teeth 34. Operculum, outer surface (2 1 mm diameter). 35. Operculum, inner surface (1.7 mm diameter). 
36. Operculum, oblique lateral view of inner side (2 2 mm diameter). 37. Opercular peg, oblique lateral view. Scale bars: 28, 30- 

33 = 10 Mm: 29, .37 = .50 fim. 



rac, rachidian teeth. 



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THE NAUTILUS, Vol. 102, No. 1 




r\ipig ^^®9 
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pispec es 



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R. Bieler and P. M. Mikkelsen, 1988 



Page 13 



testis. Testis simple, elongated sac, yellow to bright or- 
ange, along inner right side of visceral coil, totalling 50- 
60% of its length (figure 44). 

Spermatozoon (56-62 nm total length, n = 3) with 
slightly twisted head comprising elongated nucleus and 
pointed acrosome (~ 14% of total length), very long, 
narrow midpiece (« 58%) and long tail (* 28%). Atyp- 
ical sperm not found. 

Female reproductive tract (figures 13, 46) adjacent to 
rectum along right side of mantle cavity. Female open- 
ing, about level with anus, situated on muscular papilla 
hanging freel\ in mantle ca\it\ at distal end of capsule 
gland. From there, closed sperm duct leading posteriorly, 
initially forming very thick-walled muscular vagina (fig- 
ure 52), with lumen almost filled with large gland cells. 
Sperm duct giving rise to small, ball-shaped, dorsally- 
situated sperm pouch w ith weakK muscular walls. Thin- 
walled ducts leading from sperm pouch and muscular 
vagina joining shortly before opening into slit-like lumen 
of massive capsule gland. \'agina-t>pe muscular wall 
gradualK disappearing, while thin-walled part forms 
sperm channel ("sperm groo\e " in "ventral channel of 
authors), in communication with gland but partially sep- 
arated by lateral fold (figure 46, cross-section B, If). An- 
terior part of capsule gland w ith large, turquoise-staining 
gland cells; remaining capsule gland staining dark blue 
in sections. [M least in the anterior part, the gland cells 
are arranged in a comple.\ pattern (see Ponder, in press). 
A detailed histological description was hampered by the 
extremely strong staining reaction of the glands.] Pos- 
teriorly, glandular mass continuing, with communicating 
lumina, as albumen gland, .\lbumen gland massive, thick- 
walled, with narrow, slit-like lumen, pinkish-orange in 
living animals, staining turquoise in sections; folded as 
S-shaped loop on right side of animal, partK in parallel 
with posterior part of capsule gland. Posteriormost part 
of albumen gland pressing against, or, in large females, 
extending beyond posterior mantle wall. Sperm channel 
separating at junction of capsule and albumen glands, 
forming closed o\iduct. penetrating posterior mantle wall, 
and forming large, widened, non-glandular coil, con- 
taining (from sections and dissected specimens) both 
sperm and eggs. At its posterior end, coiled oviduct giving 
rise to 2 proximal sperm pouches, as inconspicuous, sub- 
equal, ball-shaped sacs, closeK' adjacent, and partialK' 
hidden under la\ers of connective tissue and kidnev. 



In ripe females, unoriented sperm, oltcn in large quan- 
tities, found throughout length of sperm channel and in 
anterior sperm pouch (functional bursa copulatrix). 
Packed oriented sperm, with heads embedded in walls, 
found in the two posterior pouches (receptacula seminis). 

Visceral oviduct very thin-walled, leading from coiled 
oviduct to ovary. Ovar\ situated at right side of visceral 
mass, structure not grape-like, extending over central 
80% of digestive gland, covering half to all ot right side 
of coil (figure 46). 

(During copulation, sperm are apparently deposited 
in the \agina, temporariK stored in the bursa copulatrix 
and then stored in the two posterior receptacula. Fertil- 
ization occurs in the anterior part of the coiled oviduct, 
after which fertilized eggs pass through the ciliated lu- 
mina of the albumen and capsule glands where they are 
surrounded by capsular and mucous material. Eggs ap- 
parently enter the mantle cavity through the vagina, as 
no other female opening was found.) 

Sex and size relationships (figure 53): Specimens less 
than 7.6 mm shell diameter (n = 25) were all males. 
Specimens of shell diameter greater than 8.6 mm (n = 
6) were all fully functional females without male repro- 
ductive structures. Specimens 7.8-8.5 mm shell diameter 
(n = 6) were "transitional" in appearance (figure 45); 
visceral and pallial reproductive organs were clearly fe- 
male, however, with visible remnant of pallial vas def- 
erens and "wound" (= penis scar) at attachment site of 
penis. Pallial vas deferens of smallest "transitional" spec- 
imen present as unciliated duct, partialK opened at prox- 
imal end, extending between penis scar and distal end 
of capsule gland. No positive connections observed. This 
specimen also with nearly transparent gonad containing 
small amount of whitish ovarian tissue in early part of 
coil, and with no sperm in the female system (it appar- 
ently had not yet mated in its female phase). Largest 
"transitional" specimen with fully-developed ovary, faint 
longitudinal marking on integument in position of vas 
deferens, and weak penis scar. Simultaneous possession 
of both ovarian and testicular tissues not observed. 

"Transition" somewhat correlated with date of col- 
lection (figure 53). Collections made between March and 
mid-August exclusively male. Fully developed females, 
"transitional" individuals, and males encountered in each 
of four collections in later part of August and September. 



Figures 38-43. Cyclostremiscus beauii. 38. Left lateral view of visceral coil, showing position of stomach. Dashed line indicates 
location of dorsal incision. 39. Stomach, dorsal view, opened at incision shown in figure .38. .\rrows inside stomach indicate flow^ 
of particles, 40. Gastric shield. 41. Right jaw, inner side, posterior end up. showing details of jaw elements. 42. Head with mid- 
dorsal incision show ing buccal mass, circumesophageal nerve ring, and base of penis. Connective tissue and minor nerves removed. 
43. Central nervous s\stem, left side, oblique lateral view. Scale bars: 39, 42 = 0.5 mm; 41, 43 = 0.1 mm 

bg. buccal ganglion; bm, buccal mass; cpc, cerebropedal connective; cs, crystalline style; ct, ctenidium; dg, digestive gland; dgo, 
opening of digestive gland; es, esophagus; eso. opening of esophagus; gon, gonad; gs, gastric shield; in. intestine; ino. opening of 
intestine; ki. kidney; leg, left cerebral ganglion; Iplg. left pleural ganglion; Ipg. left pedal ganglion; mo, mouth; mpg. metapodial 
ganglion; on, optic nerve; os, osphradium; pch. posterior chamber; peb, base of penis; pipe, pleuropedal connective; plspec, 
pleurosupraesophageal connective; ppg, propodial ganglion; pvd, pallial vas deferens; ra, radula; ras, radular sac; rcg, right cerebral 
ganglion; rplg, right pleural ganglion; sag, salivary gland; sbeg, subesophageal ganglion; speg, supraesophageal ganglion; ss, style 
sac; st, stomach; stc, statocyst; stl, statolith; tns, swelling of tentacular nerve; ty, typhlosole. 



Page 14 



THE NAUTILUS, Vol. 102, No. 1 




spc'^ spc 

Figures 44-46. Cyclostremiscus beauii. reproductive system (semi-diagrammatic) 44. Male phase 45. "Transitional female 
phase. 46. Female phase, with three cross-sections (A, B, C) through female glands. Small arrows indicate reflected organs 

ag, albumen gland; an, anus; be. bursa copulatrix; eg, capsule gland; co, coiled oviduct; cont. connective tissue; ct, ctenidium; hg, 
hypobranchial gland, If. lateral fold, ov, ovary; pe, penis; pr, prostate gland; prom, opening of prostate gland into mantle cavity; 



R. Bieler and P. M. Mikkelsen, 1988 



Page 15 



Mating and larval development unknown. Protoconch 
morphology suggests planktonic veliger stage (see Dis- 
cussion). Sex change within single individual not ob- 
served. 

Habits and habitat: To our knowledge, this is the first 
record of habitat type for this species, i.e., within the 
burrows of thestoniatopod L(/sio.sq(n7/a scabricaiida, oth- 
er published records (,see synonym) ) refer only to empty 
shells. Wells et al. ( 1961 ) recorded Cyciostremiscus beatiii 
from the stomachs of Astropecten articutatus (Say, 1825), 
however it is unclear whether the material was alive 
when swallowed b> the starfish. Another specimen from 
gut contents of A. articutatus, in the UNC-IMS collec- 
tions, from 24 m depth off North Carolina, contained 
dried animal tissue and is the only other verified live- 
collected specimen known to the authors. Although bur- 
rows of other local species, e.g., callianassid shrimps, 
polychaetes, hemichordates, and sipunculans, were also 
sampled, these vitrinellids were not collected in associ- 
ation with an\- burrower but Lijsiosquilla during this 
stud\ . 

The snails probabK feed on algae, bacterial films and 
detritus in the burrows. They are capable of handling 
larger items in their alimentary tract, as evidenced by 
various shell pieces and foraminiferan shells found in the 
stomach. From the absence of glandular esophageal 
pouches and the presence of a crystalline style in the 
stomach, it can be inferred (Yonge, 1930) that free pro- 
teolytic enzymes, capable of digesting animal matter, 
are not present in this species and would not be expected 
in this group. 

Individuals were almost invariably collected in groups 
of more than one animal per burrow sample; the max- 
imum number encountered in one burrow sample was 
seven. Cyciostremiscus heauii was twice found crawling 
openly on sand or seagrass in the vicinity of Lysiosquilla 
burrow openings. In captivity, the animals were active 
crawlers and were not distracted by light; dark, sheltered 
areas in the tank (provided by black plastic film con- 
tainers) were not preferred. During resting periods, all 
specimens attached themselves to the wall of their con- 
tainer, just above the water level, by means of highly 
\ iscous mucus produced by the anterior pedal gland. This 
behavior was not correlated with food availability or 
water qualitv'. It might reflect tidal rhythm, but material 
was not sufficient to test that hypothesis. 

Geographical distribution: Western Atlantic, from 
North Carolina to Brazil (Ceara and Alagoas; teste Rios, 
1985:41). Recorded from the Pliocene of Jamaica and 
Florida (see synonymy). 

Taxonomic remarks: Fischer (1857b;286), in the orig- 
inal description of Adeorbis heauii (in his earlier pub- 
lication, the name occurred only as a nude list name; 



1857a;173), did not give an indication of the number of 
specimens in the original lot. Moore (1964T32) men- 
tioned a "holotype" in the Paris Museum. However, the 
single A. fceauii-specimen in that type collection (MNHN 
unnumbered, vidi) is much smaller than the dimensions 
given by Fischer (teleoconch diameter 6.3 mm, height 
3.5 mm, protoconch diameter 0.42 mm, umbilical di- 
ameter 1.1 mm; protoconch whorls 2, teleoconch whorls 
2%; vs. 10.5 mm maximum shell diameter in the original 
description). The specimen is therefore considered the 
only remaining syntype of a formerly larger lot, and is 
here selected as lectotype. 

Woodring (1928:440) pointed out the clo.se resem- 
blance between Guppy's (1866) Cyctostrerna bicarinata 
and Adeorbis beauii Fischer, but separated the two be- 
cause the latter "has a more strongly sculptured base." 
Smith (1937:67) in turn separated his new species CAr- 
culus stirophorus from both "Circuhis bicarinatus Gup- 
py" and "Circuhis beauii Fischer, ' by differing numbers 
of upper and basal spiral ribs. A re-examination of the 
neotype (Woodring, 1928) of Cyclostrema bicarinata 
(USNM 115621) and the original description and figure 
of Circuhis stirophorus Smith revealed that both fall 
within the range of variation (partly caused by ontoge- 
netic change of basal sculpture) displayed by the Ft. 
Pierce population, and both are here synonymized (see 
also Moore, 1964:131). The holotype of Circuhis stiro- 
phorus could not be located. Although the original author 
indicated the type to be in the MCZ collection, it was 
never received by that institution (K. J. Boss, personal 
communication); it was also not located at the Florida 
State Museum, Gainesville (F. G. Thompson, personal 
communication). 

Cyciostremiscus beauii is the largest vitrinellid in the 
western Atlantic. Two Recent Panamic species are very 
similar in size, shape and sculpture: 

Cyciostremiscus major Olsson & Smith (1951:46, pi. 
3, figs. la,b) from Panama differs from Cy. beauii in 
having more numerous spiral ribs throughout and a more 
rounded periphery (holotype ANSP 187199; figured by 
Olsson & Smith, 1951). 

Cyclostrema gordana Hertlein & Strong (1951:110, pi. 
9, figs. 3, 4, 7; holotype CAS 064803, vidi) from the 
Gulf of California, differs conchologically from Cycio- 
stremiscus beauii in having a prominent double spiral 
rib on the base between the basal keel and the umbilicus 
(see Pilsbry, 1953: pi. 55). Hertlein and Strong (1951: 
110) gave the following measurements for the "unique 
type": maximum diameter 9.7 mm, minimum diameter 
7.0 mm, and height 3.3 mm; our examination of the 
holotype yielded, respectively, 8.6, 6.8, and 3.9 mm. 
Hertlein and Strong (1951:110) compared C. gordana to 
"Cyclostrema angulata A. Adams [1850] from the West 
Indies ' [following "Pilsbry" (= error for Tryon), 1888: 



ps, penis scar; pvd, pallia! vas deferens; pw, posterior pallial wall; re, rectum; rs, receptaculum seminis; sc, sperm channel; spc, 
sperm channel; te. testis; va, vagina; vo, visceral oviduct; vs, part of visceral vas deferens serving as vesicula seminalis; vvd, visceral 
vas deferens. 



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THE NAUTILUS, Vol. 102, No. 1 




Figures 47-52. Cycloslrcmiscus heauii, specinu-iis from Ft. Vwvcv Inlet, Florida (light micrographs or critical-point dried SEM 
preparations). 47. Cross-section through \isceral ma.ss (male), at level of ve.siciila seminalis. 48. Section through prostate, showing 



R. Bieler and P. M. Mikkelsen, 1988 



Page 17 



TW4-1 



3- 



n D 



o 

D 



D 



o o 



o o 
o 



n15 
10 

5i 



2 3 4 5 6 7 8 9 10 11 12 

Imml 

Figure 53. Cyclostremiscus beauii. Plot of maximum shell diameter vs. number of teleoconch whorls (TW). Inset: Histogram 
summarizing sex distribution of collections, Ft. Pierce Inlet population. A = collecting period March-July, B = August 1-15, C = 
August 16-31, D = September, 1987. Open circles = empty shells and unsexed specimens. Open squares = functional males. Cross- 
hatched open squares = "transitional" females with pallial vas deferens and/or penis scar. Solid squares = females without male 
structures. Star indicates lectot>pe specimen (MNHN unnumbered). 



92, they considered C. angulata a senior synonym of 
Cijclostremiscus beauii — see below] and stated that it 
differed "principally in the smaller size and more de- 
pressed form." However, a re-examination of the C. gor- 
dana t%pe showed it to differ principaJK- b\' a strong, 
second basal keel surrounding the umbilicus. It definitely 
belongs in the genus Cyclostremiscus, as advocated by 
Keen (1971), 

Tryon (1888), followed by Hertlein and Strong (1951), 
considered Cyclostremiscus beauii a synonym of Cy- 
clostrema angulata A. Adams, 1850: "C. angulata, A. 
Ad. was described as from the Philippines on the au- 
thority of Cuming, but as that great collector sometimes 
made mistakes, the localit> needs confirmation. There 
can be no doubt of the identity with this species of C. 



Beaui, Fischer (fig. 63), a West Indian species" (Tryon, 
1888:92-93). Tryon 's figures (1888: pi. 32, figs. 64, 65) 
may represent Cyclostremiscus beauii. However, as sim- 
ilar species are known from the Indo-Pacific, we feel that 
the two nominal species should not be synony mized with- 
out a re-examination of Cyclostrema angulata type ma- 
terial, which could not be located in the British Museum 
(Natural History) (London) nor in the Redpath Museum 
(Montreal). 

The type species of Cyclostremiscus, Vitrinella pan- 
amensis C. B. Adams, 1852, and several other, similar 
species (see, e.g., Pilsbry & Olsson, 1945) have distinct 
axial sculpture which is lacking in Cy. beauii, Cy. major, 
Cy. gordana, and a number of other nominal species. 
Pilsbry (1953) described the subgenus Ponocyclus in Cy- 



lumen and opening into mantle cavity. 49. Section through circumesophageal nerve ring, with saUvary glands. 50. Spermatozoon 
(SEM). Arrow indicates junction of niidpiece and tail. 51. Same, detail of two heads (SEM). Arrow indicates junction between head 
and midpiece. 52. Cross-section through vagina (left) and rectum (right). Scale bars: 47-49, 52 = 0.1 mm; 50 = 4 ^m; 51 = 2 ^m. 

cc, cerebral commissure; cont, connective tissue; ope, cerebropedal connective; dg, digestive gland; es, esophagus; hg, hypobranchial 
gland; leg, left cerebral ganglion; Ipg, left pedal ganglion; mc, mantle cavity; mw, mantle wall; pc, pedal commissure; pch, posterior 
chamber of stomach; pr, prostate gland; prom, opening of prostate gland into mantle cavity; rcg, right cerebral ganglion; re, rectum; 
rpg, right pedal ganglion; sag, salivary gland; te, testis; tm, transverse muscle band; va, vagina; vs, part of visceral vas deferens 
serving as vesicula seminalis; vvd, visceral vas deferens. 



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THE NAUTILUS, Vol. 102, No. 1 




R. Bieler and P. M. Mikkelsen, 1988 



Page 19 



clostremiscus \\ itli Adeorbis beauii as the type species, 
and distinguished it from Cyclostreniiscu.s by its lack of 
axial sculpture. As Pilsbr\ alread\ pointed out in his 
description of Ponocijclus (1953:426). there are a number 
of species showing intermediate conditions ("weak traces 
of axial sculpture") and the name is not used here. 

Due to the small number of species studied in the 
\ itrinellidae, little can he said about anatomical char- 
acters on the generic level. Aside from the large and 
elaborately-sculptured shell, the extensive ciliation of the 
head-foot, and the paddle-shaped pallial tentacle, Cy- 
clostremiscus beauii differs anatomically from Circulus 
striatus (see Fretter, 1956), Ci. texaniis (see below), and 
Cochliolepis parasitica (see Moore, 1972) in ha\ing the 
pallial portion of the intestine straight and uncoiled. 

Circulus Jeffreys, 1865 (page 315) 

Type species by monotypy: Delphinula duminiji Re- 
quien. 1848 (page 64; not "Trochus duminyi Requien" 
as usually cited) [= Yalvata? striata Philippi. 1836; see 
Jeffre\s, 1865:317, where he used the junior synon> m D. 
duminyi only to a\oid secondary homonymy with Tro- 
chus striatus Linne, 1767]. 

Circulus texanus (Moore, 1965) new combination 
(figures 54-68; tables 1, 2) 

Vitrinella texana Moore, 1964:66 [unpublished dissertation). 

Vitrinella texana Moore, 1965:76, pi. 7. figs. 4-6 [Gulf of 
Mexico]. 
? Vitrinella texana "Moore, 1964" — .■Andrews, 1971:73-74, text- 
fig. — Andrews, 1977:887 [citing "1965"], text-fig. [poor fig- 
ure; = Vitrinella sp., teste Ode, 1987.35]. 

Vitrinella (Vitrinella) feiana,— Abbott, 1974:83. 

Vitridomus texana. — Ode, 1987:37. 

Material examined: Ho]ot\pe: empt\ shell, 1.8 mm, 
USNM 636311 (Texas). Other material: 15 specimens. 
FLORIDA: Ft. Pierce Inlet: 2-3 May 1987, 1 unsexed; 
24 June 1987, 14 specimens (4 males, 7 females, 2 un- 
sexed, 1 empty shell). 

Description 

Teleoconch (figures 54-56): Shell small (1.7-1.8 mm 
diameter, 0.55-0.65 mm height), with 1V2-1% teleoconch 
w horls; almost planispiral, sculptured dorsalK' and ven- 
trally with about 18 fine spiral ribs; transparent when 
alive, opaque after death. Ribs slightly stronger, more 
widely spaced just below suture on dorsum and at pe- 
riphery, where about 3 ribs form rounded keel below 



lateral midline. Suture impressed. Ventral surface below 
keel less convex, often with 30-40 widely-spaced, low 
axial ribs which are primarily evident from inside of 
body whorl (figure 57). L'mbilicus wide (~ 25% of width). 
Outer lip ver>- slightly reflected; some specimens with 
one former varix. Aperture at oblique angle to dorso- 
ventral axis. Sutural sinus shallow. Periostracum thin, 
transparent, with spiral grooves more numerous than on 
shell surface. 

Protoeonch (figure 58): Smooth, 0.5 mm diameter, 
about 2 whorls. No sculptural demarcation separating 
protoeonch I and protoeonch II. 

External anatomy and organs of the mantle cavity (fig- 
ures 64-66): Living animal translucent white with 
buccal mass (visible through proboscis) and protrusible 
portion of ctenidium rose-pink; digestive gland orange; 
ner\e ring area opaque yellow, visible through integu- 
ment between eyes. Head with long, extensible snout, 
rounded and terminally notched at mouth. Ventral side 
of snout ciliated behind mouth opening, with cilia beat- 
ing toward mouth. Two long cephalic tentacles w ith im- 
mobile bristles on slight]} enlarged tip; motile cilia at 
least at tip and along side facing snout. Left cephalic 
tentacle fitting into shallow notch formed by shell keel 
(figure 65). Eyes black, on slight bulges at base of cephalic 
tentacles. Neck region very long, slender. Foot elongate, 
flattened, with anterolaterally recurved corners; fineK 
ciliated sole broadly rounded posteriorly, extending well 
beyond shell (figure 65). Transverse crease on sole about 
'/^ of total length from anterior edge. Anterior pedal 
mucous gland (figure 65, amg) opening at slit across 
entire leading edge; mid- ventral fold evident on posterior 
quarter of sole, but presence of posterior mucous gland 
not confirmed. Locomotion by ci]iar\ action. Operculum 
(figures 61-63) corneous, circular, multispiral {ca. 8 
whorls), with small central peg and rounded-triangular 
muscle scar on inner surface. Operculum supported by 
opercular lobes (figure 65, ol) on dorsal side of foot. Lobes 
simple, unpigmented, without tentacles. Epipodial ten- 
tacles absent. 

Two finger-shaped pallial tentacles (figure 64, Ipt, upt) 
arising separately but adjacently from inner mantle edge, 
at right side of animal. Upper (= most dorsal) tentacle 
unciliated, with immobile terminal bristles, curling into 
shallow sutural sinus, directed dorsalK during crawling. 
Lower tentacle bearing motile cilia, directed anterolat- 
erally. Ctenidium (figures 64, 65, ct) with numerous fin- 
ger-shaped filaments, attached for most of its length to 
internal surface of mantle; anteriormost filaments darker 



Figures 54-63. Circulus texanus, specimens from Ft. Pierce Inlet, Florida (SEM) (figures 54-56, USNM 846324). 54. Shell, apical 
view (2.7 mm diameter). 55. Shell, umbilical view (2.7 mm diameter). 56. Shell, apertural view (2.2 mm diameter). 57. Inside 
surface of base of bod> whorl, showing shallow radial grooves. 58. Protoeonch. 59. Larval shell, left lateral view (167 /im max. 
diameter! 60. Radula. 61. Operculum, inner surface (0.7 mm diameter) 62. Operculum, outer surface (0.8 mm diameter). 63. 
Operculum, oblique view of inner surface with opercular peg. Scale bars: 57, 58 = 0.2 mm: 60 = 5 ^m; 63 = 50 ^m. 



rac, rachidian teeth. 



Page 20 



THE NAUTILUS, Vol. 102, No. 1 








Figures 64-68. Circulus texanus. 64. Crawling female, dorsal view, drawn as with transparent shell. 65. Crawling animal, ventral 
view, drawn as with transparent shell. 66. Head of male, left lateral view, showing penis. 67. Egg mass 68. Two egg capsules, 
each with veliger larva shortly before hatching. Scale bars: 64, 65 = 0.5 mm; 66, 67 = 0.25 mm. 

ag, albumen gland; amg, anterior mucous gland; br, immobile bristles; eg, capsule gland; ct, ctenidium; dg. digestive gland, ebv, 
efferent branchial vessel; ey, eye; he, heart; in, intestine; ki, kidney; Ipt, lower pallial tentacle; ol, opercular lobe; op, operculum; 
ov, ovary; pe, penis; re. rectum; sh, shell; sn, snout; ss, style sac; st. stomach; ten, cephalic tentacle; umb, umbilicus; upt, upper 
pallial tentacle. 



in color, not attaclu'd to tnantle skirt, extending out ot 
aperture at right side of head. E.xtent of osphradium not 
ascertained. 

Alimentary system: Riiduia (figure 60) taenioglossate. 
Rachidian tooth wider than long, with acute posterior 
corners projecting laterally and concave front. Main cusp 
narrow, unserrated, with 5-6 slightly smaller flanking 
cusps on each side (number of flanking cusps var\ ing 
within single radular rihhon) decreasing laterally in size; 
base with 1 weak elevation per side (in position of basal 
denticle), midway between posterior corners and central 
ridge. Lateral tooth with asymmetrical cutting edge, 
strongly indented at front edge, with narrow , unserrated 



main cusp and subequal flanking cusps (6-7 inner, 9 
outer). Apex of inner marginal tooth with numerous long, 
thin, flanking cusps on either side of similarly-sized main 
cusp. Apex of outer marginal tooth with long, thin inner 
flanking cusps, slightly more robust than tho.se of inner 
marginal tooth; cusps at termiiuis subequal in size; outer 
margin with 2-3 cusps decreasing in size, remainder of 
outer margin smoothly rounded. Jaws composed of ele- 
ments approximately 4 ^m in length. 

Stomach traii.spinciit, with rotating style often clearly 
visible through shell. Posterior part of stomach not mark- 
edly elongated. Intestine (figures 64, 65, in) forming wide 
loop in posterior part of mantle cavity. Anus just above 
anterior end ol capsule gland in females. Fecal material 



R. Bieler and P. M. Mikkelsen, 1988 



Page 21 



as continuous rods of irregular length, rather than pellets, 
with no obvious surface sculpture. 

Reproductive system: Penis ifigure 66, pe) a posteriorly 
directed, counterclockw ise, double coil; base arising pos- 
terior to eyes, somew hat right of dorsal midline. Testis 
orange. 

Ovary (figure 64, ov) filling inner side of early whorls 
with grape-like lobes, lighter in color than digestive gland. 
Capsule gland and much darker-staining (in metln lene 
blue) albumen gland (figure 64, eg, ag) forming compact 
mass on right wall of mantle cavity, below rectum, ter- 
minating anteriorly at level of anus. Position and number 
of sperm pouches not ascertained. 

No evidence of se.xual dimorphism associated with 
possible sequential hermaphroditism. Males both consid- 
erably smaller (1.7 mm diameter at 1% whorls) or larger 
(1.8 mm diameter at 1': whorls) than females in same 
sample (1. 74-1. 78 mm at I'/s whorls). One copulation 
briefly observed, wherein male paused while crawling 
over dorsal surface of female, with heads adjacent (male 
at left) and at angle of about 30°. Position of penis not 
ascertained. 

Eggs and larval development: Three weeks after col- 
lecting, 7 females and 4 males were placed into glass 
bowl with seawater (24 °C). Overnight, 15 egg masses 
were laid; each mass contained 1-5 eggs, each in clear, 
flattened, circular capsule (egg diameter 1 10 fim: capsule 
diameter 170 ^m; n = 20). All eggs found in same de- 
velopmental stage, uncleaved, completely round, finely 
granular. Soft capsule walls of eggs laid in groups touch- 
ing each other; each "mass," even if consisting of single 
egg, coated b\ sticky, clear, gelatinous layer (figure 67). 
Eighteen additional egg masses were found in glass jar 
in which specimens had been kept since collecting. Total 
number of masses (with respective numbers of single- 
egg-capsules) was: 4 masses with 1 capsule, 4 masses with 
2, 13 masses with 3, 9 masses with 4, and 3 masses with 
5 capsules. 

One mass with 5 eggs followed through development: 
From beginning of equal, holoblastic 2-celled stage, 60 
minutes elapsed to 4-celled stage, 205 minutes to 8-celled 
stage, 385 minutes to 16-celled stage, respectively. Gas- 
trula-stage discernible after 1 1 hours; embryos rotating 
after 17 hours. 

X'eliger larvae (figure 68) fully developed at 104 hours, 
intermittently or constantly rotating, with transparent 
shell, 2 short veliger lobes, black eye spots (not situated 
on tentacles at this point), statocysts and operculum [much 
as described for Caecum glabrum by Gotze (1938:108, 
text-fig. 33), but with smaller vela]. Periphery of velar 
lobes bearing long cilia, underlain by row of shorter cilia. 
Capsule size now approximately 150 x 120 fim, greatest 
shell diameter 107 /xm, longest dimension of veliger with 
extended \ela 133 ^m. During entire process, 3 of 5 
embryos developed completely synchronously; other 2 
initially about 45 minutes, later several hours, behind. 
First veligers hatched after 130 hours and swam actively. 
For follow ing 36 hours, veligers observed either swim- 



ming or resting on bottom of bowl, with strong ciliary 
action on extended vela. No further development noted. 
Added cultures of mixed single-celled green algae ap- 
parently not excepted; all larvae died within next 2 days. 
[From the size difterence between these larvae (170 nm) 
and the protoconchs of adult shells (500 nm), a longer 
lar\al stage, w ith extensive food intake can be predicted.] 

Habits and habitat: Habitat as described for Cyclo- 
stremiscu.s hcaiiii. In the laboratory, animals of Circiilns 
texanus were very active and fed on normally-occurring 
algal and bacterial surface films. 

Geographical distribution: Circuhis texanus (type lo- 
cality: Mustang Island, near Port Aransas) was previously 
reported to be endemic to the northv\estern Gulf of Mex- 
ico, and has never been reported alive (Moore, 1964:41; 
.\ndrews, 1977:87; Ode, 1987:37). No fossil records are 
know n. 

Taxonomic remarks: Moore (1965:77) placed this species 
in the genus Vitrinclla C. B. Adams, 1850, and pointed 
out the differences in shell shape and sculpture in com- 
parison to other species. However, members of Vithnella 
(based on the type species V. helicoidea C. B. Adams, 
1850; see tables 1, 2) are generally smooth-shelled or 
weakly sculptured and are usually furnished with a ca- 
rina bordering the umbilicus (e.g.. Keen. 1971:377; Ab- 
bott, 1974:82). We feel that this species is better placed 
in Circuhis sensu lato because it agrees in shell shape 
and sculpture with the type species, Circuhis striatus 
from the eastern Atlantic. Ode's (1987:37) placement of 
this species in VHridomus Pilsbry & Olsson, 1945, cannot 
be accepted: members of Vitridomus [a "rather feebly 
defined genus (Pilsbry & Olsson, 1952:36), very similar 
to Teinostoma H. & A. Adams, 1853] ha\ e the umbilicus 
partly covered by a callus. 



DISCUSSION 

Comparison of the anatomical information on Cyclo- 
stremiscus beauii and Circuhis texanus with literature 
data revealed that published information is scarce or 
misleading, and that genus- as well as family-level taxa 
of marine near-planispiral Rissoacea are poorly defined. 
Three nominal families, Vitrinellidae Bush, 1897, Tor- 
nidae Sacco, 1896, and Circulidae (for authorship, see 
below), are currently in use for members of this group, 
and the lack of comparative data led Ponder (in press) 
to treat all three families as a single group in a phylo- 
genetic analysis of the Rissoacea. In the follow ing dis- 
cussion, we (a) review the available data on species of 
this complex, (b) compare Cy. beauii and Ci. texanus 
w ith these data and with other members of the Rissoacea, 
(c) suggest a preliminary grouping for the species for 
w hich anatomical data are available, (d) demonstrate the 
differences between Vitrinellidae and Tornidae, (e) eval- 
uate the taxonomic status of the nominal family Circu- 
lidae, and (f ) discuss some unusual aspects of the anatomy 
and reproductive biology of Cy. beauii. namely ciliation. 



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THE NAUTILUS, Vol. 102, No. 1 



tentacle shape, stomach morphology, and hermaphro- 
ditism. 

(a) Available literature data (see tables I, 2). 

Except for Fretter's (1956) study of Circulus striatus, 
and Woodward's (189S) and Graham's (1982) work on 
Tornus suhcarinatus, most of the available data on the 
anatom\ of marine near-planispiral Rissoacea are re- 
stricted to descriptions or illustrations of external mor- 
phology, especially Pilsbr\ and McGinty's (1945a, 1946b) 
sketches of western .\tlantic forms. Some of the few 
published observations on the radular structure and gross 
morphology of vitrinellid softbodies are misleading (see 
also Moore, 1964, 1972). Based on the misconception that 
these species are archaeogastropods. Rush (1897:127, 142, 
pi. 22, figs. 12a-g) erroneously constructed a rhipido- 
glossate radula for Circulus trilix (Bush, 1885) [= Cy- 
clostremiscus pentagonus (Gahh, \8S7), fide Moore, 1964; 
138]. The "supplementar\ plicated gill" (Stimpson, 1858: 
308) of Cochliolepis parasitica uas not found in that 
species by Moore (1972:101). Pilsbry (1953:427) men- 
tioned a taenioglossate radula for Cyclostremiscus beauii, 
and added "This will be figured in a paper now in prep- 
aration b\ T.L. McGinty and the author " This work was 
apparentl) never published. 

(b) Comparison of Cyclostremiscus beauii and Circulus 
texanus with other \itrinellids and rissoaceans. 

Both Cyclostremiscus beauii and Circulus texanus dis- 
play "typical" rissoacean characters (as outlined by Fret- 
ter & Graham, 1978:153; Boss, 1982:984; Ponder, 1983, 
in press) and agree to a large extent with the previous 
concept of Vitrinellidae (Fretter, 1956; Moore, 1972; see 
tables 1, 2, Group 1). As in most other vitrinellid species, 
the cephalic and pallial tentacles bear immobile bristles 
and/or motile cilia. Tracts of motile cilia on the cephalic 
tentacles occur in numerous groups in the Rissoacea; for 
lack of comparative data, it is not yet clear whether the 
arrangement of these cilia in grooves, as in Cy. beauii, 
is unusual. Photographs of critical-point dried tentacles 
of a freshwater rissoacean, the hydrobiid Tryonia ctath- 
rata Stimpson, 1865 (see Hershler & Thompson, 1987: 
27, figs. 13-17), show no grooves present in that species. 

Cyclostremiscus beauii and Circulus texanus have two 
closely-spaced pallia! tentacles on the right side. This 
arrangement is present in most studied vitrinellids (the 
exception being Cochliolepis albiceratus Ponder, 1966), 
in the type species of Tornus (Tornidae, see below), in 
Hydrococcus brazieri (T. Woods, 1876) (monotypic Hy- 
drococcidae, see Ponder, 1982), and in the rissoid genus 
Rissoina Orbigny, 1840, where some species have either 
the anterior (right) or posterior (left) pallial tentacle bi- 
lobed (Ponder, 1985:78). As described for Cy. beauii, the 
function of the upper tentacle, which usually bears im- 
mobile bristles, and is held upright and bent around the 
aperture, is clearly sensory, while the lower one, often 
distinctly ciliated and sometimes broadened, controls and/ 
or enhances water flow. 

Both species have a small central opercular peg which 
is probably not homologous with the lateral "neritid- 



type " peg in other families of Rissoacea (character con- 
sidered plesiomorphic in this superfamily), while the cir- 
cular shape of the vitrinellid operculum is considered 
derived (Ponder, 1985:5). 

The osphradium of Cyclostremiscus beauii is ver) sim- 
ilar to those described for Circulus striatus (see Fretter, 
1956:372) and for species of the Rissoidae (Johansson, 
1939:319, pi. 3, figs. 3, 4, text-fig. 5; Haszprunar. 1985: 
476, figs. 7k, 16). 

The radulae of the two species studied here ha\ e only 
one pair of basal denticles (or elevations) on the rachidian 
tooth, a feature known from some other rissoacean fam- 
ilies, such as Hydrobiidae (see Bandel, 1984:29, text-fig. 
47) and Rissoidae (see Ponder, 1985:10). This character 
was hvpothesized to be primitive in the Rissoacea by 
Ponder (1985:119). 

The position of the salivary glands relative to the cir- 
cumcsophageal nerve ring is often used in family- and 
superfamiK -level discussions (e.g., Ponder, 1983:236, 258; 
in press). However, as a highly variable character in the 
populations of Cyclostremiscus beauii studied herein, 
further data on additional species are necessarv before 
it can be reliabl) utilized to infer phylogenetic relation- 
ships of the Vitrinellidae. 

The extensive anterior section of the esophagus of Cy- 
clostremiscus beauii, with its long, coiled, dorsal folds, 
is verv similar to that of Hydrococcus brazieri (see Pon- 
der, 1982:77). 

The conspicuous ciliated tract leading from the renal 
opening to the head of Circulus striatus (see Fretter, 
1956:372) is lacking in Cyclostremiscus beauii. 

The presence of markedly vacuoled connective tissue 
in Cyclostremiscus beauii is a character shared by Cir- 
culus striatus and Tornus subcarinatus (see Graham, 
1982:147). 

The nervous system of vitrinellid species is tvpically 
rissoacean and shows moderate (Cyclostremiscus beauii) 
to high (Circulus striatus; Fretter, 1956:377) concentra- 
tion. The RPG ratio of Cy. beauii, averaging 0.49. lies 
within the range of those of members of the rissoacean 
family Pomatiopsidae (see, e.g., Davis ii Mazurkiewicz, 
1985:45, table 8). 

The structure of the male and female reproductive 
systems encountered during this stud\ is ris,soacean (e.g., 
Johansson, 1956). As in most other rissoaceans (e.g.. Pon- 
der, 1985:6), the albumen and capsule glands form a 
single, continuous mass [a character not recognizable from 
Fretter's (1956) description and figures of Circulus stria- 
tus]. The N'itrinellidae differ from the Rissoidae (e.g.. 
Ponder, 1985: text-fig. 2) wherein a non-homologous 
structure, an expanded part of the upper oviduct, pos- 
terior to the bursa copulatrix and/or receptacula seminis, 
functions as an albumen gland. This "upper oviduct 
gland " of the Rissoidae is homologous with the coiled 
part of the visceral oviducts of the Hvdrobiidae (see 
Johansson, 1956) and N'itrinellidae [Fretter's (1956) "ren- 
al oviduct ']. 

Most members of the Rissoacea have one bursa cop- 
ulatrix and one receptaculum seminis near the posterior 



R. Bieler and P. M. Mikkelsen, 1988 



Page 23 



pallial \\ all, ifquii iiig the sperm to travel the entire length 
of the sperm ehannel (vaginal lumen, sperm groove of 
the ventral channel of authors) immediately after cop- 
ulation. Others have develeoped additional, ilistal sperm 
pouches (anterior sperm-storage structure, distal l)lind 
sac, sac-hke vestibule, spermatheca, pseudo-bursa of au- 
thors), eitlier in addition to [e.g., Ptisillina incon.^pictia 
(Alder, 1844); see Johansson, 1939:337. text-fig. 22 (as 
Rissoa); Rissoidae], or instead of the pro.ximal bursa cop- 
ulatrix [e.g., Htjala vitrea (Montagu, 1803); see Johans- 
son, 1949: text-fig. 1; Iravadiidae]. These distal sperm 
pouches are not necessarily homologous with each other 
and certainly not with the proximal bursa (see, e.g., Sla- 
voshevskaya, 1978). Vitrinellids for which such data are 
available (CArciilus striatus and Cyclostremiscus beauii) 
differ from most other rissoaceans in the presence of two 
more-or-less eciually developed, proximal receptacula 
seminis which, in position and size, look much like the 
two sperm sacs {i.e., bursa copulatrix and receptaculum 
seminis) of other Rissoacea {e.g., in the rissoid Lucidestea 
Laseron, 1956; see Ponder, 1985:67, text-fig. 3). Whether 
the presence of two proximal receptacula seminis is a 
distinguishing character for Vitrinellidae cannot yet be 
determined, as accessory receptacula have been dem- 
onstrated for members of the genus Alvania in the Ris- 
soidae (Johansson, 1956; Ponder, 1985). In both Circiilus 
striatus (see Fretter, 1956:377) and Cyclostremiscus 
beauii, the distal end of the coiled oviduct serves as the 
fertilization area, and not as an additional functional 
receptaculum as has been reported for some other ris- 
soaceans [e.g., Alvania suhsoluta (Aradas, 1847), where 
the distal end contained oriented sperm; Johansson, 1956: 
380]. 

The spawn mass of members of the Rissoidae usually 
has numerous eggs per capsule in planktonic forms, while 
one-egg-per-capsule is characteristic of "direct ' devel- 
opment (see Lebour, 1937; Thorson, 1946). This does not 
hold true for the spawn of Circiilus texanus, which gen- 
erally resembles that of the freshwater rissoacean Bithijn- 
ia tentaculata (Linne, 1758), as described and illustrated 
by Ankel (1936:164, text-fig. 142B). [The number of eggs 
there, however, is larger (4-24; Lilly, 1953:104), and 
hatching occurs at the crawling stage]. Jablonski and Lutz 
(1980:336; after Taylor, 1975) stated that the Rissoacea 
". . . follow one of two developmental pathways: those 
that hatch as crawling juveniles from relatively large eggs 
(140-320 nm) and those that hatch from relatively small 
eggs (60-130 jxm) and spend 2-3 weeks as planktonic 
veligers. Both species studied here definitely fall into 
the latter category: veliger shells of Circulus texanus 
(figure 59) were one full whorl smaller than the final 
larval shells as seen in the adult protoconch (figure 58); 
the protoconch of Cyclostremiscus beauii (figures 9-11) 
shows a distinct line and change of sculpture between 
protoconch I and protoconch II, with almost another full 
whorl of growth before metamorphosis, suggesting sev- 
eral weeks (K. Bandel, personal communication) of 
planktonic life. In his unpublished revision of western 
Atlantic Vitrinellidae, Moore (1964:18) inferred that 



"most, if not all, species appear to have a planktonic 
veliger stage of some duration." 

The sperm cells of Cyclostremiscus beauii are of the 
general type known tor other members of Rissoacea (see 
Gotze, 1938; Franzen, 1955). The twisted acrosome and 
relative lengths of head, midpiece and tail are similar to 
those described by Franzen (1955) for Caecum glabrum 
(Montagu, 1803). The head is much shorter than that of 
the two rissoids previously studied [Pusillina incon.spicua 
(Alder, 1844) (as Rissoa) and Onuba striata (J. Adams, 
1797); Franzen, 1955:406-409], and the relative length 
of the midpiece much greater than in Hydrolna ulvae 
(Pennant, 1777). 

Direct communication of the coiled ("renal") oviduct 
with the kidney (Fretter, 1956), previously considered 
unique for Circulus or the Vitrinellidae, is now also known 
for other families of the Rissoacea (Tornidae, Graham, 
1982; Truncatellidae, Fretter & Graham, 1962). 

Anatomical characters of Circulus striatus, which 
Fretter (1956:380) discussed as probably "associated with 
small size and body form," are equally expressed in the 
much larger Cyclostremiscus beauii, suggesting that 
Fretter's hypothesis was incorrect. 

(c) Preliminary grouping of "vitrinellid-like" species for 
which anatomical data are available. 

The species for which sufficient gross morphological 
data are available appear to fall into three groups, two 
of which are here considered of familial rank and one 
comprising species of incertae sedis (tables 1, 2). Mem- 
bers of Group 1, currently placed in the genera Vitri- 
nella, Teinostoma, Pieurornalaxis, Cochliolepis, Circu- 
lus and Cyclostremiscus, are here considered to belong 
to the Vitrinellidae (the placement of the species de- 
scribed as Cochliolepis albicerata Ponder, 1966, is some- 
what doubtful as it has only one pallial tentacle, and the 
morphologies of its osphradium and penis are unknown). 
Some of the characters attributed to the family Vitri- 
nellidae in Moore's dissertation (1964) were based on 
Parviturboides interruptus (C. B. Adams, 1850) (tables 
1, 2, Group 2), later considered a species of uncertain 
systematic position (Moore, 1972:107). This species dif- 
fers from the other forms here grouped in Vitrinellidae 
by the following anatomical characters: posterior foot 
margin with immobile cilia, left cephalic tentacle with 
four low swellings on proximal posterior border, penis 
with glandular area and directed straight back. Another 
species, Tomura bicaudata (Pilsbry & McGinty, 1946) 
differs in many head-foot characters (tables 1, 2, Group 
2) and is here also considered as incertae sedis. White 
(1942:92) advocated the inclusion of "Cyclostrema" bushi 
Dautzenberg & Fischer, 1907, in the Vitrinellidae/Tor- 
nidae complex and published anatomical descriptions 
and illustrations of that species. While some morpho- 
logical characters [head with well-developed eyes and 
terminally ciliated cephalic tentacles, the well-developed 
osphradium (1942: text-fig. 6), and the circular, multi- 
spiral operculum (1942: pi. 2, fig. 3)] agree well with the 
species listed here as vitrinellids, the glandular pouches 



Page 24 



THE NAUTILUS, Vol. 102, No. 1 



in the esophagus and, most of all, the pair of epipodial 
(not pallial) tentacles on the right side of the animal 
(White, 1942:90, text-fig. 5), prevent inclusion in \itri- 
nellidae. Tornm siibcarinatus (tables 1, 2, Group 3) is 
here considered a member of a separate famiK (see be- 
low). 

(d) Family relationships: Vitrinellidae-Tornidae. 

Several attempts have been made to define the key 
characters of the family Vitrinellidae [e.g., Moore, 1965: 
74, 1969:170, 1972:107ff.; Boss, 1982:991). However, be- 
cause of the small number of species fulK studied ana- 
tomicall), no single synapomorphy defining the famiK 
is clear at this time. The most obvious features of the 
Vitrinellidae are: a low-spired, translucent white shell; 
long cephalic tentacles equipped with terminal bristles 
and, in most cases, motile cilia; two closely-spaced pallial 
tentacles on the right side; a large monopectinate gill 
often projecting to the right of the head; a large linear 
osphradium; a foot with simple or only slightly indented 
anterior and posterior margins; a horn\ concentric, mul- 
tispiral operculum; and the possession of a non-glandular 
curved penis in the male. Some of these features (shell 
shape, pallial tentacles, and projecting gill) have led au- 
thors (e.g., Ta\ior & Sohl, 1962) to s\nonymize Vitri- 
nellidae and Tornidae, based on Woodward s (1898) work 
on Adeorbis [= Tormts] .siibcarinatus (Montagu, 1903). 
Others (e.g.. .\dam & Knudsen, 1969; Moore, 1972) have 
pointed out differences between vitrinellids and Tornus/ 
toniids in features of the e\ es (functional eyes are lacking 
in T. .'iubcarinatiis ) and, or the operculum (oval and pau- 
cispiral in T. siibcarinatus) and have separated the two 
families. In a redescription of the anatomy of T. sub- 
carinatus, Graham (1982:147) saw the opercular shape 
as the "single difference between the two nominal fam- 
ilies and again advocated synonymy. 

Additional anatomical characters of Tornidae that 
warrant separation from Vitrinellidae are: (1) the con- 
spicuous, elongate osphradium of the latter is not present 
in Tornus (Graham, 1982:144, found only "a small cil- 
iated groove, which may be a reduced osphradium"), (2) 
the attachment of the ctenidial axis to the mantle is short 
and the axis hardly supplied with blood vessels, and (3) 
the penis of 7". siibcarinatus bears several finger-like 
processes which are not known in vitrinellids as delimited 
here. [Fretter & Graham (1978:231) described the cte- 
nidium of T. siibcarinatus as "partly bipectinate." This 
is apparentK in error, as both Woodward (1898) and 
Graham (1982) described and illustrated that gill with 
only a single row of lamellae.] As T. siibcarinatus lives 
in a comparable habitat (under boulders on well-oxy- 
genated sand or iiuid) and is of about the same size as 
Circulits spp.. the drastic difference in osphradial type 
indicates phylogenetic difference rather than specializa- 
tion on a low taxonomic level. 

Ponder (in press) combined the "tornicl-\ itrinellid-cir- 
culid complex" as lamiK 'i'ornidae tor the purpose of 
his pin logenetic analysis of the Rissoacea. However, most 
of the characters and character states he attributed to 



that complex (metapodial tentacle present, esophageal 
pouches and glands present, penial glands present, os- 
phradium short, posterior end ot foot not simple) do not 
occur in N'itrinellidae as understood here, and must refer 
to Tornus and other tornid genera studied b> Ponder 
(Pseudoliotia Tate, 1898, Scrupus Finlay, 1927; unpub- 
lished). 

Vitrinellidae and Tornidae fit well into the Rissoacea, 
and an independent grouping of these famiilies as Tor- 
nacea (e.g., Kuroda et al., 1971; Golikov & Starobogatov, 
1975) is not justified. The family Adeorbidae Montero- 
sato, 1884, used b\- some authors for members of this 
complex, is a s\non> m of Tornidae, since Adeorbis S. 
Wood, 1842, is an objective synonym of Tornus Turton 
& Kingston, 1830 (see Iredale, 1914:172, 1915:344). 

(e) The nominal family Circulidae. 

The taxonomic status of the nominal family Circulidae 
remains problematic. The family name "Circulidae" was 
first used b> Fretter and Graham (1962:642. ".\ppendix 
I"), in a list of taxa treated in that publication. The text 
reads merely "Circulidae: Circulus striatus (Philippi). ' 
Fretter and Graham (1962) did not state that they in- 
tended to create a new family, did not mention the name 
in the main bod\ of the text, even u hen the taxonomic 
position of the genus (1962:550, 618) and the composition 
of the superfamiK (1962: 622-623) were discussed, and 
in fact ne\er used any famiK name but N'itrinellidae 
when the\ referred to Ci. striatus in subsequent publi- 
cations (Fretter bi Graham, 1978:227; Graham, 1982: 
147). Lacking an\ description, definition, or bibliograph- 
ic reference, a name thus introduced is not taxonomically 
available (ICZN, 1985: Art. 13). "Circulidae" could 
therefore be regarded as a nomen nudum. 

However, Golikov and Starobogatov (1975:211) ac- 
cepted "Circulidae Fretter & Graham, 1962" as a valid 
family and stated (1975:218) that "the characteristics of 
the latter famiK are found in Fretter (1956:381)," there- 
by referring to the summar\ of Fretter s description of 
Ci. striatus. This fulfills the requirement of ICZN (1985) 
Art. 13(a)ii (Bibliographic Reference to Published State- 
ment) and, unless there are earlier such statements that 
have escaped us, this makes Goliko\ and Starobogatov 
(1975) the authors of Circulidae, with Circulus as the 
name-bearing t\ pe. The nominal family Circulidae is, 
with anatomical descriptions available for Ci. striatus 
and Ci. texanus, much better detined than the X'itrinel- 
lidae, where our knowledge ot Vitrinella is based onK 
on shell characters and Pilsbry and McGinty's (1945a, 
1946b) sketches of crawling animals (tables 1. 2). .\11 
available data suggest s>non\in\ of N'itrinellidae and 
Circulidae (tables 1, 2; Boss, 1982:991 ). In any case, the 
placement of N'itrinellidae and Circulidae in separate 
superfamilies, or even separate superorders (Golikov & 
Starobogatov, 1975), is highly exaggerated. 

(f) Unusual features of Cyclostremiscus beauii. 

The stud) of ('yclustrentisciis beauii revealed a num- 
ber of features that differ from other members of the 
N'itrinellidae (and, in part, from the Rissoacea). Most of 



R. Bieler and P. M. Mikkelsen, 1988 



Page 25 



these features may be related to either its unusually large 
size (for a \ itrinellid) or its unusual habitat in stoniatopod 
burrows. 

Ciliation and tentacle shape: The extensive ciliation of 
the gills, cephalic tentacles and lower pallial tentacle 
provides eflective respirator) and excretory currents. On 
the right side of the head, where in- and outgoing cur- 
rents are present, water flow is controlled by the paddle- 
shaped tentacle. Low tides and high water temperatures 
are likely tc create anoxic conditions in the burrows, and 
a large vitrinellid such as Cyclontremiscus heauii may 
be more strongly affected by oxygen deficiencies than, 
for instance, the smaller Circuhis texanus. and thus would 
benefit from an elaborate s\stem to produce and direct 
currents. The onl\ other known commensal \itrinellid, 
Cochliolepis parasitica, is much smaller, and lives di- 
rectly on its host [under the scales of the giant scale worm 
Pohjodontcs lupina (Stimpson, 1856); see Stimpson, 1858; 
llartman, 1945:10]. Moore (1972:104) did not find cilia 
on the gill filaments of Cochliolepis parasitica, suggesting 
tliat currents produced by the annelid are sufficient to 
supply the snail. All other species studied live under rocks 
(table 2). 

Stomach: The rissoacean stomach does not normally 
have a caecum or an elongated posterior chamber, as 
occurs in various other prosobranchs ("it is not possible 
to see any trace of it in the . . . Rissoacea"; Fretter & 
Graham, 1962:225). OnK a few exceptions are known: 
Ponder (1985:78) described the stomach of Rissoina (Ris- 
soidae) as "very long due to elongation of posterior cham- 
ber," and Ponder (in press) listed the character state 
"posterior gastric chamber not small" for the families 
Emblandidae, Truncatellidae and Stenothyridae. The 
presence of a large posterior chamber in Cijclostreiniscus 
beauii, similar in relative shape and organization to that 
of Pomatias elegans (Miiller, 1774) (Pomatiasidae), as 
described by Graham (1939:90, fig. 6D), is therefore sur- 
prising. It may allow Cij. beauii to maintain a regular 
supply of food particles by regulating fluctuations in the 
rate of food intake (as suggested by Graham, 1939:93, 
for Pomatias). especially since feeding must be strongly 
affected b> tidal, and therefore water-level, changes 
within the stomatopod burrows. A relatively large animal 
such as Cy. beauii would be more strongly affected than 
smaller vitrinellids, e.g., Circulus texanus, which were 
observed to feed in very small pockets of water (personal 
observation), thus maintaining a more-or-less continuous 
feeding activity, as has been described, e.g., for a mem- 
ber of the genus Caecum (Morton, 1975:14). 

Hermaphroditism: The most striking result of this study 
was the discovery of small functional females in Cy- 
clostremiscus beauii with apparently functionless rem- 
nants of the male reproductive system. Penial structures 
are known to occur in various female rissoaceans (par- 
ticularly hydrobiids and rissoids). Thiriot-Quievreux 
(1977:779ff.) based her hvpothesis of sequential her- 
maphroditism in four species of Ris.soa on the presence 
of more-or-less developed penes in immature specimens 



and lemales. However, a later in-depth study (Thiriot- 
Quievreux, 1982:]67ff.) of these species through annual 
cycles shov\ed a seasonal (and geographical) pattern of 
penis-size distribution in females, but no evidence of sex 
change. The survey did "not support the hypothesis of 
a successive hermaphroditism" (1982:167). In Cyclo- 
stremiscus heauii, remnants of the male apparatus are 
only evident in the smallest functional ieniales (figure 
53), with a gradual decrease of male structures with size. 
As outlined above, we take this as evidence for a sex 
change in this species rather than a simple sexual di- 
morphism in shell size. During ontogeny, the male re- 
productive system apparently disappears completely and 
is replaced by female organs. The anteriormost part of 
the pinkish-orange female albumen gland is found in the 
same relative position as the prostate (of similar color 
and reaction to staining) in the male phase. As no animal 
was found with a very early stage of development of the 
albumen gland, it cannot be decided whether these or- 
gans, or parts of them, are homologous. [Reid (1986), in 
a study of Mainwaringia Nevill, 1885, the first reported 
case of protandrous sequential hermaphroditism in the 
Littorinacea, found both a closed pallial oviduct and an 
open prostate in the intersexual and female stages, "sug- 
gesting that these structures are not strictly homologous" 
(1986:225). In that littorinid group, small penes are re- 
tained in the female phase (1986:237), and the pallial 
oviduct opens into the also-retained pallial vas deferens, 
then serving to carry egg capsules (1986:238).] With this 
admittedly small data set for Cy. beauii from only two 
localities, and the absence of data on individual devel- 
opment and longevity, settlement cues and adult mo- 
bility, several scenarios could be constructed. It is, for 
example, theoretically possible that functional males ar- 
rest growth to prolong the male phase, or that the change 
into the female phase occurs only after successful mating 
as a male. However, no evidence was found to consider 
this sex change as labile, i.e., environmentally mediated, 
as is known for other caenogastropod families such as 
Calyptraeidae and Stiliferidae (see Hoagland, 1978, for 
examples and discussion). The data are further insuffi- 
cient to demonstrate either seasonal or partner-induced 
change. The sex change in Cy. beauii appears to occur 
only once, at a predetermined size (figure 53; about 8 
mm shell diameter, 3 teleoconch whorls), and it appar- 
entlv affects most, if not all, individuals (as the general 
shift in gender over time in the Ft. Pierce population 
suggests). 

Thus Cyclostremiscus beauii is interpreted as a pro- 
tandrous sequential hermaphrodite. The term sequential 
(= consecutive, successive) is important here. In a review 
of the terms protandry, protogyny and hermaphroditism, 
Hoagland (1984:86) defined protandry as "the function- 
ing of an organism first as male, then as female, with no 
further sex change. The two sexual phases are separated 
by a phase in which male primary and secondary sex 
characters disappear, and the animal re-differentiates as 
a female." This however omits those species which, after 
an initial male-only phase, have both male and female 



Page 26 



THE NAUTILUS, Vol. 102, No. 1 



reproductive systems developed and tiiiictional (e.g., 
members of Rissoellidac and Omal(>i;\ ridae; see below), 
i.e., protandrous simultaneous hermaphrodites. [Simul- 
taneous hermaphrodites, in our understanding, do not 
necessariiK use "the same gonad to produce both eggs 
and sperm" as defined b% Hoagland (1984:85).] 

Although, at this point, we do not know whether pro- 
tandrous sequential hermaphroditism is the exception, 
rather than the rule, in this family, the occurrence of 
hermaphroditism in Cyclostremiscus beauii could be in- 
terpreted as an adaptation to its peculiar habitat char- 
acterized b\ low densit\ and relative isolation. Protan- 
drous sequential hermaphroditism has an ad\antage for 
Cy. beauii, as each individual thereby minimizes the age 
at which it first reproduces and increases the likelihood 
of finding a compatible mate in a small group. Inbreeding 
between siblings would also be reduced (see Ghiselin, 
1969, for discussion). 

Members of the approximately 25 families grouped 
under Rissoacea are general!) described as gonochoristic 
(see, e.g.. Boss, 1982:984), although possibly derived from 
ancestors that were sequential hermaphrodites (Slavo- 
shevskaya, 1984). Reported cases of hermaphroditism in 
prosobranchs (e.g., Webber, 1977:10; Fretter, 1984:15) 
include onl\ two genera that have classicalK been as- 
signed to this superfamily, Omalogyra (Omalogyridae) 
and Rissoella (Rissoellidac). Fretter (1948) described in 
detail the anatomy and reproductive biology of O. ato- 
muilPhilippi. 184'l)andfi. Jiap/!a(!a(Alder,'l848). Both 
species were found to be protandrous simultaneous her- 
maphrodites, with some likelihood of self-fertilization in 
O. atomus (1948:612, 621, 630; see also Fretter & Gra- 
ham, 1962:381, and 1978:218, 223). [Fretter & Graham, 
1964:134, refer (erroneously:') to the .same two species as 
being "protandrous con.secutive hermaphrodites. '] Both 
families have been subsequently removed from the Ris- 
soacea and have been recently placed outside the Cae- 
nogastropoda, near (Salvini-Plawen & Haszprunar, 1987) 
or in (Ponder & Waren, in press) the Allogastropoda 
(= Heterostropha of authors). Thus, Cyclostremiscus 
beauii is to our knowledge the only species in the su- 
perfamily Rissoacea for which protandrous sequential 
hermaphroditism has been demonstrated. However, as 
sequential hermaphroditism is not as easily recognized 
as simultaneous hermaphroditism, it might be more 
widely distributed in the N'itrinellidae and/or other fam- 
ilies of this group. The mismatch in descriptions of the 
male reproductive system ol Tornus subcarinatus, where 
a penis was lacking according to Woodward (1898) and 
was found by Graham (1982), might find an explanation 
after all. 

ACKNOWLEDGEMENTS 

For invaluable help in the field and laboratory, we es- 
pecially thank William D ("Woody") Lee (SMSLP), 
w hose enthusiasm, expert technique and keen eye added 
significantly to the progress of this study. Dr. M. G. 
Harasewych (USNM) furnished the .scanning electron 



micrographs for figures 1-3. Dr Richard S. Houbrick 
provided valuable field assistance (he found the largest 
temale, without his glasses). Drs. Klaus Bandel (Univer- 
sitiit Hamburg, West Germany), Donald R. Moore 
(KSM.\S) and Winston F. Ponder (Australian Museum, 
S\dne\ ) offered comments on species identifications and 
larval shells Dr Ra\mond B. Manning (USNM) was first 
to bring the molluscan associates of Lysiosquilla to our 
attention and provided information about the stomato- 
pods. Dr. Robert Robertson (ANSP), Dr. Patrick Nuttall 
and Ms. Kathie Way [British Museum (Natural History)], 
Dr. Terrence M. Gosliner (CAS), Dr. Fred Thompson 
(Florida State Museum, Gainesville), Dr. Kenneth J. Boss 
(MCZ), Dr. Philippe Bouchet (MNHN), Hugh J. Porter 
(UNC-IMS), Dr. M. G. Harasewych (USNM), and Fred- 
erick J. C'ollier (USNM) provided comments on and or 
loans of specimens entrusted to their care. We also thank 
the following: Julianne Piraino (SMSLP) for SEM assis- 
tance; Patricia Linley (HBOI) for advice on histological 
techniques; Tom Smoyer for help with photographic 
printing; Dr. Robert Hershler (USNM), Dr. M. G. Har- 
asewych (USNM), Richard E. Petit (North Myrtle Beach, 
SO, kristen Metzger (HBOI), and Carol Browder (HBOI) 
for help with literature acquisition. Drs. Kevin J. Eck- 
elbarger (HBOI), M. G. Harasewych (USNM), Robert 
Hershler (USNM), and Richard S. Houbrick (USNM) are 
thanked for valuable comments on the manuscript. The 
senior author gratefully acknowledges financial support 
provided by a NATO postdoctoral fellowship, adminis- 
tered b\ the German .Academic Exchange Service 
(D.\AD), and research facilities provided by the Smith- 
sonian Marine Station at Link Port, Ft. Pierce, FL. 

This is Smithsonian Marine Station Contribution No. 
217 and Harbor Branch Oceanographic Institution Con- 
tribution No. 622. 

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THE NAUTILUS 102(1 ):30-.35, 1988 



Page 30 



Geographical Distribution of Some Epitoniidae 
(Mollusca: Gastropoda) Associated with Fungiid Corals 



Helen DuShane 

Research Associate 
Natural History Museum 
Los Angeles Count) 
Los Angeles, CA 90007, USA 



mailing address: 
15012 El Soneto Drive 
Whittier, CA 90605, USA 



ABSTRACT 

The known ranges of three coral associated epitoniids, Epito- 
nium costulalum (Kiener, IS39), Epitonium ulu Pilsbry, 1921, 
and Epitonium hulldtttm (Snwcrln, 1844) are reviewed 



INTRODUCTION 

The family Epitoniidae has had a complicated nomen- 
clatural history, with more than 3,000 Recent and fossils 
species named. The genus Epitonium is an overwhelm- 
ingly large group about which itiucIi has been published, 
but information on habitats, ecology, growth patterns, 
and reproduction is known for relatively few species. 
Some epitoniids are perhaps the only gastropods e.xcept 
certain coraiiiophiiidae to have feeding associations with 
iuiigiid corals. 

The stony corals comprise a large group of animals 
showing a great variety of form. Wells ( 1956:F388) listed 
1 1 living genera in the family Fungiidae Dana, 1846, 
and further stated. "Within the family Fungiidae some 
animals are attached as juveniles, but as adults become 
detached and live free on rubble substrates. Others re- 
main attached throughout their lives." There are 11-12 
genera, within w liich are probably 49 species. Fungiids 
live in warm seas in water temperatures 22 °-33 °C. The 
corallum (skeleton) is rounded or elongate, varying in 
length from about 5 cm to 35 cm, and in width from 5 
cm to 20 cm Known commonly as "hat' or "mushroom " 
corals, living FuiifJia sp|). arc pink, green, off-white, or 
pale tan. Distribution of these corals is extensive, from 
the east coast of Africa, across the Arabian Sea, the Bay 
of Bengal, Western Pacific Ocean and including the 
Southern and Flastern Pacific oceans. 

Previous reports ol epitoniids associated with Fiingia 
are those of Root (1958:8), Robertson (1963:57, 1965:7, 
1970:45), Bosch (1965:267), Taylor (1977:254), Kay (1979: 
152), Bell (1982:508, 1985:161 ). Sabelli and Taviani (1984: 
92). These papers report at least three species of epito- 
niids associated w ith species ol Fungia. 

A collection of Epitonium spp. made by SCUBA divers 
in the northern Red Sea verified that Fungia is "home" 



to the tollowing species of epitoniids: Epitonium cos- 
tulalum (Kiener, 1839), originally described from an 
unknown locality; Epitonium ulu Pilsbr\. 1921, origi- 
nalK described from Hawaii; and Epitonium hullatum 
(Sowerby, 1844), originalK described from the Philip- 
pines. An unidentified Epitonium sp. associated with 
Fungia was also reported by Sabelli and Taviani (1984) 
from the Red Sea. 

The following abbreviations appearing in the text are 
defined as follows: 

ANSP — Academy of Natural Sciences, Philadelphia. P.\. 
GENEVA — Museum d Histoire Naturelle, Geneva, Sw it- 

zerland. 
NHM, L.A. — Natural History Museum. Los Angeles 

County, Los Angeles, CA. 
Bratcher (Collection — Twila Bratcher, Los .Angeles, C.\. 
Chane)' Collection — Henry Chaney, Redondo Beach, CA. 
DuShane Collection — Helen DuShane, Whittier, CA. 
Kaiser Collection — Kirstie Kaiser, Park Citv, UT. 



RECORDS 

Epitonium costulatum (Kiener, 1839) 
(figures 1, 2) 

Discussion: Epitonium costulatum (Kiener, 1839) as 
far as presentK know n lives in warm seas from 10°N to 
SCN, from the Red Sea, Bay of Bengal (India and Thai- 
land), and the Philippine Islands. This is the first report 
of the egg capsules oi E. costulatum found on the un- 
derside ol Fungia sp. from the Red Sea. Each egg capsule 
was encrusted w ilh bits of white coral-sand and attached 
to another by two slender threads (figure 2). The number 
of egg capsules per mass is approximately 100. 

Hoot (1958) was the first to report this species as an 
unidentified eiJitoniid li\ing under Fungia, in the Sulu 
Archipelago, Philippines. His six specimens are at the 
Academy of Natural Sciences of Philadelphia (ANSP 
230639) and were studied b\ Robertson (1963) who later 
(1970) compared them with the holot\pe in CJeneva 
(original locality unknown) and concluded that Root's 
specimens are indeed £. costulatum. The ANSP speci- 



H. DuShane, 1988 



Page 31 





4 

Figure 1. Apertural view of Epitoniitm costulatum (Kiener, 1839). Length 28 mm, width 13 mm, DuShane Collection. Straits of 
Tiraii, Red Sea Figure 2. Coral-sand encrusted egg mass of £. costulatum. Chaney Collection. Straits of Tiran, Red Sea. Figure 
3. Epitonium ulu Pilsbr\, 1921. Length 13.5 mm, width 5.5 mm, Bratcher Collection. Saudi .Arabia, Red Sea. Figure 4. £. ulu 
with egg mass. Length of shell 16 mm, width 11 mm, Chaney Collection. Tiran Island, Straits of Tiran, Red Sea. Figure 5. 
.Apertural view of Epitonium bullatum (Sowerby, 1844). Length 13 mm, width 9 mm, Kaiser Collection. Thomas Reef, Sinai, Red 
Sea. Figure 6. Dorsal view of E. htllatum. Kaiser Collection. 



Page 32 



THE NAUTILUS, Vol. 102, No. 1 



mens have 21-26 costae and range in length from 12 4 
to 35.3 mm (Robertson, 1963, 1970). 

Robertson (1963:60) stated, "There are some wentle- 
traps which seemingly live throughout most ot their post- 
larval lives with (relatively) large sea anemones (or cor- 
als). Such species are . . . £. aff. costulatum ..." "This 
F.pitonitim. the oiiK wentletrap so far found w ith a coe- 
leiiterate other than a sea anemone, presurnabK feeds 
on Fungia." We now know that £. costulatum lays its 
gelatinous egg masses under Fungia on a sandy, rubble 
substrate, in depths of 2-30 m. Feeding observations are 
lacking. 

Subsequent to 1958, other specimens have been col- 
lected from various Indo-Pacific localities. 

Recent records: Ba\ of Bengal, India. One specimen, 
length 31 mm, with 17 costae and 12 whorls. DuShane 
Collection. 

Raya Island, Bay of Bengal, Thailand. One specimen, 
length 34.5 mm, with 23 costae and 1 1 \\ horls. DuShane 
Collection. 

Phuket Island. S\V Thailand, Collected by fishermen 
under "hat" coral, 15-20 m, 5/22/85. Three specimens, 
lengths 23.5, 24.5, 25 mm, with 19, 20, 26 costae and 7, 
9, 12 whorls. DuShane Collection. 

Phuket Island, S\V Thailand. Collected by fishermen 
under "hat" coral, 15-20 m, 5/22/85. One specimen, 
length 23.5 mm, with 20 costae and 10 whorls. NHM, 
L.A. 124505. 

Tiran Island (SW), Straits of Tiran, Red Sea (27°57'N, 
34°32'E). Collected by Henry Chaney, 10/31/85, SCU- 
BA, 2-5 m in rubble under Fungia sp., four specimens, 
three specimens, Chaney Collection, lengths 13, 20, 29 
mm, with 18, 23, 28 costae and 8, 9, 13 whorls; one 
specimen, DuShane Collection, length 28 mm, with 24 
costae and 12 whorls. .\\\ C^haney collected specimens 
were live-taken, two were pink in color, with gelatinous 
egg masses as described above. 

Thomas Reef, Sinai, Red Sea (27°59'N, 34°27'E). Col- 
lected by Kirstie Kaiser, 10/30/85, SCUBA, 6.1 m in 
sand pockets under live, detached Fungia sp., one spec- 
imen, length 23 mm, witli 21 costae and 11 whorls, water 
temperature 25 °C, live shell attached to coral. Kaiser 
Collection. 

Little Hiva, Maldive Islands. Collected by Henry Cha- 
ney, 8/31/86, SCUBA, 1 m reef sand, under Fungia 
repanda Dana, 1846, one specimen, live-taken, length 
14 mm, with 20 costae and 11 whorls. DuShane Collec- 
tion. 



Epitonium ulu Pilsbry, 1921 
(figures 3, 4, 7-10) 

Discussion: Epiloniuiit ulu l'ilsbr>, 1921, a widely dis- 
tributed species, lives in warm seas from 30°N to 5°S, 
from the Red Sea, to the Maldive Islands, New Guinea 
and Haw aii. It is specific to certain species of the stony 
coral genus Fungia. Masses of beige colored egg capsules 



are laid on the concave underside of Fungia spp. attached 
to both the coral and to adult Epitonium. 

Bosch (1965) reported that Fungia .scutaria Lamarck, 
1816, from Kaneoke Ba\, Oahu. Hawaii, was infested 
with both eggs and adults of Epitonium ulu. The snails 
had large amounts of pink tissue containing symbiotic 
algal cells and nematoc\sts the\ had ingested. 

Taylor (1977) studied the growth rate of Epitonium 
ulu feeding on the sea anemone Aiptaaia sp. and found 
that the intracapsular development time for this species 
varied from five days to several weeks. She found that 
after hatching the ju\ eniles added approximately 0.2 mm 
of shell length per day. 

Bell (1982) observed that £. ulu produced a "mean of 
32 capsules per day, each capsule containing 500-600 
eggs." She later stated (Bell, 1985) that "embryos com- 
plete intracapsular de\elopment in six da\s at 26-27''C 
and hatch as planktotrophic veligers. ' 

Epitonium ulu has been collected recentK from five 
localities. 

Recent records: Maldive Islands. Robertson (1965) re- 
ported an Epitonium sp. living in association with Fun- 
gia. He now identifies the four specimens as Epitonium 
ulu (personal communication, 1986). 

Papua New Guinea. Collected by Tw ila Bratcher, 1980, 
from under Fungia sp. (figure 3), 11m, water temper- 
ature 28 °(]. length 16 mm, w ith 30 costae and 11 w-horls. 
Bratcher Collection, and one specimen, length 12 mm 
with 30 costae and 12 whorls. DuShane (Collection. 

Tiran Island (SW), Straits of Tiran, Red Sea (27°57'N, 
34°32'E). Collected b\' Henry Chaney. 10/31/85. SCU- 
BA, 2-5 m, with egg mass, under detached Fungia on 
rubble substrate. Five live-taken specimens, all exuding 
mucus, were collected. Four specimens, Chanc\ Collec- 
tion, lengths 3, 4, 5, 16 mm, with 10, 11, 24, 33 costae 
and 9, 10, 12, 12 whorls; one specimen, DuShane Col- 
lection, length 6.5 mm, with 21 costae and 11 whorls, 

Sinafir Island, Saudi Arabia, Red Sea. (Collected b\ 
Twila Bratcher, 10/30/85, SCUBA, 9 m, with eggs under 
detached Fungia. Two live-taken specimens, lengths 9, 
13.5 mm, with 17, 24 costae and 9. 10 whorls. Bratcher 
Collection. 

Thomas Reef, Sinai, Red Sea (27°59'N, 34°29'E). Col- 
lected by Kirstie Kaiser, 10/30/85, SCUBA, 6 m, water 
temperature 25 °C. .\ttached with eggs to underside of 
detached Fungia. 1\ ing in small sand pockets with sand 
and light rubble, l-'our live-taken specimens, lengths 6, 
8, 15, 19 mm, with 17, 20, 23, 32 costae and 10, 10, 13, 
14 whorls, all exuding mucus when collected. Kaiser 
Collection. 

All three S(CUB.'\ divers (Bratcher. Chaney, Kai.ser) 
collected egg masses (figures 4, 8-10) associated with 
specimens of Epitonium ulu from the Red Sea. The eggs 
were off-w liite w ith light purple spots. One egg sac con- 
tained 4()()-()()0 embr\()s in different stages of de\elop- 
ment. Each translucent egg sac w ithin the cluster is pa- 
pillose over the entire external surface. The papillae are 
softly rounded. From each egg sac three transparent 
twisted threads are attached at each end of the oval egg 



H. DuShane, 1988 



Page 33 








8 





10 




.113 mm 



Figure 7. Epitonium ulu Pilsbry, 1921 with egg masses on concave underside of Fungia sp. Figure 8. Egg capsules of £. ulu 
that appear to be empty and transparent. Figure 9. Developing, opaque embryos of £. ulu located on the inner surface of the 
transparent sac Figure 10. Prehatching veligers of £. ulu. Diameter of a single snail 0.113 mm. 



sac, and eventually twist together to form a single knotted 
mass. Each connecting thread appears to be many times 
the length of the sac itself. 

The egg capsules e.xamined under a microscope (300 x ) 
showed the young capsules in three different stages of 
development. Within each capsule development was uni- 
form. The first stage capsules appeared to be empty and 
transparent (figure 8). The second stage showed devel- 
oping, opaque, white embryos that appeared to be lo- 
cated on the inner surface of the sac, but not filling the 
entire compartment (figure 9). The third stage consisted 
of well-developed veligers, thus crowding the capsule 
(figure 10). One capsule in the third stage of development 
contained 400-600 free swimming veligers of equal size 
and development. The diameter of a single veliger was 
0.113 mm. The gibbous shell is transparent e.xcept for 
the darkened area of the columella. In each embrvonic 



shell the animal appeared as a uniform gelatinous mass 
with two dark spots. 

Epitonium (Globiscala) bullatuni (Sowerby, 1844) 
(figures 5, 6) 

Discussion: Epitonium {Globiscala) bullatum (Sower- 
by, 1844) lives in warm seas from 35°N to 30°S, from the 
Red Sea, East Africa, New Guinea, Australia, the Phil- 
ippines, and Japan. Kilburn (1985:330-331) reported this 
variable species from off Southern Mozambique, and in 
addition to living under species of Fungia, "under rocks 
in low tide pools, associated with the actinian Fseudac- 
tinia flagellifera (Hertw.) on w hich it feeds; on occasion 
it ma\ be partly covered by the basal disk of the anem- 
one. Juvenile bullatum shelter among coralline algae, 
probably feeding on the small anemones that are at- 



Page 34 



THE NAUTILUS, Vol. 102, No. 1 



tached to the fronds." He synonymized Scala (Globis- 
cala) papyracea do Boiiry, 1912 from Natal, Epitonium 
(Glolmcala) woolacuttae Kersiake, 1958 from South 
Queensland and GlobUcala kashiuajimensis .\zuma, 1962 
from Japan with £. bullatum (Sosserby, 1844). 

Recent records: Thomas Reef, Sinai, Red Sea (27°59'N, 
.34°27'E), collected by Kirstie Kaiser, 10/30/85, SCUBA, 
6. 1 m, water temperature 25 °C. one live specimen from 
under live, detached Fungia. length 13 mm, with 40 
almost imperceptible costae, 7 whorls, protoconch whorls 
lost, spiral striations weak when viewed under a micro- 
scope (10 X ), umbilicate, color white. A second live spec- 
imens was eaten by a wrasse (Labridae), before it could 
be collected. An egg mass was observed and photo- 
graphed with this species but not collected. 

New Guinea. Three specimens in the DuShane Col- 
lection (ex Withrow Collection), taken in 30 m, lengths 
11, 11.5, 13 mm, with 8, 9, 9 whorls. 

Epitonium sp. 

An unidentified species of £pi7o;!nu>i was reported taken 
from under Ftingia paunwtensis (Stutchbur\-, 1833) off 
Saudi Arabia in the Red Sea b\ Sabelli and Taviani 
(1984). Three specimens in varying growth stages were 
collected with an egg mass at a depth of 2 m. The authors 
described their specimens as umbilicate with a very thin 
shell, 22 obsolete costae and interspaces crossed by fine 
spiral grooves. This is similar to the description of Epi- 
tonium ulu, but comparison with E. ulu is needed. 

The extensive paper by Jousseaume (1911) treating 58 
species of epitoniids from the Red Sea is of little help in 
identifying species from that area. Unfortunately, his 
figures are too small to be of value, and some of his 
descriptions are vague. 

CONCLUSIONS 

With an upper surface exposed to the light and a pro- 
tected concave under surface, species of the genus Fun- 
gia serve as host for several species of epitoniids with 
very delicate shells, thin, fragile walls, and numerous 
costae that are easiK shattered. Whether the wentletraps 
feed on the mucus from the host polyps or the polyps 
themselves is unknown. Taylor (1975) concluded that 
Epitonium ulu can, under laboratory conditions, feed 
on the sea anemone Aiptasia. All other evidence indicates 
that E. ulu deposits eggs only under Fungia. The coral 
is used as the spawning site, the eggs being laid in ge- 
latinous masses, each capsule connected to another by 
two threads. The ability of these epitoniids to locate and 
utilize the underside of F'ungiidae requires further study. 
An increasingly high degree of specificity is being found 
w ith certain epitoniids seemingK having permanent as- 
sociations with Fungia corals. 

ACKNOWLEDGEMENTS 

My thanks to Robert Robertson for encouragement, and 
for reading the manuscript and making useful sugges- 



tions; to William K. Emerson and James H. McLean for 

reading the manuscript and offering advice; to the divers, 
Twila Ikatcher, Henry (^hane\ , and Kirstie Kaiser, who 
returned (rom the Red Sea with epitoniids that formed 
the backbone of this paper; to the personnel of the .\llan 
Hancock Foundation Library, Universit>- of Southern 
California, for help in locating several rare volumes; to 
Da\id Nhilliner and Bertram Draper for the excellent 
photographs; and to Kirstie Kaiser for the drawings. 

LITERATURE CITED 

Azuma, M, 1962. Descriptions of five new species of Japanese 
Epitoniidae. Venus 22(2);130-136, text figs. 1-6. 

Bell, J. L. 1982. Larval development and metamorphosis of 
the prosobranch mollusc, Epitonium ulu, associated with 
a solitary coral. Pacific Science 16:508. 

Bell, J. L. 1985. Larval growth and metamorphosis of a proso- 
branch gastropod associated with a solitar\ coral. Pro- 
ceedings of the Fifth International Choral Reef Congress, 
Tahiti 5:159-164, 6 figs. 

Bosch, H. F. 1965. .\ gastropod parasite of solitary corals in 
Hawaii. Pacific Science 19:267-268, 1 fig. 

Boury, E. de. 1912. Description de Scalidae nouveaux ou peu 
connus. Journal de Conch\ liologie 60t2):87-108, 1 pi. 

Dana, J. D. 1846. Zoophytes. United States Exploring Ex- 
pedition during the vears 1838-1842 under the command 
of Charles Wilkes, L .S.N., Vol. 7, p. 1-741 Philadelphia. 

Dana, J. D. 1849. Descriptions of fossils. United States Ex- 
ploring Expedition, 1838-1842, under the command of 
Charles Wilkes, U.S.N. Geology. By James D. Dana. \'ol. 
X, .Appendix, p. 681-729, pars. Atlas, xxi, pis. pars. Phila- 
delphia, 1849. 

Jousseaume, F. P. 1911[1912]. Faune malacologique de la 
Mer Rouge. Memoires de la Societe Zoologique de France 
24:180-246, pis. .5-7. 

Kay, E. A. 1979. Hawaiian marine shells Reef and shore 
fauna of Hawaii. Section 4: Mollusca. Bernice P. Bishop 
Museum Special Publication 64(4). Bishop Museum Press, 
Honolulu, Hawaii, 653 p., 2 figs. 

Kersiake, J. 1958. A new Epitonium from eastern .Australia. 
Proceedings of the Royal Zoological Society of New South 
Wales for 1956-57, p.' 157-158, 1 fig. 

Kiener, L. 1838-39. Species general et iconographie des co- 
quilles vivantes; Genre Scalaire. Rousseau, Paris 7:1-22, 
pis. 1-7 (text: 1839; plates: 1838). 

Kilburn, R. N. 1985. The family Epitoniidae (Mollusca: Gas- 
tropoda) in southern Africa and Mozambique, Annals of 
the Natal Museum 27(l):239-337, 171 figs. 

Lamarck, de M. de. 1816. Histoire naturelle des animaux sans 
vertebres, Paris 2:1-568. 

Pilsbry, H. A. 1921. Marine mollusks of Hawaii. Proceedings 
of the Academy of Natural Sciences of Philadelphia 72: 
360-382. 

Quelch, J. J. 1886, Scientific results of the vo\age of Chal- 
lenger. Report on the reef corals collected b\ H.M.S. Chal- 
lenger during the years 1873-76. London. Zoology 16:1- 
203. 

Robertson, R. 1963. Wentletraps (Epitoniidae) feeding on sea 
anemones and corals. Proceedings of the Malacological 
SocietN of London 35(2&3):51-63, pis. 5-7. 

Robertson, R 1965 Coelenterate-associated prosobranch gas- 
tropods .•Viinual Reports of the .American Malacological 
Union for 1965, 32:6-8. 



H. DuShane, 1988 



Page 35 



Robertson, R. 1970. Review of the predators and parasites of 
stony corals, with special reference to symbiotic proso- 
branch gastropods. Pacific Science 24(l):43-54. 

Root, J. 1958. Rapa rapa in the Snlu Sea. Hawaiian Shell 
News 7:7-8, 

Sabelli, B. and Marco Taviani. 
Fungia-associated epitonid 
20(l-4):91-94, 

Sowerby, G. B., II. 1844 (July) 

of Scalaria, collected by Mr. H. Cuming, to be figured in 
the fourth part of Thesaurus Conchyliorum. Proceedings 
of the Zoological Societ) of London, pt 12:1-38. 

Stuchbury, T. 1833. .\\\ account of the mode of growth of 
young corals of the genus Fungia. Transactions of the 
Linnean Society of London 16:493-498, pi. 32, figs. 6a, b. 



1984 Red Sea record of a 
sic]. Bollettino Malacologico 

Descriptions of new species 



Taylor, J B 1975 Planktonic prosobranch veligers of Ka- 
neolie Bay Pli D. dissertation, I'niversity of Hawaii, Ho- 
nolulu, 599 p. 

Taylor, J. R. 1977. Growth rate in juvenile carnivorous proso- 
branchs (Mollusca: Gastropoda) of Kaneohe Bay, Oahu, 
Hawaii. Proceedings Third International Coral Reef Sym- 
posium Rosenstiel School ut Marine and Atmosplieric Sci- 
ence, University of Miami, Miami, Florida 3(l):253-259, 
8 figs. 

Wells, J. W. 1956. Treatise on invertebrate paleontology. Part 
F. Coelenterata. Geological Society of America and Uni- 
versity of Kansas Press, xvii -I- 498 p., 2,700 figs. 



THE NAL TILLS 102(1 ):36-.39, 1988 



Page 36 



A New Species of Vasum (Gastropoda: Turbinellidae) 
from off Somalia 



William K. Emerson 
Waller E. Sage, III 

Department of ln\ertebrates 
American Museum of Natural History 
New York, NY 10024-5192, USA 



ABSTRACT 

Vasum stephanti new species is described from moderately 
deep water off Cape Gardafui (Ras Asir), Somalia and com- 
pared with congeners. 



INTRODUCTION 

In the past few years, deep-sea commercial fisheries op- 
erations off the northeastern coast of SomaUa have re- 
sulted in the discovery of several new or otherwise in- 
teresting species of moilusks. Lorenz (1987:11) described 
P.seudosiiunia wieseonim new species from the region 
off (^ape Ras Hafun in about 300 m, and recorded the 
presence oi Festilyria j estiva (Lamarck, 1811), S(romi>us 
oldi Emerson, 1965, and Cijpraea broderipii Sowerby, 
1832, from the trawl samples. Waller (1986:39-46) de- 
scribed Somalipecten crannwrorurn new genus, new 
species, from "off Somalia, depth 150-300 m ", obtained 
from Taiwanese fishermen, and also provided a list of 
associated species. Another species recently described 
from off Somalia is Vulutocorbis rosavittoriae Rehder, 
1981. 

Through the good offices ot John Bernard, our atten- 
tion was called to an umiamed species of Vasum trawled 
in Somalian waters. We take pleasure in describing this 
new species in honor of Adolphe Stephant, w ho obtained 
the specimens from Danish shrimpers and generously 
provided Mr. Bernard with the data and specimens. Oth- 
er species of moilusks reported by Mr. Stephant (in Hit., 
August 19, 1987) to have been taken during these trawl- 
ing operations include h'cstihjria jcstiva, titwrnbits oldi, 
S. piicatus (Roding, 1798), Phalium microstoma (von 
Martens, 1901), P. bituberadosnm (von Martens, 1903), 
Ficus investigatoris (E. A. Smith, 1894), Cytnatium ran- 



zanii (Bianconi, 1851), Bufonaria fernandesi Beu, 1977, 
Vasum crosseanum (Souserbie, 1875), Tudicula zartzi- 
barica Abbott, 1958, Metula bosu:ellae Kilburn, 1975, 
CucuUaca labiata (Lightfoot, 1786), and Chlamys tonn- 
sendi (Sowerb>, 1895). 

SYSTEMATICS 

Family Turbinellidae Swainson, 1840 
Subfamily Vasinae H. & .\. .\dams, 1853 

Genus Vasum Roding, 1798 

Remarks: See Abbott (1959) and \okes (1966) for re- 
views of this subfamily. 

Vasum stephanti new species 
(figures 1-6) 

Diagnosis: Similar to Vasum tubiferum (.-^nton, 1839) 
in general appearance, but differs in having a more tri- 
angular outline, three equalK well-de\ eloped columellar 
plaits (in place of three major, plus one or two minor 
plaits), a nearly uniformly milk-w hite shell w ith a white, 
glazed aperture and parietal wall (compared to an or- 
ange-brown to yellowish shell with the parietal wall a 
light tan w ith \er\ large splotches of chestnut to purple- 
brown), and in the presence of two or three rows of spines 
at base of shell (instead of one row). 

Description: Shell moderately large for genus, attaining 
108-f- mm in length. Solid, heavy, turbinate, and strongly 
spined. Spire ele\ated with a short, smooth, bulbous nu- 
cleus of 1'2 whorls (figures 5, 6). Postnuclear whorls 7 
(adult specimens lack complete spires), the body whorl 
with 7 to 8 well-developed, curved to strongly recurved, 
flaringly grooved and terminally open, subsutural spines. 



Figures 1-6. V'a.sinn stephanti new species. 1, 2. Paratype, AMNll 225988. 3, 4. Holot\pe, AMNH 225987. 5, 6. Paratype, 
AMNH 225989 (details of spire). All from type locality: off Cape Gardafui, Somalia, see text; figures 1-4 approximately x %. 
figures 5, 6, x 2. 



W. K. Emerson and W. E. Sage, III, 1988 



Page 37 




Page 38 



THE NAUTILUS, Vol. 102, No. 1 



A row of similar but much shorter spines below the first 
row, followed by 5 coarse spiral cords and intervening 
raised lines. Base of shell with 2 to 3 spiral rows of 
moderateK developed, groove spines and lower surface 
with weakK developed spiral lines. Parietal wall thick- 
ened, slightK raised, glazed. (Columella with 3 plicae, 
posterior 2 better developed; first posterior plica semi- 
bifid in 1 specimen. Outer lip moderateK thin, slightK' 
reflected, crenulated. Umbilicus funnel-shaped and in 
most specimens widely open. Base color of shell milky 
white, spire stained buff. CJolumella glazed. Aperture 
white, with a slight bluish tinge. Periostracum moder- 
ateK thick, tannish brown, and somewhat foliaceous. Soft 
parts not seen. Operculum brown, corneous, unguiculate, 
apicalK nucleate, filling most of the aperture with foot 
fulK u ithdrawn. Inner surface marginally thickened on 
basal and abcolumellar sides, central area depressed and 
with irregular concentric rings; outer surface scabrous. 

Type locality: 13-16 km east, 80-96 km south of Cape 
Gardafui (Ras Asir), Somalia, trawled by shrimp fisher- 
man in 183 to 220 m, December, 1986. 

Range: Known only from the type localit\ and in the 
Gulf of .'Kden off the Bari coast of Somalia. 

Material examined: Holotype, AMNH 225987, 102.36 
mm, ex E. Schelling Collection (figures 3, 4); paratype 
2, AMNH 225988 (figures 1, 2), paratype 5, AMNH 
225989 (figures 5, 6), paratype 8, AMNH 225990, ex J. 
Bernard Collection; paratypes 1, 3, 4, 7, 9 J. Bernard 
Collection; paratypes 6, 10 A. Stephant Collection, all 
from the t\pe locality; referred specimen, H. Lee Col- 
lection, Alula, Bari coast, Somalia (.see Table 1). 

Remarks: As noted above, Vasum stephanti new species 
most closely resembles in shell morphology the endemic 
Philippine (Cuyo-Palawan group) species V. tubiferum 
(Anton, 1839:70; Kobelt, 1876:155, pi. 9, fig. 3; Abbott, 
1959:20, pi. 4, fig. 1; Spring.steen & Leobrera, 1986:105, 
pi. 28, fig. 6). Anton's taxon and the closeK related V. 
turbinellus (Linne, 1758:750; Abbott, 1959:17, pi. 1, figs. 
2, 3), which ranges from East Africa to the western Pa- 
cific, are inhabitants of shallow water, as are the other 
four Indo-Pacific species assigned to Vasum {sensu stric- 
to) by .Abbott (1959). One of these, V. rhinoceros (Gme- 
lin, 1791 ), from Kenya and Tanzania, is somewhat similar 
but has a lower-spired, heavier shell with massive nod- 
ules, thickened and reflected outer lip, and a brown- 
blotched to light yellow parietal wall (Abbott, 1959:21, 
pi. 4, figs. 3, 4). Strongly spino.se specimens with im- 
mature outer lips of the Brasilian V. cassiforme (Kiener, 
1840), cited from low tide to 60 m (Rios, 1985:115), are 
superficially similar to the new species. (See Abbott & 
Dance, 1982:209, 210 for polychrome illustrations of these 
taxa.) The Australian Altivasurn Hedle\, 1914, and sev- 
eral species of Indo-Pacific Tudicula H. & A. Adams, 
1864 are known to occur in moderate depths (to 220 m). 
Some of the spinose species originalK assigned to Tu- 
dicula (e.g., T. zanziharica Abbott, 1958, Iron) the west- 
ern Indian Ocean, and T. rasilistorna Abbott, 1959), from 



Table 1. Vasum stephanti new species. Shell measurements 
in mm and number of whorls; width measured including spines, 
n = 12. Spires incomplete e.xcept for 5, 10 



l.rliUtl] 



Wiiltli 



= \\1 



Paratype 1 
Parat) pe 2 
Holotype 
Paratype 3 
Parat) pe 4 
Paratype 5 
Paratype 6 
Paratype 7 
Paratype 8 
Paratype 9 
Referred 
Parat\|)c II) 



107 68 


106.75 


102.36 


97.72 


97.10 


95.53 


86.22 


78.75 


73.68 


66.87 


63.47 


56,92 



97 SS 
97.11 
82.70 
82.08 
84.19 
77.93 
73.47 
64.10 
57.18 
67.25 
52.78 
,55 79 



6 

6 

6 

6 

6 

8'/4 

6 

5 

5 

5 

5 

4'/4 



off Queensland, .\ustralia, may prove to be referable to 
Vasum when the soft anatomy is known. The weakK 
spinose V. crosseanum (Souverbie, 1875), from the In- 
dian Ocean, appears to be closely related to T. rasili- 
storna. The development of long spines in the species 
described herein may reflect the deeper-water habitat. 

ACKNOWLEDGEMENTS 

We are grateful to Edward T. Schelling, Shalimar, Flor- 
ida, and John H. Bernard, Crossville, Tennessee, respec- 
tively, for generously donating the holotype and two 
purat) pes to the American Museum of Natural Histor\-. 
John H. Bernard, Harr\ G. Lee, Jacksonville, Florida, 
and Adolphe Stephant, Lorient. France, kindK lent spec- 
imens. We thank each of these collectors for pro\ iding 
useful information. R. Tucker Abbott and M. G. Hara- 
sewych oftered helpful suggestions and critically re- 
viewed the manuscript. Stephen Butler, AMNH, con- 
tributed the photographs. 

LITERATURE CITED 

Abbott, R, T. 1958. .\ new Recent species of Tudicula from 
Zanzibar (Gastropoda: X'asidae). Notulae Naturae of the 
Academy of Natural Sciences of Philadelphia 305:1-4. 

.\bbott, R. T. 1959. The family Vasidae in the Indo-Pacific. 
Indo-Pacific MoUusca 1(1): 15-32. 

Abbott, R, T. and S P. Dance. 1982. Compendium of sea- 
shells, E, P. Dutton, New York, ix -(- 411 p. 

.•\dams, H and .-^ ,\dams. 1864. Descriptions of new species 
of shells chiefl) from the Cumingian collection. Proceed- 
ings of the Zoological Society ot London, for 1863:428- 
435 (published April, 1864). 

Anton, H. E. 1839. Verzeichniss der Concliylien welch sicli 
in der Sammlung von Herrmann Eduard Anton befinden. 
Halle, xvi -I- llOp 

Gmelin, J. F. 1791. Caroli a Linne Systema naturae per regna 
tria naturae, 13th ed Leipzig, Vol. 1, pt. 6, cl. 6. Wrmes, 
p. 3021-3910. 

Hedley, C. 1914 Report on the MoUusca obtained 1)\ the 
F.I.S. "Endeavour " from the Great .\ustralian Bight and 
from north and south of Gabo Island. Biological results of 
F".I.S. "Endeavour ". Sydney, 2:65-74. 



W. K. Emerson and W. E. Sage, III, 1988 



Page 39 



kiener, L. C;. 1840[-41]. Species general et icoiiographie des 
coquilles vivantes. . . . Paris, Genre Turbinelle 6(59-71): 
1-50, 1841; pis. 1-21, 1840 (Vasum cassijormc (Kiener) 
dates from the citation on plate 9 of "Turhinclla ca.ssi- 
formis \'alene." [= Valenciennes]; the text was pnhlislied 
in 1841). 

Kobelt, H. C 1876. Systematisches Conchylien-Cabinet von 
Martini und Chemnitz. ... Niirnberg. Purpur.schnecken, 
Band 3, Abt. 3a, pt. 251:121-164. 

Linne. C, von. 1758. Systema naturae per regna tria naturae, 
10th ed. Stockholm, \ol. 1, Regnum animale, 284 p. 

Lorenz, F , Jr. 1987 Description of a new Ovulidae species 
from Somalia (Gastropoda: Ovulidae). La Conchiglia, Rome 
19(214-215):11-12. 



Rios, E. C. 1985. Seashells of Brazil. Fundagao Sociedade do 

Rio Grande, Funda^ao Universidade do Rio Grande, Mu- 

seo Oceanografico. 329 p., 102 pis. 
Souverbie, S. M. 1875. Description d une espece nouvelle 

appartenant au genre Titrhinella. Jonrnal de (>'onchylio- 

logie 23(4):297-298. 
Springsteen, F. J. and F. M. Leobrera. 1986. Shells of the 

Philippines. Carfel Seashell Museum, Manila, 377 p. 
Vokes, E. H. 1966. The genus Va,si(ni (Mollusca: Gastropoda) 

in the New World Tulane Studies in Geology 5(l):l-36. 
Waller, T. R. 1986. A new genus and species of scallop (Bi- 

valvia: Pectinidae) from off Somalia and the definition of 

a new tribe Decatopectinini. The Nautilus 100(2):39-46. 



THE NAUTILUS 102(1 ):40-45, 1988 



Page 40 



Density, Spatial Distribution, Activity Patterns, and 

Biomass of the Land Snail, Geophorus bothropoma Moellendorff 

(Prosobranchia: Helicinidae) 



Kurt Auffenberg 
Troy Auffenberg 

Malacology Division 
Florida State Museum 
University of Florida 
Gainesville, FL 32611 



ABSTRACT 

The results of preliminary work done in the Republic of the 
Philippines during July, 1981 on the density, spatial distribu- 
tion, activity patterns, and biomass of the Philippine Island 
land snail Geophorus bothropoma (Prosobranchia, Helicinidae) 
are presented. 

Density was estimated at 0.55/m^ utilizing the quadrat meth- 
od. Using the parameters of mean crowding and patchiness, 
the species was shown to be uniformly distributed spatially, 
possibly due to mutual repulsion. Individual activity distance 
per 24 hour period varied greatly (0.0-275.0 cm) and was 
possibly correlated with rainfall Utilizing the convex polygon 
and ellipsoid methods, minimum individual activity ranges for 
the duration of the study were estimated to be 2.4-4.2 m^. 
Biomass (live weight) of adults was estimated at 0.13 g/m'^. 
The species is diurnal and does not home. 

Key words: Gastropoda; Prosobranchia; Helicinidae; Geopho- 
rus; density; spatial distribution; activity; biomass; Philippine 
Islands. 



INTRODUCTION 

There are relatively few published studies dealing with 
the density, spatial distribution, activity patterns, or bio- 
mass of land snails (see Discussion section for citations). 
Of those that do exist, almost all have dealt with pul- 
monates of temperate regions, while tropical terrestrial 
prosobranchs remain virtually unstudied. The present 
paper reports on the results of a preliminary study con- 
ducted July 7-17, 1981 on Geophorus bothropoma Moel- 
lendorff, 1895 (Quadras & Moellendorff, 1895:148; Wag- 
ner, 1908:152, pi. 29, figs. 20-23), a large (13.0-16.0 mm 
in diameter), limestone rock-dwelling helicinid endemic 
to the Caramoan Peninsula, Luzon, This represents one 
of the very few times quantitative ecological methods 
have been utilized for the study of terrestrial mollusks 
in all of Southeast Asia. In addition, the present paper 
introduces to ecological malacology several analyses used 
by zoologists working in groups other than mollusks (i.e., 



birds, Odum & Kuenzler, 1955; frogs, Turner. 1960; in- 
sects, Alexander, 1961, Iwao, 1970; mammals, Stumpf & 
Mohr, 1962). While the database with which we worked 
was rather small, the ease ot anaKses and the applica- 
bility of the results suggest that such approaches to sim- 
ilar questions in terrestrial snail ecolog) would be highly 
beneficial. 



STUDY AREA 

The study area is located in the Republic of the Phil- 
ippines, Luzon Island, Camarines Sur Province. Cara- 
moan Municipality, 1.0 km south of Barrio Ilawod. The 
work was conducted 10 m off the trail between Ilawod 
and Gota Beach on a large (8 x 9 x 3 m high) limestone 
boulder with almost vertical sides. 

Much of the Caramoan Peninsula consists of well- 
developed limestone karst of Miocene reef origin. Be- 
cause the area has been relativeK recently uplifted, the 
mountains are rather steep. Hills and ridges often consist 
of masses of large limestone boulders with chasms tens 
of meters deep. Runoff is rapid and though rainfall may 
be high seasonally, the surface environment is often rel- 
atively dry. Surface water collects in solution pits on 
limestone and in hollows of trees. 

Due to the rugged topography, the area remains blan- 
keted in a multi-canopied, tropical evergreen forest. 
Vegetational communities are extremely diverse. The 
forest is dominated by tall dipterocarp trees in both upper 
(30-40 m) and lower (5-20 m) canopies; a dense her- 
baceous layer of mainK ferns exists near the ground 
surface. 

The Caramoan Peninsula has no definite dry season. 
While it tends to rain throughout the \ear (ca. 2,900 
mm), most rainfall occurs from September through Jan- 
uary, with a minor peak in July associated with the onset 
of the typhoon season. 

As in other tropical rainforests, the Caramoan Pen- 
insula shows little seasonal temperature variation. Mean 



K. Auffeiiberg and T. Auffenberg, 1988 



Page 41 



iluil\ temperature fluctuations are greater tPian mean 
seasonal variation. However, there is a marked vertical 
tliermocline within the forest, in which temperatures 
from the surface to 1 m are most stable; daily variation 
increases at greater heights above the ground. Due to 
the dense vegetation at the study site, the effects of solar 
railiation and wind are minimal. Therefore, Geophorus 
butliropoma lives in the most stable part of the local 
warm, moist tropical forest environment. The above eco- 
logical data is from Auffenberg (in press). 

METHODS 

An 8 m- (2 m tall, 4 m wide) vertical grid of 1 m- quadrats 
was constructed against the east side of the boulder to 
measure density and activity. All adult Geophorus both- 
ropoma within this grid (N = 14) were numbered with 
small dots of fingernail polish on the dorsal surface of 
the shell. This increased shell visibilit\ , but is not believed 
to have increased predation due to the nocturnal habits 
of most potential predators. No additional individuals 
were located during subsequent visits. The site was vis- 
ited nine times during the next 11 days at 10 a.m. for 
appro.ximateK 1 hour observations. In addition, the site 
was visited sporadically throughout the daylight hours 
and seven times at night (8-10 p.m.). This provided a set 
of successive censuses, not a random quadrat sampling. 

Densit\ estimates are based on the ciuadrat sampling 
method. This method simply involves counting individ- 
uals occurring within a quadrat of fixed size. 

Analyses of spatial distribution are based on m* (mean 
crowding) and m* m (patchiness) of Lloyd (1967). Mean 
crowding is given by the formula: 



q 



- 1 



(1) 



where x, = the number of individuals in the jth quadrat 
and q = the total number of quadrats. 

Mean crowding is defined as the mean number of other 
individuals per quadrat per individual. It expresses the 
degree of spatial crowding experienced by an individual 
because of others of the same species (or different species; 
see Lloyd, 1967). A particularly important feature of this 
statistic is that it is relatively independent of spatial dis- 
tribution t\ pe, number of samples, and size of the means. 
The most complete discussion concerning the utility of 
this statistic is by Iwao (1977). It should be stressed that 
this statistic does not measure crowding in an ecological 
sense, as man\' variables (i.e., food availabilit\', territo- 
rialit) ) must be examined before this can be determined 
(Lloyd, 1967). 

Patchiness (m* /m) is the ratio of mean crowding to 
mean densit\ (Iwao, 1968). This index provides a relative 
measure of aggregation. It equals unity (1) in random 
distribution, is greater than unity in contagious distri- 
butions and is less than unity in regular (uniform) dis- 
tributions (Iwao, 1977). 



Anal) sis of the activity of Geophorus buthropoma was 
made by subdi\idiiig the I m- quadrats, plotting the 
exact location of individuals and measuring straight line 
distances (nearest cm) from the last observation with a 
flexible tape. This provided a crude estimate of the min- 
imum distance tra\eled since the last ob.servation. Cal- 
culations are based only on data provided by two or more 
consecutive daily observations (N = 27). 

Estimations of activity ranges (N = 5) are based on 
the convex polygon method and the ellipsoid method, 
which is based on a covariance matrix of capture loci 
(Jeunrich & Turner, 1969). The convex polygon method 
involves plotting all the recapture points, drawing the 
smallest convex polygon containing these points and then 
determining the area. This method, defined by the equa- 
tion: 



A = 



x.y^+i - Xi+iYi 



(2) 



is relatively simple but is biased b\ sample size. The 
values tend to increase as the number of capture points 
increases. 

The ellipsoid method is not biased by sample size and 
assumes that individuals are active outside the observed 
range of activity (see Jennrich & Turner, 1969, for dis- 
cussion). This method is defined by; 



67r|S| 



(3) 



where |S| is the determinant of the capture point co- 
variance matrix; 



S = 



defined by the equations; 



S S 

s s 



(4) 



1 



x;^ 



S„ 



n - 2 



o 2 (y. - y)^ 



S. = S,. = — !— 2 (X, - x)(y, 
n — 2 " 

X - - 7v X, , 

y = - 2j y. 



(5) 



Live adult snails (N = 90) were collected near the 
study site and their weights (in groups of 10 individuals) 
were taken to the nearest milligram on a single beam, 
double pan scale. Fifty specimens were then macerated; 
the shells and opercula were rinsed, dried for several 
days and reweighed. 

Voucher specimens of Geophorous bothropoma are 
housed in the mollusk collection of the Florida State 
Museum (UF 56667). 



Page 42 



THE NAUTILUS, Vol. 102, No. 1 



Table 1. Densit\ and spatial distribution parameters based on quadrat method, n = individuals recaptured per visit, m/m- 
(lciisil\, m* = mean crowdiiij;, m* m = patchiness. Visit 6 is (irnitted from densit\ and spatial distribution analyses 













Visits 










X 


SE 






1 


2 


3 


4 


.5 


6 


7 


8 


9 


SD 


n 


9 


6 


5 


5 


2 





1 


3 


4 


3.89 








m/m" 


1.13 


0.75 


0.63 


0.63 


0.25 


— 


0.13 


0.38 


0.50 


0.55 


0.11 


0.32 


m* 


1.11 


1.00 


40 


1 20 


00 


— 


00 


67 


0.00 


0.55 


018 


0.52 


m*/m 


0.98 


1.33 


6-1 


1 90 


(100 


— 


1 ) 1 )( 1 


1.76 


0,00 


83 


0.28 


0.79 



RESUl/rS 

Density and spatial distribution: Froiumnct'd \ariatiun 

in population density was observed during this study 
(table 1). Densities were calculated in eight quadrats for 
nine visits (N = 72 quadrats). \'isit 6 yielded no recap- 
tures and is omitted from density and spatial distribution 
analyses. Depending on the individual visit analyzed, 
mean crowding and patchiness reveal either random, 
contagious, or uniform spatial distributions. However, 
the means of these statistics suggest that individuals are 
uniformly distributed, for the averages of both mean 
crowding and patchiness are less than unity. 

The regression of mean crowding on mean density 
(figure 1 ) also suggests a uniform spatial distribution (« = 
y intercept = —0.14, /3 = slope = 1.25) with possible 
"mutual repulsion" (see Iwao, 1968, for discussion of 
regression analysis). 

Activity: Marked individuals were observed to be active 
during all da\ light hours. None were active during visits 
alter dark. .\ typical activit\' sequence consisted of an 
individual moving along a meandering path for several 
minutes, sometimes feeding on algal growth, stopping 
for several minutes, then continuing. Some individuals 




1.8 2.0 



remained at the same exact location tor 1 or 2 da\s, while 
for no apparent reason others remained active. 

Fifty-seven recaptures were recorded (50.9^ of pos- 
sible recaptures). Distances traveled per day varied greatly 
between individuals (0.0-275.0 cm). Because distances 
traveled rarely conform to normal distributions only the 
range of distances is given. 

Population activity variation was possiblv correlated 
with rainfall. Light rain did not initiate activity in in- 
active individuals, nor did it seem to affect individuals 
already active. However, snails ceased moving about if 
the rainfall became intense. Above average activity levels 
were observed between July 8 and 11 (50.0-275.0 cm, 
N = 11 recaptures, three visits) following intense rainfall 
associated with two small typhoons in the area. This in 
turn was followed by 5 days (July 12-16) of relatively 
little rainfall and reduced snail activity (0.0-65.0 cm, 
N = 10 recaptures, four visits). A heavy rain fell during 
the night of July 16, resulting in a marked increase in 
snail activity on July 17 (65.0-203.0 cm, N = 6 recap- 
tures, one visit). 

Perhaps the most interesting observation concerning 
the activity of this species is that it does not home. In- 
dividuals simply wander from one resting site to another, 
usually shallow depressions or crevices in the rock. They 
avoid resting sites in leaf-filled solution pits, which are, 
however, usually inhabited bv Cijclophorus ceratodes 
Moellendorff, 1895, Japonia ciliata (Sowerby, 1843), ]. 
stephanophora (Moellendorff, 1895), and several small 
pulmonate species (personal observation). 











2- 




,^ 








j 


%^ 






1 ' ^ 


\ 

\ 


1- 


4=- 


■:::y^cT^^--- 


_...^.....,;» 




r^ a 


1 .-- 





Fipurp I . Rejiression of mean density (ni, nr) on mean crowd- 
ing (ni*) of Gi'(>i)li(inis hothropoma in study area. Curved lines 
represent 95% confidence limits. 



12 3 4 

Figure 2. Five representative activity ranges of Geophorus 

bothropojna plotted on 8 m- grid HIack dots represent recap- 
ture sites. Each polygon depicts an activity range of an indi- 
vidual snail ov er an 1 1 day period. 



K. Auffenberg and T. Auffenberg, 1988 



Page 43 



Estimates of individual activity ranges (N = 5) (figure 
2) are quite consistent. The convex polygon method (cor- 
rected for sample size bias; Jennricli & Turner, 1969) 
yielded activity ranges of 2.3-3.8 m- (x = 3.1 ± 0.3, 
SD = 0.5 m-). the ellipsoid method yielded slightly (but 
insignificantly), larger area estimates of 2.7-4.2 m- (x = 
3.3 ± 0.3, SD = 0.6 m-). 

Biomass: Mean total live weight per adult individual 
in the study area was 0.59 g (N = 90). Mean shell weight 
(including operculum) per individual was 0.35 g (N = 
52), yielding a mean live biomass per individual of 0.24 
g. The mean density estimated by the quadrat method 
(table 1) of this species in this area is 0.55 individuals/ 
m-. Thus the estimated biomass is 0.13 g/m-. 



DISCUSSION 

Density and spatial distribution: Previous studies on 
species comparable in size to Geophorus bothropoma 
have been few (i.e.. Mason, 1970; Richardson, 1975; 
Cameron, 1982), and none concerns terrestrial proso- 
branchs. The only prosobranchs with available data are 
the small hydrocenid Georissa monterosatiana (Godwin- 
.•\usten & Nevill, 1879) and the diplommatinid Opis- 
thostoma retrovertens Tomlin, 1938 of Malaysia. Berry 
(1966) found these species occurring in higher densities 
on moss covered rocks than on moss-free rocks, but did 
not address spatial distribution. In the Caramoan area 
we found Geori.'isa rufescens (Moellendorff, 1887) in ex- 
tremely localized and highly aggregated populations on 
vertical rock faces. This is very different from the large 
Geophorus bothropoma, which is found on virtually every 
vertical rock face and is uniformly distributed spatially. 

In this stud\-, intersample variation in estimated mean 
densities is probabK largely due to the limited size of 
the study area. The choice of study area size and number 
of samples must have some biological basis to estimate 
population density and spatial distribution adequately. 
This is usually done by preliminary sampling or guessing 
(Ivvao, 1977). However, we do feel confident that the 
proper quadrat size (1 m^) was chosen because the ob- 
served dail\ activity was approximately 1 m (x = 93.7 ± 
13.31 cm). Density estimates were possibly also biased 
by the rugged microhabitat. The limestone karst provides 
man> cracks, crevices, and solution pits in which a snail 
could remain unseen, despite our thorough searches. 

Utilizing Iwao's (1968) interpretation of m*/m regres- 
sions, we determined that individuals of Geophorus both- 
ropoma were uniformK distributed and may have ex- 
hibited "mutual repulsion. Infra-specific competition 
has never been properly substantiated in studies on ter- 
restrial mollusks and we cannot substantiate it here. 
Nevertheless, density was relatively low, despite the 
seemingly adequate food and resting site resources that 
could have been expected to sustain substantialK' higher 
snail densities. 

.Activity: Previous studies have shown terrestrial mol- 
lusks to be very cyclic in their activity patterns (i.e.. 



Barnes & Weil, 1944, 1945; Wells, 1944; Dainton, 1954a,b; 
Blinn, 1963; Henne, 1963; Cameron, 1970; Crawford- 
Sidebotham, 1972; Baker, 1973; Bailey, 1975; Shachak 
et al., 1975; Heatwole & Heatwole, 1978; Deisler, 1987). 
It is apparent from these studies that the initiation and 
continuation of activity is a variable response to complex, 
simultaneous climatic changes (some minute, perhaps 
immeasurable) in the microenvironment. More field and 
laboratory studies must be undertaken before this will 
be understood. In previous studies snail movements were 
most numerous shortly before or after nightfall and short- 
ly before dawn. However, Geophorus bothropoma is 
diurnal rather than crepuscular, and the congener G. 
trochiformis (Sowerby, 1842) was also observed active 
throughout the day. We believe this behavioral pattern 
is not a general trend in tropical terrestrial prosobranchs. 
Cyclophorus ceratodes Moellendorff, 1895 and Japonia 
ciliata (Sowerby, 1843) and J. stephaiiophora (Moellen- 
dorff, 1895) living in the same study area are decidedly 
crepuscular in their activity patterns (personal observa- 
tion). 

Few data are available on activity and home site ranges 
of terrestrial mollusks. Heatwole and Heatwole (1978) 
found individual home site ranges of the camaenid Car- 
acolus caracoUus (Linne, 1758) to vary greatly (0.08- 
59.0 m-). Since Geophorus bothropoma does not home 
we prefer to call the area enclosed by the recapture sites 
(figure 2) the activity range. The term homesite range 
implies that the individual possibly returns to a certain 
area after foraging outside that area. We assume that 
the activity range of Caracolus caracoUus is much larger 
than the enclosed polygon determined by recapture sites. 

Long-term study is needed to provide a meaningful 
estimate of the activity range of Geophorus bothropoma. 
The data presented here only provide an estimate of the 
minimum size of the activity range expected during a 
wet part of the year. Activity ranges are probably sig- 
nificantly smaller during the dry season. 

Homing is apparently based on chemoreception (see 
Cook, 1979, for review). The advantages and/or disad- 
vantages of homing have not been properly addressed. 
A permanent resting site should provide relative safety 
from predators and fulfill the physiological requirements 
of the individual at rest. The physiological and/or be- 
havioral basis for Geophorus bothropoma being unable 
to (or not having to) home is not known. Although the 
operculum provides protection from small predators and 
desiccation, this does not explain this species' lack of 
homing behavior. Geophorus bothropoma most com- 
monly rests on vertical rock faces. Even when inactive, 
it does not retract fully into the shell, leaving the oper- 
culum non-functional. Because Cyclophorus ceratodes 
Moellendorff, 1895 and Japonia ciliata (Sowerby, 1843) 
and J. stephanophoru (Moellendorf. 1895) were found 
to return to the same solution pits after foraging (personal 
observation), non-homing is not a general trend in trop- 
ical terrestrial prosobranchs. 

Biomass: Few data on the biomass of terrestrial mol- 
lusks are available (Strandine, 1941; Jennings & Bark- 



Page 44 



THE NAUTILUS, Vol. 102, No. 1 



ham, 1975; Richardson, 1975; Cameron, 1982) and none 
concern prosobranchs, nor do they pro\ ide eas\ or direct 
comparisons. Man\ more species must be studied before 
general trends can be demonstrated. 

Fragments of one marked specimen were found in a 
solution pit, possibly preyed on by the nocturnal shrew, 
Suncus marinus (Linne, 1766), an introduced species. 
This species was seen on two occasions at the study site. 
Many shells of Geophorus bothropoma gnawed open at 
the periphery were found in solution pits nearbs . The 
diurnal, snail-eating monitor lizard Varanti.s grayi Bou- 
lenger, 1885 rarely preys on this species (Auffenberg, in 
press). 

Although short-term data such as those presented here 
add much to our knowledge of this species, onl\' with 
long-term life-histor\ studies will ([uestions be answered 
with confidence. Combined with morphological infor- 
mation, sound behavioral and ecological data uill un- 
doubtedly contribute greatK to a better understanding 
ot the systematics and e\olution of Philippine helicinids. 

ACKNOWLEDGEMENTS 

We would like to thank the U.S. Fish and Wildlife Service 
and the New York Zoological Society for herpetological 
grants to Walter Auffenberg that allowed us the oppor- 
tunit) for the present stud>. The Philippine Parks and 
Wildlife office was very helpful in issuing appropriate 
permits. The illustrations were drawn by Armando Gar- 
cia and Marcos Guinaldo. We appreciate the patience, 
time, and insight provided bv Irvy Quitmyer, Mark 
Brenner, and Michael W. Binford of the Florida State 
Museum during the computer assisted regression anal- 
ysis. Glenn A. Goodfriend provided suggestions during 
early data analysis. Robert .\. (>ameron provided many 
constructive suggestions for analyses and format. We also 
thank Arthur J. Cain, Robert Hershler, and Fred G. 
Thompson for reviewing the manuscript. Special thanks 
are extended to our father, Walter Auffenberg, for his 
tireless assistance in the field and for his time and guid- 
ance during the writing of the manuscript. 

LITERATURE CITED 

Alexander, R D. 1961, .Aggressiveness, territoriality and sex- 
ual behaviour in field crickets (Orthoptera: (iryllidae). Be- 
haviour 17:130-223. 

Auffenberg, W. In press. The behavioral ecology of Gray's 
Monitor Lizard. University Presses of Florida 

Bailey, S. R. 1975. The seasonal and daiU patterns of loco- 
motor activity in the snail Helix asperse Muller, and their 
relation to environmental variables. Proceedings of the 
Malacological Society of London 41:415-428. 

Baker, A. N. 1973. Factors contributing towards the initiation 
of slug activity in the field Proceedings of the Malaco- 
logical Society of London 40:329-333. 

Barnes, H. F. and J. W. Weil 1944 Slugs in gardens: their 
numbers, activities, and distribution Part 1 Journal of 
Animal Ecology 13:140-175 

Barnes, H. F. and j VV Weil 1945. Slugs in gardens: their 



numbers, activities, and distribution. Part 2. Journal of 
.■\nimal Ecology 1471-105 

Berry, .A J 1966 Population structure and Quctuations in the 
snail fauna of a Malayan limestone hill. Journal of Zoology, 
London 150:1 1-27. 

Blinn, W. C. 1963. Ecolog\ of the land snails Mesodon thy- 
roidus and Allogona profunda. Ecology 44(3)498-505. 

Cameron, R. A. D. 1970. The effect of temperature on the 
activity of three species of helicid snail (Mollusca: Gastrop- 
oda). Journal of Zoolog\ . London 162:303-315. 

Cameron, R A D 1982 Life histories, densit\ and biomass 
in a woodland snail communit\ . Journal of Molluscan Stud- 
ies 48:159-166 

Cook, A. 1979. Homing in gastropods. Malacologia 18:315- 
318. 

Crawford-Sidebotham, T. J. 1972. The influence of weather 
upon the activity of slugs, Oecologia 9:141-154. 

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Iwao, S, 1977, The m*-m statistics as a comprehensive method 
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THE NAUTILUS 



Volume 102, Number 2 
April 29, 1988 
ISSN 0028-1344 

A quarterly devoted 
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T H E t7N AU T I L U S 



CONTENTS 



Volume 102, Number 2 

April 29, 1988 

ISSN 0028-1344 



Joseph R. Pawlik 
John B. O'Sullivaii 
M. G. Harasewych 



The egg capsules, embryos, and larvae of Cancellaria 
cooperi (Gastropoda: Cancellariidae) 



Martin Avery Snyder 



Latirus nuirtini (Gastropoda: Fasciolariidae), a new species 
from Honduras 



54 



Robert Hershler 
Lee-Ann C. Havek 



Shell variation of springsnail populations in the Cuatro 
Cienegas Basin, Mexico: Preliminary analysis of 
Limnocrene Fauna 



56 



Eva Pip 



Niche congruency of freshwater gastropods in Central 
North America with respect to six water chemistry 
parameters 



65 



B. W. Kilgour 


Factors affecting the distribution of sphaeriid bivalves in 
Britannia Bav of the Ottawa River 


73 








Fred G. Thompson 
Harry G. Lee 


Hypselostoma holimanae new species, a pupillid land snail 


78 






Manuel llaimovici 


Eledone gaucha, a new species of eledonid octopoti 
(Cephalopoda: Octopodidae) from southern Brazil 


82 






88 









Mcriot NolQCicai Uboratory 
LIBRARY 

MAY 61988 



Woods Hole, Mass. 



THE NAUTILUS 102(2):47-53, 1988 



Page 47 



The Egg Capsules, Embryos, and Larvae of Cancellaria cooperi 
(Gastropoda: Cancellariidae) 



Joseph R. Pawlik 

Scripps Institution of Oceanography 
UniversitN of California, San Diego 
La Jolla, CA 92093, USA 



John B. O'Sullivan 

Monterey Bay Aquarium 
886 Cannery Row 
Monterey, CA 93940 



M. G. Harasewych' 

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



ABSTRACT 

The egg capsules, and the embr\onic and larval development 
of the cancellariid gastropod Cancellaria cooperi Gabb are 
described. Egg capsules are spatulate in form, having long, 
narrow stalks that support the eggs above the surrounding sand. 
Egg capsules contain 4,000-5,000 eggs (165 ^ni in diameter), 
which undergo typical prosobranch development to hatch as 
planktotrophic veligers after 27 da> s at 15 °C Larvae in culture 
grew from 305 ^m to 890 ^m in shell length over 30 days, but 
died before metamorphosis. Limited comparative data suggest 
that long stalked egg capsules are known only in members of 
the Cancellariinae, that opercula, absent in all adult Cancel- 
lariidae, are present or prominent in the late larval stages of 
at least some species, and that developmental type cannot be 
inferred from protoconch morphology using the criteria of Shu- 
to (1974) in a majority of cancellariid species. 

Key words: Reproduction; development; larvae; eggs; egg cap- 
sules; Cancellariidae; Cancellaria. 



INTRODUCTION 

Little is known about the reproductive biology and early 
development of most of the approximately 200 species 
that constitute the neogastropod family Cancellariidae. 
The few published reports (Morch, 1869; Thorson, 1935, 
1944; Knudsen, 1950; MacGinitie, 1955; Kilburn & Rip- 
pey, 1982; Bouchet & Waren, 1985) are limited to de- 
scriptions of egg capsules and, in some cases, ova or larval 
shells attributed to cancellariids, usually on the basis of 
the proximit\' of living snails. 

During studies of the diet and feeding behavior of 
Cancellaria cooperi Gabb, 1865, a species that is attract- 
ed to and speciBcall) parasitizes the Pacific electric ray 
Torpedo calif arnica .\>res, 1855 (O'Sullivan e/ ai. 1987), 
a number of these snails were observed producing egg 
capsules. The present study supplements our know ledge 
of the natural history of this cancellariid with descrip- 



Author for correspondence. 



tions of its egg capsules and embryonic and larval de- 
velopment, and reviews the available data on the 
reproductive biology and larval development of the Can- 
cellariidae. 

MATERIALS AND METHODS 

Twenty-three specimens of Cancellaria cooperi were col- 
lected on the artificial reef "Torrev Pines #1", off San 
Diego, CA (32°53'12"N, 117°50'50"'w) at depths of 20- 
22 m using SCUBA. The animals were maintained in an 
aquarium containing sufficient sand for complete snail 
burial (4-8 cm depth) and supplied with a continuous 
flow of seawater (12-16 °C). Snails were allowed to feed 
on electric rays on a bimonthly basis, and had been 
maintained in this manner for at least 6 months prior to 
the onset of oviposition. 

Individual egg capsules were freed from the aquarium 
bottom and maintained in beakers containing continu- 
ously aerated, 1 ixm filtered seawater at 15 °C. Developing 
embryos were removed with a glass pipette through an 
incision cut along the narrow edge of the egg capsule. 
Hatched larvae of Cancellaria cooperi were cultured 
following procedures described by Paige (1986), except 
that 1 /um filtered natural seawater containing 40 mg/ 
liter each of the antibiotics streptomycin sulfate and so- 
dium penicillin G was used instead of artificial seawater. 
Larvae were fed a mixture of the green flagellates Iso- 
chrysis galhana (Park, 1949) and Pavlova lutheri (Droop) 
Green, 1975 at 10^ cells/ml. Prior to photography, larvae 
were narcotized in a 1:3 mixture of a saturated aqueous 
solution of chlorobutanol in seawater. 

Eggs and larvae for SEM examination were fixed in 
2% formalin in seawater, and stored in 70% ethanol. 
Specimens were critical-point dried and photographed 
using a Hitachi S-570 Scanning Electron Microscope. 

RESULTS 

Beginning April 21 and continuing through Ma\ 19, 
1986, a total of 15 egg capsules were laid by at least 
three snails (mean shell length = 62.3 mm), with nine 
capsules being produced sequentially b>- a single female. 



Page 48 



THE NAUTILUS, Vol. 102, No. 2 






Figure 1. Cancellaria cooperi Gabb. Apertural and right side 
view of female specimen collected by SCUBA in 20-22 m at 
artificial reef "Torrev Pines #1", off San Diego, CA {32°53'12"N, 
!17°50'50"W), USNM 846054. 1.0 x. 



Oviposition generally occurred at night or in the morn- 
ing. Snails emerged partialK from the sand, with their 
raised foot spread anteriorK and enfolded posteriorly, 
the emerging stalk protruding from the folded portion 
of the foot. Over the course of several hours, the egg case 
emerged from the ventral pedal gland and was released, 
whereupon the stalk straightened and the egg case was 
supported well above the surface of the sand. 

The spatulate capsules (figure 2; table 1) consisted of 
lenticular egg cases supported on long, narrow (250 fim 
diameter) stalks. Egg capsules were roughly rectangular 
in transverse section, with narrow keels running along 
the outer edge of each corner. A pre-formed hatching 
aperture, containing a membranous plug (figure 2, mp), 
was situated at the uppermost end of each egg case. 
Capsules were affixed to the bottom ot the arjuarium by 
holdfasts that spread from the base of the stalk. 

Each egg case contained an estimated 4,000-5,000 
spherical eggs (figures 3, 4; diameter = 164.5 ± 6.7 ixm, 
N = 10), suspended in a clear, gelatinous matrix, all of 



mp 





Figure 2. Egg capsule of Cancellaria cooperi Gabb. Frontal 
and side views 1.0 x. detail of hatching aperture 6.0 x. mp, 
mucous plug; li, membranous sack. 

which was enveloped in a membranous, transparent sack 
(figure 2, li). The lenticular walls of the egg case were 
slightly concave, creating a constriction along the mid- 
line of the case that displaced the eggs to either side. 

Spiral cleavage commenced within 12 hours of ovi- 
position, and the 8 cell stage (figure 5) was reached by 
the second day. Thereafter, the embryos became in- 
creasingly irregular in form (figures 6-8). All the embryos 
within an egg case underwent de\elopment; there were 
no nurse eggs or unfertilized eggs in the cases examined. 
By 10 da\s after deposition, the stomodeal in\agination 
was e\ident (figure 9). .\fter 12 da\s, the protoconch and 
operculum, both of conchiolin, were clearly discernable 
(figures 10, 11), and torsion was complete after 14 days 
(figures 12-14). The velar lobes were formed and in- 
creasing in size by the 16th day (figures 15, 16). On the 



Figures 3-14. Embryonic and larval development of Cancellaria cooperi Gabb at 15 °C. 3. Living etnbr\os, 1 day old 4. One- 
(l.i\-(]ld embryo, critical-point dried. Scale bar = 25 ^m. ,5. lJ\iiig embrxos, 2 days old. 6. Living embr\o, 4 days old. 7. Four- 
da\-old embryo, critical-point dried Scale bar = .50 fim. 8. Living embr\os, 8 days old. 9. Ten-dav-old embr\o. critical-point 
dried. Scale bar = 50 /nm 10. Li\ing embryos, 12 days old 1 L Hight side and ventral views of same 12-da\-old embr\o, critical- 
point dried. Scale bar = 50 ^m. 12. Living embryos, 14 days old \'.i. 14. X'cntral (13) and dorsal (14) views of 14-da\-old embrsos, 
critical-point dried. Scale bar = 50 ^m 

asr, apical sensory region; dml, dorsal maiille lip; Ik, larval kidney; o, operculum; pb, polar body; pc, protoconch; pt, prototroch; 
s, stomodeum. 



J. R. Pawlik et ai, 1988 



Page 49 




Page 50 



THE NAUTILUS, Vol. 102, No. 2 




Figures l,'j-22. Embryonic and lar\al development of Cancellaria cooperi Gabb at 15 °C. 15. Living embryo, 16 days old 16. 
Sixteen-da\-old t'nibr\o, critical-point dried. Scale bar = .50 ;um. 17. Li\ing. newK -hatched larva with carmine particles in the 
gut. 18. Shell ultrastrticture of newK-hatched larva, plane of fracture parallel to growing edge. Scale bar = 2 nm 19. .^pertural, 
dorsal and apical views of shells of newly-hatched larvae. Scale bar = 100 ^in. 20. Living larva, 30 days after hatching 21. Shell 
and operculum of larva, 30 days after hatching. Scale bar = 250 fim. 22. .-Vpical portion of adult shell (LACM 40-95.1), arrow- 
indicating transition from protoconch to teleoconch, scale bar = 500 fim. 

mo, month; o, ()j)crculuni; po, posterior ciliary band; pr. preoral ciliar\ band; sto, storiiach; vl, velar lobe. 



20tli day after capsule depobitiou, eiiibr\ os began to move 
more freely within the egg case. The gelatinous matrix 
in which the developing embrvds had been suspended 
became less viscous, and the swimming enil)ryos aggre- 
gated randondy w illiin the case. ,\ftor an average of 27 



da\s at 13 °(.' the nicnibrancjus plug occluding the hatch- 
ing aperture dissolved, and swimming larvae escaped. 
Shells ol the new K hatched veligers (figures 18, 19) had 
a mean diameter of 304,6 /um (N = 10, SD = 3.5), and 
were 3.9 ^m thick. The veligers possessed an operculum 



J. R. Pawlik et a/., 1988 



Page 51 



Table 1 . Measurements of egg capsules of Cancellaria cooperi 
Gabb. All measurements in mm (N = 10) 



Character 



Mean 



Range 



SU 



Total length 


79.6 


70.5-S3.6 


4.8 




Capsule length 


30.1 


27.8-32.8 


1.9 




Capsule width 


11.4 


10.7-12.4 


0.6 




Capsule thickness 


1.8 


1.6-2.0 


0.2 


_i 


Stalk length 


49.5 


37.7-54.0 


6.0 


O 
> 


Maximum diameter 








of holdfast 


9.1 


7.4-lO.N 


1.2 





(figure 19, o), and a fully functional alimentary system. 
When added to a small dish ot seawater containing sus- 
pended carmine particles, the veliger larvae rapidly filled 
their guts with these particles (figure 17, sto). 

Attempts to rear these veliger larvae through meta- 
morphosis were unsuccessful. At 20 °C, larvae grew rap- 
idly, reaching a mean shell diameter of 644 ^m (figure 
20) 22 days after hatching. Thereafter, the larvae began 
to die, with only a single larva surviving 30 days after 
hatching (shell diameter = 890 nm. figure 21). Larvae 
that had survived more than 22 da\s after hatching de- 
veloped propodia, but did not metamorphose in the pres- 
ence of fresh or frozen Torpedo mucus, or in the presence 
of sand from an aquarium containing adult snails. No 
crawling or search behavior associated with the onset of 
metamorphic competence was observed. Protoconchs of 
adult shells of Cancellaria cooperi indicate that the larval 
shell reaches 3.3 whorls (shell diameter = l,160;um) prior 
to metamorphosis (figure 22; table 2). 

DISCUSSION 

Egg capsules of Cancellaria cooperi resemble those of 
man\ neogastropods (e.g.. D'Asaro, 1970; Radwin & 
Chamberlin, 1973; Bandel, 1976), with notable modifi- 
cation in the length of the supporting stalk. This elon- 
gation of the stalk appears to be an adaptation for un- 
stable sediments in which capsules are deposited. Under 
natural conditions, egg capsules are most likely attached 
to buried stones or shells, with the long stalk supporting 
the egg case well above the sand surface, preventing its 
burial, and possibly protecting the eggs from small, bot- 
tom-dwelling predators. The compressed, spatulate form 
of the egg case may serve to increase the surface area 
available for diffusion of gases and waste products be- 



5.0 



4.0 



3.0 



2.0 



1.0 



PLANKTOTROPHIC 
8 



• 1 

4o05 

7o 
o9 



o10 



o6 



-^ 



LECITHOTROPHIC 



0.0 



0.1 



0.2 



0.3 



0.4 



0.5 



0.6 



D / VOL 



Figure 23. Relationship of the number of whorls (Vol) and 
the ratio of maximum diameter to number of whorls (D/V'ol) 
of cancellariid protoconchs. Solid circles denote species for which 
type of development is known or inferred on the basis of num- 
ber of ova per capsule. Open circles denote species for which 
type of development is unknown. 1, Cancellaria cooperi, 2, 
Trigonostoma foveolata. 3, .\clnicte viridula: 4, Cancellaria 
reticulata: 5, Cancellaria spengleriana. 6, Trigonostoma sca- 
lare: 7, Olssonella smithii. 8, \arona ntitraeformis: 9, Scalptia 
obliquata; 10, Cancellaria similis. 



tween the egg case and the surrounding seawater. Cap- 
sules were most frequently laid with the broad face of 
the case oriented into the stream of flow ing seawater. 

The egg cases of Cancellaria cooperi are most similar 
in morpholog} to those reported by Knudsen (1950: fig. 
18j for Cancellaria sp., and to his account of the egg 
cases of Cancellaria spengleriana (Deshayes, 1830). All 
have the characteristic long stalk, but the latter two are 
described or figured as being hemi-elliptical rather than 
lenticular in profile, and triangular rather than rectan- 
gular in transverse section. Egg cases of Trigonostoma 
joveolata (Sowerb> , 1848), a species occurring "in sand 
or gravel among rocks in low-tide pools , are similar to 
those of Cancellaria cooperi in shape, but are smaller in 
size, and have a proportionally shorter stalk (Kilburn & 
Rippey, 1982:115). Capsules of Admete viridula (Fabri- 
cius, 1780), a subtidal boreal species, are attached directly 
to the substrate and lack a stalk [Thorson, 1935; fig. 71 



Table 2. Protoconch measurements of the three cancellariid species for which the mode of development is known or inferred 
based on number of ova per capsule. Measurements are presented in the format mean/standard deviation D = diameter in mm: 
Vol = number of whorls or volutions, measured according to Jablonski and Lutz (1980:332). 



Species 



D 



\nl 



D Vol 



Cancellaria cooperi. LACM 40-95.1; LACM 39- 

94.1; LACM 39-116.2 (\ = 4) 
Trigonostoma foveolata: XM C8393; NM C8394; 

\M B7651 (N = 5) 
.\dmetc viridula: USNM 189720 (N = lOt 



1.16/0.02 

1.33/0.21 
0.78/0.05 



3.29/0.02 

2.24/0.21 
2. 12 '0.10 



0.35, 0.01 

0.59 0.08 
0.37 0.01 



Page 52 



THE NAUTILUS, Vol. 102, No. 2 



Table 3. Protoconch measurements for cancellariid species for which mode of development is not known Data are presented in 
the format of mean/standard deviation. D = diameter in mm; \'(il = number of w horls or volutions, measured according to Jablonski 
and Lutz (1980:332). 



Species 



1) 



V„l 



D \ ol 



Cancellaria reticulata: L'SNM 619108 (N = 10) 
C-ancellaria spengleriana. USNM 664965; USNM 

.344417 (N = 6) 
Trigonnstoma scalare: USNM 846304 (N = 1) 
Olssonella smithiU USNM 806986; USNM 450577; 

USNM 667720 (N = 5) 
Sarona mitraeformis: Petit collection (N = 3) 
Scalptia ohiiquala: L'SNM 629063 (N = 4) 
Cancellaria similis. USNM 664967 (N = 5) 



0.99/0.11 


2.90/0.06 


0.34/0.01 


1.08/0.07 
0.96 


3.02/0.09 
2.0 


0.36/0.02 
0.48 


0.85/0.04 
1.06/0.03 
0.89/0.03 
1.26 0.06 


2.62/0.16 
3.60/0.15 
2.45/0.09 
2.75 0,00 


0.33/0.02 
29 0.00 
n )(i 0.01 
(14(1 0,02 



(as Velutina iindala Brown, see Thorsoii, 1944); Bouchet 
& Waren, 1985: fig. 687]. The egg capsules of all can- 
cellariids studied to date have roughly elliptical, parallel 
sides w ith strongly to weakly keeled margins, and a me- 
dial, dorsal hatching aperture. Capsules with very long 
stalks appear to be restricted to the subfamily Cancel- 
lariinae. 

De\elopment oi Cancellaria cooperi is similar to that 
ot Thais hacniaatuma jiuridana as described by D'Asaro 
(1966), although C. cooperi takes about 80% longer to 
reach comparable developmental stages, and does not 
produce a noticeable sinusigeral ridge. It is interesting 
to note the presence of an operculum, particularly prom- 
inent in the planktotrophic larval stage (figure 21), in a 
family noted for the absence of opercula in adults. 

The present study comprises the first direct observation 
of oviposition and development of any species of can- 
cellariid, although the mode of development can be de- 
duced in several cases from previousK' published data 
on capsule contents. Thus, Cancellaria sp. (30-40 ova/ 
capsule, 500 /jm in diameter; Kiiud.sen, 1950:109), Tri- 
gonostovia foveolata (16 larvae/ capsule; Petit & Hara- 
sewych, in preparation), Admete viridula (6-7 larvae/ 
capsule; Thorson, 1935:67) and Admete sp. (6 larvae/ 
capsule, capsule referred to by MacGinitie, 1955:51, 
USNM 664468) all likely undergo lecithotrophic devel- 
opment, as indicated by the low number of large larvae 
or ova per capsule. To date, Cancellaria cooperi is the 
only cancellariid known to undergo planktotrophic de- 
velopment. 

In the absence of direct information, gastropod larval 
development may be inferred from the morphology of 
the protoconch at the apex of the adult shell. Thorson 's 
"apex theory" (Thorson, 1950; Jablonski & Lutz, 1980, 
1983) asserts that a large, rounded, paucispiral proto- 
conch indicates non-planktotrophic larval development, 
while a narrow, polygyrate, sculptured protoconch sug- 
gests planktotrophic de\elo[)meiit. In more tiuantitative 
studies of this relationship, Shuto (1974) found that the 
ratio ol the maximum protoconch diameter (D) to the 
number of protoconch whorls (Vol) was a reasonable 
indicator of developmental type. \'alues greater than 1.0 
were indicative ol lecithotrophic larvae, while values 
below 0.3 were more characteristic of planktotrophic 



larvae, especialK it the protoconch consisted of three or 
more whorls. Species with D \'ol ratios between 0.3 and 
1.0 usually have lecithotrophic larvae if protoconchs con- 
sist of less than 2'/i whorls. \ plot of \'ol r,s. D, \'ol values 
for 10 species of cancellariids (figure 23) indicates that 
although protoconch morphologv appears to be an ac- 
curate indicator of the mode ol larval development for 
those species of Cancellariidae that fall within the di- 
agnostic regions proposed b\ Shuto (1974), develop- 
mental type cannot be inferred for the majority of can- 
cellariid taxa using these criteria. 

As more information on the life histories and diets of 
additional species of cancellariids becomes available, the 
relationship between the mode of larval development 
and the mobility and patchiness of prey species distri- 
bution may prove fruitful ground for investigation. 

ACKNOWLEDGEMENTS 

We thank R. R. McConnaughey for assistance in col- 
lecting and maintaining animals, and R. E. Petit for 
providing information on the Cancellariidae, and for 
critical review of the manuscript. Susann Braden took 
the Scanning Electron Micrographs. 



LITERATURE CITED 

Bandel, K. 1976. Morphologie der Gelege und okologische 
Beobachtungen an Buccinaceen (Gastropoda) aus der siid- 
lichen Karibischen See. Bonner zoologische Beitrage 27: 
98-133. 

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

D'Asaro, C. N. 1966. The egg capsules, embryogenesis. and 
early organogenesis of a common oyster predator, 7'/iais 
haetnastorna floridana (Gastropoda: Prosobranchia). Bul- 
letin of Marine Science 16(4):884-914. 

D'Asaro, C. N. 1970. Egg capsules of prosobranch mollusks 
from south Florida and the Bahamas and notes on spawn- 
ing in the Jaborator), Bulletin of Marine Science 20:415- 
440 

Jablonski, D. and R A. Lutz. 1980. Molluscan larval shell 
morphology: ecological and paleontological applications. 



J. R. Pawlik et al, 1988 



Page 53 



In: Rhoads, D. C. and R. A. Lutz (eds.). Skeletal growth 
of aquatic organisms: biological records of environmental 
change. Plenum Press, New York, p 323-377 

Jabloiiski, D. and R. A, Lutz. 1983. Larval ecolog\ ot marine 
benthic invertebrates: paleobiological implications. Bio- 
logical Reviews 58:21-89. 

Kiiburn, R and E Rippey. 1982. Sea shells of southern Africa 
McMillan South Africa, Ltd., Johannesburg. 249 p 

Knudsen, J. 1950. Egg capsules and development of some 
marine prosobranchs from tropical west Africa. Atlantide 
Report 1:85-130. 

MacGinitie, G. E. 1955. Distribution and ecology of the ma- 
rine invertebrates of Point Barrow. Alaska. Smithsonian 
Miscellaneous Collections 128(9):1-201. 

Morch. O. \ L. 1869. Catalogue des Mollusques du Spitz- 
bergen recuillis par H. Kroyer. Memoire de la Societe 
malacologique de la Belgique, Tome 4, Bru.xelles. 

O'Sullivan, J. B., R. R. McConnaughey. and M. E. Huber. 1987. 
A blood-sucking snail: the Cooper's nutmeg, Cancellaria 
cooperi Gabb, parasitizes the California electric ray. Tor- 
pedo californica .\yres. Biological Bulletin 172(3):362-366. 



Paige, J. A. 1986. The laboratory culture of two aplysiids, 
Aplysia brasiliana Rang, 1828, and Bursatella leachii plei 
(Rang, 1828) (Gastropoda: Opisthobranchia) in artificial 
sea water. The Veliger 29(l):64-69. 

Radwin, G E. and J. L. Chamberlin. 1973. Patterns of larval 
development in stenoglossan gastropods. Transactions of 
the San Diego Society of Natural History 17(9):107-117. 

Shuto, T. 1974. Larval ecology of prosobranch gastropods and 
its bearing on biogeography and paleontology. Lethaia 7: 
239-256. 

Thorson, G. 1935. Studies on the egg-capsules and develop- 
ment of Arctic marine prosobranchs. Meddelelser om 
Gronland 100(5):1-71, 

Thorson, G, 1944. The zoology of east Greenland: marine 
gastropoda prosobranchiata. Meddelelser om Gronland 
121(13):1-181. 

Thorson, G. 1950. Reproductive and larval ecology of marine 
bottom invertebrates. Biological Reviews 25:1-45. 



THE NAUTILUS 102(2):54-55, 1988 



Page 54 



Latirus martini (Gastropoda: Fasciolariidae), 
a New Species from Honduras 



Martin Avery Snyder 

7-4.5 Newtown Road 
\'illanova, PA 19085 



ABSTRACT 

Latirus martini new species is described from shallow water 
off KoalaM Island. approximateK 40 miles north ot the Hon- 
duras mainland. This ne« ta.xoii differs from its nearest Carib- 
bean relative, Latirus angulatus (Roding, 1798), by its sculp- 
ture, coloration, shape, and size. The 13-16 axial ribs on the 
body whorl constitute roughly twice the number found in other 
Caribbean members of this genus 



INTRODUCTION 

In the summer of 1985 10 specimens of a new Latirus 
were taken under coral rubble b\ divers off the north 
coast of Roatan Island, Honduras. These shells were ob- 
tained by Mr. Thomas Honker of Florida, who kindly 
passed them on to the author. The specimens were all 
collected alive, the soft parts discarded, and the opercula 
glued back in place on cotton. For this reason no soft 
parts were examined, although the opercidum is de- 
scribed. Since that time, some additional specimens have 
been taken from the same locality. 

Specimens of the new species are deposited in the 
collections of the Academx of Natural Sciences of Phila- 
delphia (ANSP), the Delaware Museum of Natural His- 
tory (DMNH), and the National Museum of Natural 
History, Smithsonian Institution (USNM). .additional 
specimens are retained in the collection of the author. 

Family Fasciolariidae Gray, 1853 
Subfamily Perislerniinae Trvon, 1880 

Genus Latirus Montfort. 1810 

Type species: Latirus auraiitiacus Montfort, 1810, In 
nionut) p> [= L. gibbulus (Gmelin, 1791 j]. 

Latirus martini new species 
(figures 1, 2) 

Description: Shell heavy, small to medium in size (,21.7 
mm to 38.4 mm in length), somewhat squat, with spire 
iiearK ' s length of shell; profile inflated, w ith 8-9 whorls; 
protoconch (figure 2) of 1 '/2 whorl, smooth, bulbous, 
translucent pale orange-brown in color; aperture oval to 



squarish, greater in length than siphonal canal; anal canal 
weakly developed; axial sculpture of 13-16 prominent 
ribs, crossed by numerous, pronounced spiral cords with 
grooves between; cords alternateK larger and smaller, 
first t\\ o on body whorl about ecjual in thickness, forming 
ridge and deep suture; approximateK 45 cortls on body 
whorl running onto anterior end of siphonal canal, 25 
cords on the penultimate whorl, with every fifth cord 
somew hat stronger than others; spiral cords darkish or- 
ange-brow n turning to dark brow n between axial ribs; 
grooves between cords light caramel brown w ith yellow 
cast; shell appears dark orange-brown, with lighter axial 
ribs; columella with 3-4 distinct plicae (figure 1); teeth 
translucent, shiny, white; outer lip crenulated, with in- 
dentations corresponding to spiral sculpture; smooth por- 
tion of inner lip extending approximately to opposing 
side of the first axial rib, followed anteriorly by 14-16 
irregular w hite lirae on inside of bod\ w horl; operculum 
\ellow -brow II in color, chitinous, nearK filling aperture. 

Type locality: 15-20 feet, under coral rubble, north 
coast, Roatan Island, Honduras, summer, 1985. 

Type material: Holotv pe, ANSP 361064 (27.5 mm); 
paratype 1, DMNH 169442 (23.8 mm); paratype2, USNM 
859070 (26.9 mm); paratypes 3-5 in the author's collec- 
tion (34.0 mm. 24.8 mm, 23.6 mm). .\11 paratv pes from 
the t\ pe locality. 

Etymology: The species is named for the author's eldest 
son. 

Discussion: This species appears to be most closeK re- 
lated to the highly variable Latirus angulatus (Roding, 
1798). Several forms of the latter were illustrated by 
Bullock (19(58, 1974). Specimens of /.. angulatus from 
the Swan Islands, approximateK 120 miles north ot Hon- 
duras (Bullock, 1968: fig. 5, plate 2), and a specimen 
from the northern coast of South .\merica (Bullock, 1974: 
fig. 21) are superficialK similar to L. martini. The col- 
oration is similar but the shape and sculpture are com- 
pletel) typical of L. angulatus and thus readiK distin- 
guishable from L. martini. Latirus angulatus has 7-9 
axial ribs, about half the iiund)er of /.. martini. The spiral 
cords on L. angulatus luc weaker than those of L. mar- 
tini. The tvpical coloration is also quite different. In a 



M. A. Snyder, 1988 



Page 55 




Figures 1. 2. Lalinis martini new species. 1. Holotype, ANSP 
361064. 2. EarK « horls of parat\ pe, DMNH 169442, both from 
north coast of Roatan Island, Honduras, under coral rubble in 
5-7 m. 

dark brown specimen of L. angulatits, the raised cords 
and that portion of the whorl near the suture are dark 
brown, in areas between cords where there is fine or- 
namentation, the shell is light cocoa-tan to orange-brown. 



Generally, L. angulatus is a more elongated shell with 
the spire usualK- more than half the length of the shell. 

Possilile confusion could also occur with Leucozonia 
nas.sa (Gnieliii, 1791 ) which is somewhat similar in pro- 
tile and coloration. This shell lacks the strong a.xial sculp- 
ture of L. martini, and has a thick, black-brown oper- 
culum. A characteristic narrow \\ hite band at the base, 
commonly terminating in a small spine on the outer lip, 
serves to distinguish this species from L. martini. 

Finally, confusion might arise with the recently-de- 
scribed species Latirus vermeiji (Petuch, 1986). Latirus 
vermeiji has less pronounced surface sculpture, is dis- 
tinctlv orangish in color, and has a caramel-orange col- 
ored inner lip, whereas the inner lip in L. martini is 
yellowish brown. The roughly even spiral cords in L. 
vermeiji are more pronounced with every other cord 
being white in color. 

LITERATURE CITED 

Bullock, R. C. 1968. The fasciolariid genera Latirus, Doli- 
cholatirus. and Teralatinis (Mollusca: Gastropoda) in the 
Western .Atlantic. Master s thesis, Universit\ of Maine, 
Orono, 109 p., 8 pis. 

Bullock, R. C. 1974. A contribution of the systematics of some 
West Indian Latirus (Gastropoda, Fasciolariidae). The 
Nautilus 88(3):69-79, 26 figs. 

Gmelin,J. 1791. Systema naturae, 13th ed. Part 6:3021-4120. 

Montfort, P. D. de. 1810, Conchyliologie systematique et 
classification methodique des Coquilles ... 2. Paris, 676 p. 

Petuch, Edward J. 1986. New South American gastropods in 
the genera Conus (Conidae) and Latirus (Fasciolariidae). 
Proceedings of the Biological Society of Washington 99( 1 ): 
8-14. 

Roding, P. 1798, Museum Boltenianum, Hamburg, 199 p. 

Tryon, G. W. 1881. Manual of conchology 3. Tryon, Phila- 
delphia, 310 p., 87 pis. 



THE NAUTILUS 102(2):56-64, 1988 



Page 56 



Shell Variation of Springsnail Populations in the 
Cuatro Cienegas Basin, Mexico: Preliminary Analysis of 
Limnocrene Fauna 



Koberl Hershler 

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



Lee-Ann C. Ilayek 

Mathematics and Statistics 
National Museum of Natural History 
Smithsonian Institution 
Washington, DC 20560, USA 



ABSTRACT 

Geographic variation in shell morphometry is analyzed for 
three locally endemic springsnail (Gastropoda: Hydrobiidae) 
species occurring sympatrically in nine limnocrenesof the Cua- 
tro Cienegas Basin, Coahuila, Mexico. Despite some correlation 
of size-related variables across species, groupings of populations 
based on multivariate analyses were not very similar among 
species, nor were they strongly concordant with current drain- 
age configurations in the basin. Groups of populations of Mexi- 
pyrgu.s churinceanus Ta\ lor having different patterns of shell 
sculpture and color banding (and once considered separate 
species) were not separated similarK on basis of shell size and 
shape. Inter-population differentiation of these snails was ap- 
proximately equivalent to that of Symphuphilus minckleyi 
Taylor, whereas Mexithaunia quadripaludium Taylor was less 
variable 



INTROULCniON 

One of the more remarkable aciuatic faunas of the New 
World occurs in the small (1,200 km-), intermontane 
valley of Cuatro Cienegas, Coahuila, Mexico, which har- 
bors at least 26 locall) endemic forms (Cole, 1984; McCoy, 
1984; Minckley, 1984; Hershler, 1985). Local aquatic 
taxa show a great diversity in extent of differentiation 
relative to adjacent biota, ranging from slightK differ- 
entiated pojMilations to highly divergent genera, sug- 
gesting both long-term persistence of aquatic habitat and 
multiple invasions of the valley over a broad time scale 
(Minckley, 1969). .\quatic organisms are deployed among 
diverse, spring-fed acjuatic habitats that comprise h\'e to 
seven local drainage systems (Minckle\ , 1969; LaBounty, 
1974), providing what has been termed a "matchless 
natural laborator> " (Taylor & Minckley 1966:22) for eco- 
logical and cvolutionars- study. 

Spriiigsnails (C»astropoda: Hydrobiidae) of the basin 
are diverse [nine genera (five endemic), 13 species (nine 
endemic); Hershler, 1985], and occur abundantly in a 
large number of easiK' accessible sites, providing an ex- 
cellent (>pportunit\ to study geographic variation of pop- 



ulations, the analysis of w hich is considered crucial to 
understanding the speciation process (Gould & Johnston, 
1972; Endler, 1977). To date, researcii on this snail fauna 
has largeK been taxonomic (Ta\ lor, 1966; Hershler, 1985), 
although geographic variation of one endemic species 
was partly analyzed (Hershler, 1985; Hershler & Minck- 
ley, 1986). Taylor (1966) and Taylor and Minckley (1966) 
noted apparent diversity in extent of differentiation 
among local species: Mexipyrgus Taylor, an endemic 
restricted to large (> 25 m-) springpools (limnocrenes) 
and stream outflows, is variable enough to have been 
origiiialK considered as six nominal species (Ta\lor, 1966; 
synonymized to monotypy by Hershler, 1985), whereas 
other snails appear morphologically uniform, at least in 
the portion oi the basin that has been well studied (i.e.. 
all but the southeastern lobe; Hershler, 1985). It was 
suggested that heightened differentiation of Mexipyrgus 
resulted from marked discontinuity of its habitat: 

. . it seems that habitat of this genus is more likely to 
be discontinuous than that of other snails in the area. 
Mexipyrgus lives in soft flocculent ooze or mud in the 
lagunas, thus not in the shallows where wave action re- 
moves the fine particles Extensive marsh\ areas with small 
streams connecting larger water bodies provide no suitable 
widespread habitat. (Taylor, 1966:188) 

Variation within Mexipyrgus has largeK- been dis- 
cussed in terms of shell sculpture and color banding 
(Taylor, 1966), characters absent from or poorl) devel- 
oped in other local snails, and there has been no attempt 
to contrast intraspecific variation among members of the 
snail launa using a set of common characters, such as 
shell morphometric variables. In this paper we provide 
such a comparison between Mexipyrgus churinceanus 
Taylor and the two other species (both local endemics) 
common in basin limnocrenes. Mexithaunia quadripa- 
ludium Ta\ lor and S'yniphophilus ininckU'yi Ta\ lor (see 
figure 2). Specifically, we seek to answer the following 
questions: 1) Do these distantly related snails (Hershler, 
1985) show commonalitN of pattern of geographic vari- 
ation? 2) Does shell morphometric variation among pop- 



R. Hershler and L.-A. C. Havek, 1988 



Page 57 




Figure 1. Map of the central portion of the Cuatro Cienegas 
Basin, show ing major drainage features and collecting localities 
(numbered as in table 1 ), The inset (lower right) shows locations 
of the state of Coahuila (dashed line) and Cuatro Cienegas (dot) 
within Mexico. 



ulations of Mcxipyrgus churinceanus exceed that of the 
tv\o other limnocrene species (above) that occur s\ m- 
patrically with this species? 3) Are Mexipyrgus popu- 
lations that are well differentiated in terms of shell sculp- 
ture and color banding patterns similarl\ separated by 
shell size and shape variation? 

MATERIALS AND METHODS 

Nine limnocrenes were sampled (shown in figure 1 and 
described in the Appendix), representing the majority of 
springpools in the study area (all of basin excluding south- 
eastern lobe) where all three forms are common. Only 
isolated sources (Sites 1-4, 9) or, in cases where springs 
were connected by stream, upflow pools (Sites 5-8) were 
considered, in order to reduce possible effects of gene 
flow from contiguous populations (Hershler & Minckley, 
1986). These springs belong to four separate drainage 
systems of the basin and harbor forms of Mexipyrgus 
referable to four nominal species (see belowj. 

Mexipyrgus churinceanus was collected by sieving soft 
substrates, while the other two species, associated with 
hard substrates, were gathered by w ashing travertine and 
macrophytes in a bucket. Material was fixed in dilute 
formalin, and preserved in 70!^ ethanol. 





Figure 2. Scanning electron micrographs (printed at same 
enlargement) of cleaned shells ol springsnails from Laguna Tio 
Candido, Cuatro Cienegas, Mexico: a, Mexipyrgus churin- 
ceanus (shell height, 6.34 mm); b, Nymphophilus minckleyi 
(6.45 mm); c, Mexithauma quadripaludium (6.22 mm). 



For each species, about 15 live-collected, fully mature 
adults, recognizable by their complete and thickened 
inner shell lips, were randomly selected from collections 
from each site and dried for morphometric analysis. Af- 
ter whorls were counted (WH), shells were imbedded in 
clay in standard apertural aspect (figure 2) and shell 
outlines were draw n using a camera lucida mounted on 
a WILD M-5 dissecting microscope (12 x or 25 x ). Points 
on these drawings were digitized and values for the fol- 
lowing "standard" shell parameters (1-4) and shell shape 
descriptors (5-8) (from Raup, 1966, and elsewhere) were 
generated: 

1) Shell height (SH) 

2) Shell width (S\V) 

3) Length of bod\ whorl (LEW) 

4) Width of body whorl (WBW) 

5) Translation rate (T) 

6) Whorl expansion rate (W) 

7) Distance of generating curve from coiling axis (D) 

8) Aperture shape (SA) 

A calculated variable (S), consisting of the addition of 
SH and SW, was generated to serve as a more realistic 
measure of size than either shell length or width. Cal- 
culation of shape parameters largely followed methods 
of Kohn and Riggs (1975), with the exception being W, 
which was measured as the mean of a series of squared 
ratios of perpendicular distances from coiling axis to 
sutures (shell in apertural and not apical aspect) at half 
whorl intervals. The apertural suture was not used, due 
to frequent loosening of coiling during last half whorl of 
growth, nor were sutures used from eroded adapical sec- 
tions of the spire. Digitizing was done using the CONCH 
software program (Chapman et ai, 1988; methodology 
fulK described therein) and a GTCO Micro-Digi Pad 
12x12 linked to a KAYPRO 2000 microcomputer. 

Descriptive statistics for all morphological variables 
were obtained for each species and locality. The hy- 
potheses of homogeneity of mean differences and vari- 
ances across localities were tested for each species. An 
ANO\"A model was selected for each variable of each 



Page 58 



THE NAUTILUS, Vol. 102, No. 2 



Table I. Descriptive statistics for each species at each locahty. Data given are mean, standard deviation, and sample size (in 
parentheses). L = locality; variable abbreviations are civen in te\t 



\ ariable 


L 


SH 


sw 


S 


LBW 


WBW 


WH 


W 


D 


T 


S.\ 


Mexipyrgus churinceanun 




















1 


5.14 


2.62 


7.76 


3.37 


2.39 


6.18 


1.90 


076 


6.17 


1.32 


(15) 


0.39 


0.20 


0.57 


0.21 


0.21 


0.37 


0.16 


O07 


0.49 


O06 


2 


6.13 


3.21 


9.34 


3.85 


3.00 


7.03 


1.81 


0.67 


5.87 


1.36 


(15) 


0.30 


0.11 


0.35 


0.19 


012 


057 


0.19 


O04 


0.53 


O06 


3 


4.76 


2.48 


7.24 


294 


2.32 


6.57 


1.75 


068 


6.13 


1.35 


(15) 


0.32 


0.21 


49 


017 


019 


048 


0.14 


0.05 


0.79 


0.06 


4 


5.02 


2.72 


7.74 


3.44 


2.47 


6.23 


1.97 


073 


5.99 


1.35 


(14) 


0.23 


0.12 


029 


0.16 


012 


027 


0.16 


0.06 


027 


O07 


5 


3.99 


2.17 


6.16 


2.60 


2.02 


6.28 


1.98 


068 


6.19 


l.'jl 


(15) 


0.22 


0.17 


0.38 


0.12 


016 


0.38 


036 


0.07 


064 


0.07 


6 


7.30 


3.98 


11.3 


4.68 


3.55 


6.55 


1.86 


0.61 


5.77 


1.36 


(14) 


0.56 


0.33 


084 


024 


0.36 


041 


021 


O04 


067 


0.04 


7 


6.48 


3.17 


9.66 


4.13 


2.88 


7.28 


2.05 


O70 


6,67 


1.36 


(15) 


0.30 


0.18 


041 


0.24 


0.24 


044 


0.40 


0.07 


0,65 


006 


8 


5.49 


2.54 


8.03 


3.. 57 


2.24 


7.03 


1.92 


0.73 


7,13 


1.41 


(15) 


0.18 


0.14 


025 


014 


0.09 


061 


0.28 


0.06 


087 


06 


9 


7.16 


3.62 


lO.S 


4.39 


3.45 


6.58 


1.76 


O70 


6,44 


1.41 


(15) 


0.46 


0.22 


0.66 


019 


026 


0.32 


0.10 


O07 


0.73 


O09 


Mcxithaiima quadri 


\i(iludiuiu 




















1 


7.39 


6.44 


13.8 


6.61 


4.72 


4.57 


2,20 


0.62 


3,81 


1.11 


(15) 


0.55 


0.41 


0.90 


047 


O30 


0.24 


022 


0.05 


044 


0.06 


2 


5.82 


5.07 


109 


5.09 


3.64 


4.47 


2.26 


O60 


3.14 


1.13 


(15) 


0.41 


0.28 


0.66 


042 


0.24 


0.23 


0.32 


0.04 


0.40 


0,06 


3 


5.73 


4.83 


106 


5.04 


3.54 


4.63 


2,28 


065 


3.50 


1.13 


(15) 


0.30 


0.33 


0.59 


031 


019 


0.25 


0.33 


0.05 


O60 


0.06 


4 


6.20 


5.45 


11.7 


5.45 


3.99 


4.37 


2.05 


062 


.3..52 


1.07 


(13) 


0.57 


0.36 


087 


0.58 


025 


0.26 


021 


0.06 


0.60 


0.08 


5 


5.77 


5.09 


109 


5.14 


3.68 


4.19 


2.36 


0.61 


3..38 


1,12 


(9) 


0.61 


0.55 


1.12 


052 


041 


0.11 


041 


0.08 


048 


0,05 


6 


6.72 


6.00 


12,7 


5.76 


4.51 


4.44 


2,07 


061 


3,34 


1,03 


(9) 


0.60 


0.49 


1,07 


047 


047 


041 


0.11 


05 


048 


0,0.5 


7 


5.23 


4.50 


9.73 


4.56 


3.27 


4.32 


2.14 


0.63 


2,85 


1,12 


(15) 


0.36 


022 


0.55 


0.34 


0.20 


0.26 


0.19 


0.04 


0.21 


()()5 


8 


6.07 


5.15 


11.2 


5.18 


3.78 


4.67 


2.20 


062 


.3.17 


! 10 


(15) 


0.48 


0.30 


0,72 


042 


0.25 


029 


0.25 


0.07 


0.37 


0,06 


9 


6.40 


5.51 


11,9 


5.47 


4.15 


4.63 


2.16 


0.64 


3.51 


1.08 


(15) 


0.44 


029 


067 


0,35 


0.27 


0.30 


0.19 


004 


056 


O05 


S'yinphophiltis minckleyi 




















1 


6.66 


5.30 


12.0 


.5.19 


4.00 


4.80 


2.06 


057 


3.24 


1.12 


(15) 


0.38 


019 


0.53 


0.28 


0.20 


034 


037 


O03 


0.29 


0.05 


2 


5.99 


4.56 


10.6 


4.51 


3.76 


5.03 


1.88 


059 


3.82 


1.09 


(15) 


0.47 


031 


073 


034 


0.22 


O09 


019 


O05 


0.43 


0,03 


3 


5.12 


4.09 


9.21 


3.96 


3.15 


4.54 


2.01 


0.57 


3.35 


1 10 


(12) 


0.43 


0.26 


067 


0.31 


0.20 


0.37 


026 


0.03 


0.36 


004 


4 


5.88 


4.89 


10.8 


4.50 


3.79 


5.39 


1.89 


054 


3.32 


1.05 


(7) 


0.39 


015 


042 


0.25 


0.09 


0.35 


012 


0.02 


033 


004 


5 


5.56 


4.34 


9.90 


4.38 


3.48 


5.28 


2.08 


0.60 


3.60 


1.14 


(15) 


0.34 


0.23 


0.50 


0.28 


0.19 


0.38 


0.29 


0,03 


048 


0.03 


6 


7.17 


5.98 


1.3.1 


5.29 


4.84 


5.68 


1.86 


054 


3,19 


1,13 


(15) 


0.30 


0.26 


0.45 


0.24 


018 


0.26 


0.16 


0.04 


042 


0.05 



R. Hershler and L.-A. C. Havek, 1988 



Page 59 



Tabic I. Oititiimcd 



Variable 


L 


SH 


SW 


S 


LBW 


WBW 


W'H 


W 


D 


T 


SA 


t 


.5.69 


4.53 


10.2 


4.13 


.3.74 


5.22 


1.76 


0.58 


3.45 


1.07 


til) 


0.29 


0.27 


54 


0.19 


0.26 


0.14 


0.04 


0,05 


0.24 


0.05 


8 


7.03 


5.55 


12.6 


5.32 


4.35 


.5.23 


1.88 


0.57 


3.34 


1.18 


(10) 


0.25 


0.36 


0.55 


0.28 


0.27 


0.18 


0.29 


0.04 


0.35 


0.07 


9 


7.48 


5.85 


13.3 


5.61 


4.61 


5.47 


1.83 


0.59 


3.21 


1.17 


(15) 


0.46 


30 


0-6S 


0.35 


0..30 


0.27 


0.13 


06 


0.32 


006 



species unless very significant heterogeneity of variance 
existed, in which case the generahzed Welch test was 
used to consider mean differences. Pearson correlations 
were computed across species pairs for population means 
of each variable. Principal component analysis (PCA) 
was applied separately to each species data matri.x to 
assess and compare groupings of specimens without a 
priori assumptions. Because the units of measurement 
were distinct and non-comparable, the anaKses were 
performed on correlation matrices. Discriminant anal\ sis 
(DA) was used to determine assignment of specimens to 
the locality groupings on basis of shell size and shape. 
This anal\ sis w as computed in a stepwise manner in order 
to identify measurements contributing to significance of 
discrimination. Selection criterion was ma.ximization of 
Mahalanobis D-squared between closest pairs of locali- 
ties. The a posteriori procedure of classification analysis 
yvas performed to determine possible error of specimen 
assignment to locality . When SL and SW yvere replaced 
by their sum (S), slightly better localit) separation re- 
sulted in the multivariate analyses and these results are 
reported. 

Computations yvere performed using SYSTAT (Wil- 
kinson, 1986) on an IBM-.XT. and SPSSX Ver. 2 on an 
IBM 4381 VM/CMS system at the Smithsonian Institu- 
tion. 



prevalent among iV. minckleyi populations and partic- 
ularly pronounced for those of Mexithaiima quadripa- 
ludium. 

For each of the three species, only three principal 
components y\ere significant and y ielded meaningful in- 
formation. Eigenstructuresshoyv that the first component 
in each analysis is dominated by size and size-correlated 
variables (table 3), and explains 39-46% of the total vari- 
ation. For Mcxipyrguschitrinceaniis, tyvo shape variables 
(T, SA) dominate the second component yvhich explains 
almost 20% additional variability, and a sixth variable 
(W) is the sole measure of importance on the third axis, 
explaining 14%. For the other tyvo species, size (PCI) 
explains 10% more variation than for Mexipyrgus chu- 
rincearms, yvhereas shape (PC2 and 3) explains only 
slightly less. Weights for shape parameters are spread 
over both the second and third axes, making interpre- 
tations of these more difficult. 

Figure 4, consisting of plots of the first three PC's for 
each species with locality means (centroids) indicated, 
alloyvs comparison of relative locations of populations in 
PC space among species. Spread of centroids is largely 
along PCI, as expected. Each plot has one or ty\o tight 
clusters of a feyv centroids, w ith cluster segregation more 



RESULTS 

Descriptive summary statistics for each species by lo- 
cality are in table 1. Results of locality tests for both 
mean differences and variance are in table 2. Hetero- 
geneity- of variance yvas more pronounced for Mexipyr- 
gus churinceanus (significant for six variables at P < 
0.01 level) than for the other species. Inter-locality vari- 
ation was marked for each species, and in ail but three 
cases (LB\\', Mexipyrgus churinceanus; W, D, Mexi- 
thauma quadripaludium ) the hy pothesis of mean equal- 
ity of variables across localities was rejected at 0.05 level. 
As an example, inter-locality variation in the size-indi- 
cator variable, S (= SH -I- SW). is shoy\ n in figure 3, yvith 
significant differences (P < 0.05) indicated by non-over- 
lapped confidence intervals. Size range for each species 
is considerable; significant differences (i.e., absence of 
overlap in figure 3) are especially numerous among pop- 
ulations of Mexipyrgus churinceanus, y\ ith overlap more 



Table 2. Results of .\iialysis of \ariance, or Welch's test; and 
Bartlett's tests for homogeneity of variance for the nine local- 
ities (**: P < 0.01; *: 0,01 < F < 0,05; ns: P > 0,05). 

\ ariable 
SH SW S LBW WBWWH W D T SA 



Mexipyrgus churinceanus 
Mean ** ** ** ns 

\ari- 
ance ns * ** * 

Mexilhaiima quadripaludium 
Mean ** ** ** ** 

Vari- 
ance ns ns ns ns 

Symphophihis minckleyi 
Mean ** ** ** ** 

X'ari- 

ance ns ns ns ii'- 



** ** ** ** ** *♦ 



ns ns ns 



** ** 



ns ns ' ns ' ns 



** ** ** ** ** ** 



* ** * 



Page 60 



THE NAUTILUS, Vol. 102, No. 2 



12 - 



10 



E 
E 

CO 



6 - 



Mexipyrgus c h u rincean us 



_i 1 I ; L. 







1 




3 


5 7 


9 






Mexith a u m a qiiadripalu din m 




14 


- 




E 


12 


♦ 




10 










1 




10 


1 


\ 

< 


i 



14 



E 
E 



12 



10 - 



13 5 7 9 

Ny m phop h ilu s ni in c k leyi 



_j I { I I I 



13 5 7 9 

LOCALITY 



obvious for Mexipyrgus churinceanus and xV. minckleyi 
than for Mexithauma quadripaludium. 

Results of discriniiiiant aiiaK ses on looalit\ are in table 
4. Size (S) is the most hea\ il\ weighted \ariable on DFl 
for Mexithauma quadripaludium and \. minckleyi, with 
LBW weighting negatively; while LBW and S are ap- 
[)ro.\imatel\ ec)ual!\ and positiveK weighted on this 
tuiiction for Mexipyrgus churinceanus. .\s \s itli the prin- 
cipal components anaKsis, the Brst discriminant axis ex- 
plained almost lO^c more variation for Mexipyrgus chu- 
rinceanus than for the other two species. Note that the 
shape parameters W and D were not correlated \\ ith any 
of the functions for any of the species. 

Entry of the first variable (S) alone for Mexipyrgus 
churinceanus \ielded significant (P < 0.01) separation 
of mean values for all but a single locality pair (1, 4). 
While entr\ of four additional \ariables significantK sep- 
arated this final pair (P < O.O.j), the significance level 
decreased to 0.082 after addition of all remaining vari- 
ables. Entr\ of the first \ariable for Mexithauma quad- 
ripaludium (WBW ) and \. minckleyi (S) resulted in 
significant .separation of all but eight and fi\e pairs, re- 
spectiveK; further addition of remaining \ariables left 
four and one pairs still unseparated. 

Classification error rates for indi\idual specimens in- 
dicated considerable overlap of populations, and varied 
across localities as follows: Mexipyrgus churinceanus, 
53-100?c correct classification [87/c (overall)]; Mexithau- 
nm quadripaludium. 40-87rc [62rc (overall)]; A', minck- 
leyi. 60-100'7 [84^t (overall)]. Mexithauma quadripa- 
ludium was the poorest classified overall. \\ ith less than 
60% classification in five of nine localities. Only three 
discriminant functions were significant for this species 
{P < 0.05), compared to six for the other two. .additional 
anaKses using onK shape parameters yielded consider- 
ably poorer classification [ranging from 26-39'^c (overall) 
for each species], and confirmed the low discriminating 
power of these variables (see Hershler & Sada, 1987). 

Differentiation among drainage systems was also ex- 
amined. The study area encompasses tour local drainages 
(as recognized b\ LaBount\. 1974, with localities allo- 
cated to these as follows (following notes in Appendix): 
Becerra System (Localit> 1 ); El Garabatal (2-4); Rio Mes- 
quites S\stem (5-8); and Tio Candido System (9). Clus- 
tering of populations in PC-space (figure 4) does not 
closeK follow partitioning of localities into drainages, as 
imlicated b\ considerable spread of centroids represent- 
ing localities from El Garabatal i2-4), and from the inter- 
connected springs at Los Remojos (6-8). 

For further analysis, localities were re-grouped into 
drainage systems. .\ di,scriminant anaKsis on each species 



Figure 3. Piols of S (Sll + S\V) vs. locality for each species. 
Filled circles represent medians; and bars denote simultaneous 
confidence intervals around the median, con.structed so that if 
parentheses do not overlap, population medians are different 
at 95% confidence level. 



R. Hershler and L.-A. C. Havek, 1988 



Page 61 



Table 3. Results of Principal Compoiit'iits .\iial\scs on cacli species. Only factor coefficients having weights > 0,25 are listed. I 
= Mexipyrgus churinceanus, 11 = Mcxilhauina quadripalndium. Ill = Nijmphophihts inincklcyi, Kig. = eigenvalue; % V. = % of 
variance explained. 







I 






11 






III 






PCI 


PC2 


PCS 


PCI 


PC2 


PC3 


PCI 


PC2 


PC3 


s 


0.30 






0.30 






0.28 






LBW 


0.30 






0.30 






0.27 






WBW 


0.29 






0.30 






0.27 






r 




0.55 












44 


-0.45 


SA 




0.44 






0.43 


0.59 




0.36 


0.31 


w 






-0.77 




0.55 








0.69 


D 










0.43 


-0.64 




60 




VVH 










-0.32 


-0.37 






-0.32 


Eig. 


3.22 


1.51 


1.10 


3,19 


1.22 


1.00 


3.49 


1.35 


1.11 


% V, 


.39 2 


19.8 


13.8 


45.5 


17.4 


144 


45 6 


16.9 


13,9 



using ail \ariables \\ as able to separate the four drainage 
groups only for Mexipyrgus churinceanus: only two 
functions were signiBcant for the other species. Resuhs 
of these analyses are in table 5. Note the low weights of 
S on DFl for Mexithauma quadripalndium and N. 
minckleyi compared to these for Mexipyrgus churin- 
ceanus, and the relatively small amount (ca. 40%) of 
variation explained by the first function for all three 
species. Classification error rates for individual specimens 
were similar for the three species: Mexipyrgus churin- 
ceanus, 61-93% [68% (overall)]; Mexithauma Cjuadri- 
paludium, 46-93% [60% (overall)]; N. niinckleyi, .57- 
100% [77% (overall)], with highest classification for 
Drainage 1 (87-100%) and lowest for Drainage 3 (46- 
61%). 



There is similarity of pattern of size variation for the 
three species: populations having large-sized shells are 
concentrated in the southeastern portion of the study 
area (figure 3, Localities 6-9). For standard shell mea- 
surements, population means were significantly corre- 
lated for Mexipyrgus churinceanus and Nymphophilus 
minckleyi (r > 0.61, P < 0.05 for S, SL, SW, WBW) 
and Mexithauma quadripalndium with Nymphophilus 
minckleyi (r > 0.60, P < 0.05 for S, SL, SW, WBW, 
LBW). There were no significant correlations between 
Mexipyrgus churinceanus and Mexithauma quadripa- 
ludium. Population means for whorl number and shape 
parameters were not correlated across species with one 
exception (Mexithauma quadripalndium with Nympho- 
philus minckleyi, r = 0.64, P < 0.05 for W). 



Table 4. Results of Discriminant AnaKses on locality for each species. Standardized function coefficients (> 0.25) and pooled 
within group correlation coefficients (> 0.25) of each function with each original variable are listed. 1 = Mexipyrgus churinceanus; 
II = Mexithauma quadripalndium; III = Nymphophilus minckleyi; Eig. = eigenvalue; % V. = % of variance explained; C.C. = 
canonical correlation 







1 






II 






III 






DFl 


DF2 


DF3 


DFl 


DF2 


DF3 


DFl 


DF2 


DF3 










Standardized 


coefficients 










s 


0.64 




0.89 


0,94 


-0.66 




1,51 




-1.61 


LBW 


0.69 


0.90 


-0.95 


-0.56 


1.88 


2.02 


-1.09 


-1.43 


1.63 


WBW 


-0.35 


-1.13 




0.56 


-0.94 


-2.15 


0.35 


0.91 




D 




















T 






0.39 






0.65 




0.30 




SA 














0.34 






WH 




49 


0.61 






0.79 




0.48 


0.77 


Eig. 


13.3 


2.19 


0.61 


3.13 


0.56 


0.45 


8.95 


2.61 


0.56 


% V. 


79.2 


13.0 


3.6 


70.9 


12.7 


10.1 


69.3 


20.2 


4.3 


C.C. 


0.96 


0.83 


0.62 


0.87 
Correlation 


0.60 
coefficients 


0.56 


0.95 


0.85 


0.60 


s 


0.89 


-0.38 




0.88 


0.31 




0.84 


-0.36 




LBW 


0.94 






0.74 


0.54 




0.61 


-0.53 


0.38 


WBW 


0.64 


-0.65 




0.93 






0.83 






T 




0.34 


0.31 














SA 










0.46 








0.34 


WH 






0.72 






0,77 


0.30 


0.38 


0,78 





Page 62 



THE NAUTILUS, Vol. 102, No. 2 




1 J 

Mexipyrgus churiiiceanus 




Mexithauma quadripaludium 




(5)@ 



3 



® 



Nymphophilus minckleyi 



Despite these correlations, concordance of pattern 
among plots of PC scores for the three species is not 
impressive, although there is similarit) in order along 
size-related PCI (figure 4). Note that the tightest clus- 
tered centroids for Mexipyrgus churinceanus (Localities 

1, 3, 4) are vvideK separated for the other two species. 
SiniilarK . di\ t-rgeiit "outK iiig centroids lor gi\en sf>ecies 
{Mcxithaiinia quaclripaludium, I, S. minckleyi, 3) are 
not so differentiated in the other species. 

As mentioned abo\e, the populations of Mexipyrgus 
considered in this stud\ are referable to four nominal 
species (fide Ta\ lor. 196(-)) on basis of shell sculpture and 
color banding; Mexipyrgus churinceanus (Localities 1, 

2, 4); Mexipyrgus rnojarralis (5); Mexipyrgus lugoi (6- 
S); and Mexipyrgus carranzae (9) (the form present at 
Localit) 3 is distiiicti\e and not clearK referable to any 
nominal species). These nominal species are poorly seg- 
regated on the PC axes: extent of separation of rnojarralis 
(5) and carranzae (9) from other centroids, for instance, 
is exceeded by that seen among three populations refer- 
able to churinceanus in Los Remojos spring complex (6- 
8). 

DISCUSSION 

For all three species, most inter-population variation in- 
volved shell size and size-correlated variables. Common- 
ality of geographic variation patterns was indicated by 
significant correlation of population means tor some of 
these variables across two of three species pairs. The 
extent of this commonality was not, however, impressive 
when groupings of populations based on multivariate 
anal) ses were examined. 

A strong correlation between geographic variation pat- 
terns and current drainage configurations was not ap- 
parent for any of the species, suggesting that in this 
example development of intraspecific di\ersit\ of shell 
inorphometrv ma\ be related to ecological as well as 
historical factors (see Chernoff, 19S2, for general dis- 
cussion of this subject), although we acknowledge the 
possibilitv that the poor correlation w ith current drainage 
configuration ma\ be obscured b\ historicalK complex 
basin hydrograpln (Miiickle\, 19(i9; Hershler \ Minck- 
ley, 1985). Springpools concentrated around the northern 
tip of Sierra de San Marcos (where the study area is 
located) are higliK unilorm in water quality (Miiickley 
in Cole, 1968): single measurements taken b\ us during 
the study indicated, for instance, that temperature and 
conductivity ranged among the nine springpools from 
25.5-34.5 °C (seven localities diftering b\- < 4 °C) and 
1,825-3,500 micromhos cm (seven localities differing by 
< 430 micromhos, cm), respectively. Pools do dilter con- 



Figure 4. 'rliree-dimeii.sioiial plots ot PC cenlroids for the 
three species t\ axis. PCI; Y, PC2; Z. PC:.'3). standardized and 
viewed in perspective. Sizes of balls indicating centroids are 
scaled to heighten perspective. 



R. Hershler and L.-A. C. Hayek, 1988 



Page 63 



Table 5. Results of four-group Discriminant Analyses on drainage system for each species. Standardized function coefficients (> 
0.25) and pooled within group correlation coefficients ( > 0.25) of each function with each original variable are listed. I = Mexipyrgus 
churinceanus: II = Mexithauma quadripaludiutn; III = \'ymphophilus minckleiji; Eig. = eigenvalue; % V. = % of variance 
explained; ('.C. = canonical correlation 







I 






11 






III 






DFl 


DF2 


DF3 


DFl 


DF2 


DF3 


DFl 


DF2 


DF3 










Standardized coefficients 










s 


2.28 


0.72 


2.39 


-0.49 


2.11 


0.84 


0.87 


1.76 


-2.78 


LBW 


-2.5.3 


0.71 




1.43 


-3.12 


-0.46 


1.41 


-1.49 


1.35 


WBW 


1.05 


-0.98 


-1.87 




1.23 


-0.91 


-1.50 




1.71 


W 


















0.61 


D 


0.52 


-0.46 


0.66 




0..57 








-0.67 


T 




0.55 








0.87 








SA 


0.38 








0.42 






0.67 




WH 


-0.36 




-0.63 






0.51 




0.53 




Eig. 


3.21 


1.51 


1.10 


3.24 


1.26 


1.03 


3.49 


1.35 


1.11 


% V. 


40.2 


18.8 


13.8 


40.4 


15.8 


12.9 


43.6 


16.9 


1.3.9 


c.c. 


0.642 


0.499 


0.431 


0,710 
Correlation 


0.426 
coefficients 


0.262 


0744 


0.717 


0.217 


s 


0.58 


0..55 




0.89 


0.32 




0.49 


0.63 




LBW 


0.48 


0.58 




0.96 






0.61 


0.57 




\VB\\' 


0.65 


0.36 




0.80 


0..50 




0.83 






VV 


















0.61 


D 




-0.36 


0..59 












-0.50 


T 




0.47 




0.40 




0.68 






-0.33 


SA 


31 








-0..32 






0,46 




WH 




(144 


-0.47 




0,31 


0.48 




0,68 





siderablv in other potentially important parameters such 
as size, substrate composition, and abundance of mol- 
luscivorous cichlid fishes and the relationship between 
these features and shell geographic variation merits fur- 
ther study . 

Groupings of populations of Mexipyrgus churinceanus 
on basis of shell size and shape were not strongly con- 
cordant \\ith allocation of these to nominal species de- 
fined b\ shell color banding pattern differences. Fur- 
thermore, both univariate and multivariate analyses 
showed that these populations were no more differen- 
tiated in terms of shell size and shape than were those 
of seemingK monomorphic Xymphophilus winckleyi. 
These results suggest that evolution within Mexipyrgus 
has been mosaic, with development of striking diversity 
in shell color banding patterns and a few other features 
(including aspects of shell sculpture, and penial lobation 
pattern) coupled w ith unremarkable divergence in shell 
morphology. 

It is intriguing that these snails and A', minckleyi. 
which differ greatly in microhabitat and presumed po- 
tential for gene flow between populations at spring sources, 
have similar levels of intraspecific shell morphometric 
divergence, whereas Mexithauma quadripaludiutn. 
which broadly- overlaps in niche with the latter (Hershler, 
1984), is less variable. The possibility that these patterns 
reflect differing times of origin of lineages within the 
basin is not currently testable due to absence of fossil 
evidence. 



ACKNOWLEDGEMENTS 

Fieldwork was supported by Smithsonian Research Op- 
portunities Fund #1233F650. .Assistance in the field was 
pro\ ided by Dr. W. L. Minckley , Dr. P. Marsh, A. Riggs, 
and C. Riggs. The assistance of staff of the Scanning 
Electron Microscopy Laboratory at NMNH in photo- 
graphing shells is gratefully acknowledged. Ms. Molly 
Kelly Ryan, staff illustrator of the Department of Inver- 
tebrate Zoology (NMNH), generously prepared figures 
on very short notice. Paul Greenhall (NMNH-IZ) per- 
formed the laborious task of gathering shell morpho- 
metric data. Drs. M. G. Harasewy ch, R. Dillon, and an 
anonymous reviewer provided useful criticism of the 
manuscript. 

LITERATURE CITED 

Chapman, R. E., M. G. Harasewych, and R. Hershler 1988. 

CONCH: an interactive computer program for the analysis 

of shell coiling parameters. In preparation. 
Chernoff. B. 1982. Character variation among populations 

and the analysis of biogeography, American Zoologist 22: 

425-439. 
Cole. G. A. 1984. Crustacea from the bolson of Cuatro Cie- 

negas, Coahuila. Mexico. Journal of the .\rizona-Nevada 

.\cademy of Science 19:3-12. 
Endler, J. .\. 1977. Geographic variation, speciation, and dines. 

Princeton University Press, \J, ix -t- 246 p. 
Gould, S. J. and R. F, Johnston. 1972. Geographic variation. 

.\nnual Review of Ecology and Systematics 3:457-498. 



Page 64 



THE NAUTILUS, Vol. 102, No. 2 



Hershler, R. 1984. The h\(lrobiid snails (Gastropoda: Risso- 
acea) of the (^uatro C^ienegas Hasin: systematic relation- 
ships and ecology of a unique fauna. Journal of the .Ari- 
zona-Nevada .-Keademy of Science 19:61-76 

Hershler, R. 1985. Systematic revision of the Hydrobiidae 
(Gastropoda: Rissoacea) of the Cuatro Cienegas Basin. 
Coahuila, Mexico. Malacologia 26:31-123. 

Hershler, R. and VV. L. Minckley. 1986. Microgeographic 
variation in the banded spring snail (Hydrobiidae: Mexi- 
pyrgits) from the (;uatro Caenegas Basin. (;()ahuila. Mex- 
ico. Malacologia 27:357-374. 

Hershler, R and U VV. Sada. 1987. Springsnails (.Gastropoda: 
Hydrobiidae) of Ash Meadows, .•\margosa Basin, Califor- 
nia-Nevada. Proceedings of the Biological Society of 
Washington 100:776-843. 

Kohn. A. J. and A. C. Riggs. 1975. Morphometry of the Conus 
shell. Systematic Zoology 24:346-359 

LaBounty. J. F. 1974. Materials for the revision of cichlids 
from northern Mexico and southern Texas. LS.\ (Pcrci- 
formes: (^ichlidae). Ph.D. thesis (unpublished), Arizona 
State University. Tempe. 120 p. 

McCoy, C. J. 1984. Ecological and zoogeographical relation- 
ships of amphibians and reptiles of the Cuatro Cienegas 
Basin. Journal of the .'Vrizona-Nevada Academy of Science 
19:49-59 

Minckley, V\' L. 1969. Environments of the bolson of Cuatro 
Cienega.s. Coahuila, Mexico L'niversity of Texas at El Paso. 
Science Series Number 2, p. 1-65. 

Minckley, W. L. 1984. Cuatro Cienegas fishes: research re- 
\ iew and a local test of diversity versus habitat size. Journal 
of the Arizona-Nevada Academy of Science 19:1.3-21. 

Minckley, VV. L. and G. A. Cole. 1968. Preliminary limnologic 
information on waters of the Cuatro Cienegas Basin, Coa- 
huila. Mexico. Southwestern Naturalist 13:421-431 

Haup. D. M 1966. Geometric analx sis ot shell coiling. Journal 
of Paleontology 40:1178-1190.' 

Taylor. D. VV. 1966. A remarkable snail fauna Irom Coahuila, 
Mexico. Veliger 9:152-228. 

Taylor, D. VV. and W. L. Minckley. 1966. Neu world for 
biologists. Pacific Discovery 19:18-22. 

Wilkinson, L. 1986. SYSTAT:' the system for statistics. SYS- 
T.-VT, Inc., Evanston, IL. 



APPENDIX 

Springpools sampled are numbered as in figure 1. Lo- 
cality names are either those of Minckley (1969) or reflect 
local usage. Locality data represent air distances troni 
Cuatro Cienegas. Dates of collection are given in paren- 
theses. Catalog numbers (USNM) for voucher material 
(dry shell plus alcohol specimens) from each locality are 
given in follow ing order: Symphophihis ininrklciji Tay- 
lor, Mcxipyrgiis cliiirinrcaiuis Ta\ lor, ami Mcxithauiua 
quadripaluilium Taylor. 

1. Poso de la Becerra [(south pool) 3-1X-86], 13.7 km 
W-SW of C^uatro C^ienegas. A once enormous spring area 
(over a kilometer long) signilicantK reduced in size hy 
canal development in early 19605 (see Taylor, 1966:162; 
Minckley 1969: figs. 15, 16). Currently consisting of two 
large springpools (each ca. 50 x 125 m-) connected by 
short section ot stream. Spring orifices few and large, 
occurring in tleepest (to 7 ni) portion of pools. Water liK 
stands extcnsi\ c Spring and outflow i-onstituting a major 



drainage of basin now feeding canals. USNM 857909, 
857918, 857927. 

Sites 2-4 represent three springs in the area known as 
El Garabatal, located north of Poso de la Becerra and 
east of Becerra s outflow . El Garabatal drains to the north 
and west, and may be considered either a separate, small 
drainage or a subset of the Rio Mesquites system, the 
largest drainage in the basin. 

2. Laguiias de Juan .Santos (5-IX-86), 12.8 km W-SW 
ot Ciuatro Cienegas. Largest of El Garabatal springs con- 
sisting of series of relatively shallow (depths to 2.0 m), 
inter-connected lagunas fed by springs emerging along 
pool margins. Pool area somewhat larger than that of 
Poso de la Becerra. Extensive marshes bordering near- 
entirety of spring's perimeter. Water lily stands exten- 
sive. Stream outflow extending several hundred meters 
before terminating in shallow marshv area. USNM 
857912, 857921, 857930. 

3. Unnamed small spring, know n to biologists working 
in area as North Spring (3-IX-86), 12.0 km W-SW of 
Cuatro Cienegas. Small springpool ca. 12 x 18 m. about 
a meter deep, w ith ca. 20^ coverage b\ water lily . Out- 
flow entering second pool (not a spring); di.scharge from 
latter e.vtending 50 m before disappearing into hole. 
USNM 857910, 857919, 857928. 

4. Unnamed large spring (.5-IX-86), 11.8 km W-SW 
ot Cuatro Cienegas. Roughly circular springpool, ca. 70 
m across; depths not exceeding 3.5 m. Water lily dense 
in center of pool. Outflow feeding small marsh. USNM 
857916, 857925, 857934. 

5. "West" Laguna in El Mojarral (4-I\-86). 9.0 km 
SW of Cuatro Cienegas. Moderate-sized spring (26 x 59 
ni; Arnold, 1972:12), with depths to 4.5 m. Orifices few 
in number, cavernous. Water lily stands few, relatively 
thin. Water exiting spring \ia both a shallow, surficial 
stream and single, tubular, subsurface \ent. Stream out- 
flow entering "East Laguna, which in turn drains into 
Rio Mesquites. USNM 857908, 857917, 857926. 

Sites 6-8 are inter-coiuiected springpools known lo- 
cally as Los Remojos. 9.0 km S-SW ot Cuatro Cienegas. 
Site 6 drains into a large pool recei\ ing discharge from 
a second pool fed by outflows from Sites 7 and 8. System 
draining into Rio Mesquites. 

6. Northernmost of Los Remojos springs (5-IX-86). 
Pool ca. IS X 32 m. Depths generalK > 1.5 m; water 
lily common. USNM 857913, 857922,' 857931. 

7. Intermediate Los Remojos spring (5-IX-86). Pool 
ca. 28 X 47 m. shallow (< 1.0 m). Water lib uncommon. 
USNM S57914. 857923. 857932, 

8. Southeriunost of Los Remojos springs (5-IX-86). Pool 
moderately large (22 x 42 m), with depths increasing 
in southern end of pool to 4.0 m. Water lib common. 
USNM 857915, 857924, 857933. 

9. Laguna Tio (-andido (5-IX-86), 12.5 km S-SW of 
C.uatro Cienegas. Large spring (ca. 45 x 100 m) exten- 
sively vegetated by water lily and other macrophytes. 
Depths generally from 2.0-4.0 m. Outflow extending 
eastward, comprising part of a major system positioned 
south of Rio Mesquites drainage. USNM 85791 1, 857920, 
857929. 



THE NAUTILUS 102(2):65-72, 1988 



Page 65 



Niche Congruency of Freshwater Gastropods in Central 

North America with Respect to Six Water Chemistry Parameters 



Eva Pip 

Department of Biology 
University of Winnipeg 
Winnipeg, Manitoba R.3B 2E9 
Canada 



ABSTRACT 

Occurrences of freshwater gastropods were studied at 430 sites 
in central North America with respect to total alkahnity, total 
dissolved solids, chloride, sulphate, phosphorus, and dissolved 
organic matter. Mean niche positions for the gastropods were 
calculated for the six combined water chemistr\ parameters 
by comparing mean values for each species using agglomerative 
hierarchical cluster analysis. Total niche relations were exam- 
ined by calculatmg and summing the amount of overlap in the 
observed ecological tolerance ranges for the six parameters for 
each species pair, and appKing cluster and principal compo- 
nent anaKses. The results showed that the species occupied a 
broad spectrum of niches, with progressively increasing eco- 
logical range. Gastropods with low mean inorganic values and 
narrow niche widths were interpreted as specialists for waters 
with low concentrations of dissolved inorganic materials. Species 
which clustered together in terms of central tendencies of oc- 
currence often differed from each other in terms of overall 
niche similarity, and vice versa. Competition may have been 
more important in waters with low inorganic concentrations. 



INTRODUCTION 

A number of workers have demonstrated the importance 
of water chemistry in the distribution of freshwater gas- 
tropods {e.g.. Boycott, 1936; Hubendick, 1947; Aho, 1966; 
Harman & Berg, 1971; Dussart, 1976; Pip, 1978, 1985, 
1986; Okland, 1979; Okland & Okland, 1980). However, 
aside from the pioneering study of Finnish lakes by Aho 
et al. (1981). relatively little is known regarding niche 
relationships between freshwater gastropod species that 
coexist within a given geographical area. 

The ecological niche of a species can be described as 
an infinite h) perspace representing ail physical and biot- 
ic aspects of the environment in which the species occurs. 
Nevertheless, practical considerations restrict most niche 
studies to a few selected parameters that are thought to 
be important for the species (Levins, 1968 in Aho et al., 
1981). .According to the latter v\orker, the niche of a 
species with respect to a given parameter can be ap- 
proximated by observing the species distribution over a 
range of values of the parameter in a number of envi- 
ronments. 



The objective of the present study was to examine 
niche relationships for the freshwater gastropods in cen- 
tral North America in six water chemistry dimensions: 
total alkalinity, total dissolved solids, chloride, sulphate, 
moly bdenum reactive phosphorus, and dissolved organic 
matter (DOM). These factors were chosen because a pre- 
vious study (Pip, 1987) showed that these variables con- 
tribute towards variation in species richness of freshwater 
gastropod communities within the study area. In the 
present study a niche was defined as the range between 
the minimum and maximum values observed for each 
species in the study area, and was thus analogous to the 
concept of the "realized niche" discussed bv Hutchinson 
(1957). 



MATERIALS AND METHODS 

A total of 430 sites was examined in central North Amer- 
ica (47°-54°N and 94°-106°W) during the May-Septem- 
ber, 1972-85 seasons. All sites contained water year-round. 
Ponds (< 10 ha) comprised 41.0% of the sites sampled, 
lakes (> 10 ha) 41.5%, rivers (> 2 m deep) 10.3%, and 
creeks (< 2 m deep) 7.2%. Each site was examined for 
snails by wading or canoeing; snails on macrophytes were 
obtained by dredging with a rake or by using SCUBA. 
Search time at each site was limited to 1 hr. 

Surface water samples were collected at most sites, but 
at depths of >3 m where macrophytes did not reach the 
surface a van Dorn sainpler was used. The samples were 
placed on ice in darkness and frozen within a maximum 
of 48 hr. Samples were analyzed using methods rec- 
ommended b\ the American Public Health Association 
(1971). 

While most of the sites were sampled only once, ap- 
proximately 50 of the locations were resampled at dif- 
ferent times of the growing season and in different \ ears. 
For such sites, extreme low and high water chemistry 
values were used for statistical anaKses. Because of en- 
vironmental heterogeneity , large lakes were sampled at 
different locations which were treated as separate sites. 

Mean and niche comparisons were made using ag- 
glomerative hierarchical cluster analysis (Sneath & Sokal, 



Page 66 



THE NAUTILUS, Vol. 102, No. 2 



1973). Ward's method, a minimum variance technique 
(Wishart, 1969) was used for clustering. Cluster fusion 
distances were calculated by means oi the squared Eu- 
clidean distance measure: 

Distance,, = S, (Xj — y,)- 

Because Ward's method is an intense!} clustering pro- 
cedure, ranking of the species in the dendrograms was 
checked for misclassifications by comparing with the 
ranking obtained using the unweighted pair-group meth- 
od (Cliltortl k Stephenson, 1975), a weakly clustering 
strateg) (Sneath & Sokal, 1973). The two methods showed 
similar relative ranking; thus mi.sclassifications were not 
apparent. Since the relative importance of the six pa- 
rameters differed with species and situation (Pip, 1987), 
ail parameters were weighted etiualK in the cluster anal- 
yses. 

In construction of the dendrogram for mean niche 
position (tigure 1), mean values for each of the six pa- 
rameters were used to construct a data matrix for the 
comparison of taxa with respect to the mean positions of 
their niches. Because of differences in units and statistical 
distributions of the six chemical parameters, values were 
converteil to standardized Z scores, each parameter w ith 
a mean of and a standard deviation of 1. Ward s method 
was applied to the Z scores. 

The first step in niche comparison was the construction 
ot a combined niche congruenc\ matrix for the six chem- 
ical parameters. For each parameter, the amount of o\er- 
lap in the observed ecological ranges of each species pair 
was di\ided by the combined range of the two species 
(range between lowest minimum and highest maximum 
values). Parameter \alues were not standardized. The six 
overlap proportions were then summed for each species 
pair (maximum possible value 6.0). The combined coef- 
ficients formed a niche congruency matrix that was ana- 
i\ zed using cluster anaKsis as above to gi\e the dendro- 
gram in figure 2. 

The niche congruency matrix was also examined using 
principal component analysis (Tatsuoka, 1971), based on 
the error correlation matrix. Components were f)rthog- 
onalK rotated so that the lirsl principal component lay 
along the direction of greatest variance. 

Statistical programs used for dendrogram construction 



and principal component analysis were obtained from 
SPSS. Inc.. (Chicago. Illinois. 

RESULTS 

The ranges of concentrations of the inorganic chemical 
parameters encompassed at the stud\ sites have been 
presented elsewhere (Pip, 1986). The inorganic param- 
eters showed a number of significant (P < 0.05) positive 
intercorrelations at the stud\ sites (table 1) and could 
therefore be regarded as a block with main similar trends. 
However DOVl was significantK correlated onK with 
moK bdenum reactive phosphorus and total dissoK ed sol- 
ids. 

Five or more sets of w ater chemistr\ data were avail- 
able for 36 species in the present stud\ (table 2); only 
these species were included in cluster and principal com- 
ponent analyses. The bulk of the mean, maximum, and 
minimum values on w hicli anahses were based are gi\en 
by Pip (1986) for all parameters except DOM, which is 
given in table 2. 

The dendrogram for mean niche positions (figure 1) 
with respect to the six combined parameters reflected 
the lf)cations in the ecological ranges w here each species 
w as most often found. C^lnster 1 consisted of species w ith 
the lowest mean values lor total dissolved solids and low 
values for all other inorganic parameters. On the other 
hand, Armiger crista, Physa jcnncssi. and Fossaria mo- 
dicella differed the most frcmi all other species; these 
gastropods showed the highest mean values for total dis- 
solved solids, as well as high \alues for all other param- 
eters (Pip, 1986), including DOM. Other species in the 
spectrum could show low or high \alues of DOM which 
were largely unrelated to the concentrations ot inorganic 
parameters; for example Valvata tricarinata, Fossaria 
decampi. and Stagnicola catascopium show ed the lowest 
mean values for DOM ol all species except Campcloma 
decisum (table 2). 

The relations between species in terms ot niche con- 
gruency for the six parameters are represented by the 
dendrogram in figure 2. The groups that emerged formed 
a spectrum ol the ranges of chemical environments in 
which each species was observed. Clusters 1 and 2 con- 
sisted of species that showed narrow tolerance ranges for 



Table 1. Correlations between water chemistrv parameters at the stud\ sites Upper diaRonal = r. lower diaijonal = p. \ = 446- 
468. 

















\lii|\ Ixlemini 


Disso!\e<l 






Total 




Total 






reactive 


organic 






dissolved so 


ids 


alkalinity 


chloride 


Sulphate 


phosphorus 


matter 


Total dissolved solids 




X 




0.47* 


0.66* 


0.61* 


0.26* 


0.10* 


Total alkaiinitv 




<0.00I 




X 


0.39* 


0,1.5* 


0.41* 


0.07 


Chloride 




< 0.001 




<0.001 


X 


61* 


0.05 


0.04 


.Sniplialc 




<0.001 




0.001 


<0.001 


X 


0.05 


0.04 


\lul\ixlcnuni reactixe 


pliosplmni'' 


<00{)1 




<onni 


14 


14 


X 


0.1.3* 


Dis.solved organic matter 


0017 




OS 


(122 


22 


()()()4 


X 



* Significant correlation. 



E. Pip, 1988 



Page 67 



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Page 68 



THE NAUTILUS, Vol. 102, No. 2 



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E. Pip, 1988 



Page 69 



the inorganic parameters at low corieeMitratioiis (Pip, 
1986), The narrow tolerance ranges ot these species con- 
tributed towards larger interspecific differences, as seen 
from the larger cluster fusion coefficients; niche overlap 
with species that had broad ranges was small, while the 
probability that significant proportions of the niches of 
any two species w ith restricted ranges would not coincide 
was increased b) the narrowness of the respective ranges. 
As a result, relative niche differentiation was better de- 
fined in waters with low inorganic concentrations. 

Species witii broader ecological tolerance ranges, on 
the other iiand, could occur in waters with high inorganic 
concentrations, although in most cases they could also 
tolerate comparatively low values. These species formed 
the clusters on the upper part of the dendrogram in figure 
2, and were more numerous than species which were 
restricted to a narrow range of environments. The greater 
similarity between members of each of these clusters 
derived from the larger degree of overlap that necessarily 
arose as a result of the broad ranges. Cluster 6, composed 
of Stagnicola paliistris, Fossaria modicella, and Physa 
jennessi, represents species that occurred in the broadest 
range of chemical environments. 

Niche relations were also examined using principal 
component anal\ sis applied to the niche congruency error 
matrix, which extracted two major components that ac- 
counted for 55.4% and 22.7% of the variance, respec- 
tively. The remaining components each accounted for 
5% or less of the remaining variance. The niche relations 
for individual species are plotted with respect to the first 
two principal components in figure 3. The distributions 
that emerged reflected the groupings of figure 2, al- 
though clusters 1 and 2 from figure 2 showed close af- 
finities (groups A and B) in figure 3. Species in groups 
A and B showed narrow niche widths and low mean 
parameter values. Species in groups E and particularly 
F showed wide ecological ranges; these groups also in- 
cluded the most common species in the stud\ area (Pip, 
1978). 

The first principal component of the niche congruence- 
error matrix was significantly correlated w ith mean val- 
ues for the respective species of all six parameters (total 
alkalinity r = 0.49, P = 0.001; total dissolved solids, 
phosphorus, and chloride, all r = 0.47, P = 0.002; sul- 
phate r = 0.36, P = 0.016; DOM r = 0.33, P = 0.023; 
all N = 36). This large number of correlations derived 
from intercorrelations of these parameters at the sites 
examined (table 1). The strong correlations with the first 
component are interesting, in that the values in the orig- 
inal congruency matrix were not themselves correlated 
with any of the chemical parameters. The second prin- 
cipal component showed no significant correlations with 
the mean values of the parameters examined and was 
apparentl) related to an unmonitored factor. 

DISCUSSION 

Cluster analysis showed that the gastropods within the 
study area occupied a broad spectrum ot chemical niches. 



Table 2. Mean, maximum, and minimum values of dissolved 
organic matter (absorbance of acidified water at 27.5 nm) ob- 
served for the gastropods in the study area. 







Mini- 


Maxi- 




Species 


X 


mom 


mum 


\ 


Campeloma clecisum (Say, 










1816) 


0.142 


0.060 


0.200 


6 


Valvata sincera Say, 1824 


0.313 


0.106 


0.737 


7 


V. Iricarinala (Say, 1817) 


0.204 


0.018 


1.366 


70 


Cincinnatia cincinnatiensis 










^Anthony, 1840) 


0.210 


0,054 


1.366 


42 


Marstonia decepta (Baker, 










1928) 


0.209 


0.0.53 


0.665 


8 


Probythinella laciislris (Baker, 










1928) 


0.299 


0.180 


0.387 


8 


Amnicola limosa (Say, 1817) 


0.263 


0.039 


1.428 


102 


A. walkeri Piisbry, 1898 


0.212 


0054 


0,815 


23 


Lymnaea stagnalit Linne, 










1758 


0.325 


0.018 


1.676 


285 


Bulimnea megasoma (Sa\-, 










1824) 


0.356 


0054 


1,383 


45 


Stagnicola palustris MuUer, 










1774 


0.314 


0.060 


1.638 


182 


S. catascopium (Say. 1817) 


0.169 


0.053 


0.395 


13 


S. caperata (Sa\-, 1829) 


0.349 


0.252 


0.438 


6 


S. reflexa (Say, 1821) 


0.311 


0.130 


1.089 


7 


Fossaria decampi (Streng, 










1896) 


0.167 


0.060 


0.366 


9 


F. exigua (Lea, 1841) 


0.266 


0,030 


0.817 


15 


F. modicella (Say, 1825) 


0.345 


0.039 


1.400 


53 


F. parva (Lea, 1841) 


0.244 


0.095 


0.392 


6 


Phijsa gyrina Say, 1821 


0.307 


0.009 


1.638 


293 


P. jennessi skinneri Tavlor, 










1953 


0.387 


0.042 


1,306 


33 


Aplexa hypnonim (Linne, 










1748) 


0.442 


0.0.53 


1,638 


41 


Ferrissia parallela (Haldeman, 










1841) 


0.426 


0,115 


0,815 


6 


F. ricularis (Say, 1817) 


0.315 


0,054 


1,168 


53 


Helisoma trivolcis (Say, 1816) 


0.341 


0.018 


1,968 


207 


H. pilsbryi infracarinatum 










Baker, 1932 


0.254 


0.050 


0.684 


23 


H. corpulentum (Say, 1824) 


0.214 


0.173 


0.263 


5 


H campanula! urn (Say, 1821) 


0.237 


0.009 


0.817 


109 


H. anceps (Menke, 1830) 


0.280 


0.009 


1.512 


122 


Planorbula armigera (Sa\-, 










1821) 


0.435 


0,088 


1.638 


73 


P. campestris (Dawson, 1875) 


0.385 


0,140 


0,650 


7 


Promenetus exacuous (Sav, 










1821) 


0.324 


0,053 


1.428 


84 


P. umbilicatelliis (Cockerel!, 










1887) 


0.240 


0.053 


0.625 


i 


Armiger crista (Linne, 1758) 


0.483 


0.364 


0.815 


17 


Cyrauhis parvus (Say, 1817) 


0.345 


0.030 


1.676 


193 


G. circumstriatus (Trvon, 










1866) 


0.347 


0.053 


1.428 


27 


C. deflect us (Say. 1824) 


273 


0054 


1 428 


70 



In general, the species groupings in terms of niche sim- 
ilarity formed a continuous series with progressively 
greater ecological range. The niches of most species over- 
lapped to some extent, and only a few were mutually 



Page 70 



THE NAUTILUS, Vol. 102, No. 2 



PC I '55.4% 
,i.«''9 33 3 7 2l\3S\ 

'24 Z. •-.<^ 



/24" 
'if 



PC 2- 22.7% 

-1,0. — . — . — .- 





'10 


'•^ 


1 • . 


114 

\ 






, 5 •x . 




^- 1,' 



- - 36/ 



I •23 s 

I 

I 12\D 



l»16" 



."/ 



18 



25l_ 



»I.O 



•32 

^ •30 / 
"^!13/ 



/c 



•27; 



-1.0 

Figure 3. Distribution of species plotted « ith respect to the 
first two principal components of the niche congruenc\ error 
matrix, .^.ves have been orthogonally rotated. Numbers repre- 
sent the following species: 1 — Campeloma decisum, 2 — Va/- 
vata sincera. 3 — V. tricarinata. 4 — Cincinnatia cincinnatien- 
sis, o — Marstonia decepta. 6 — Prolnjthinella lacustris. 7 — 
Amnicola limom. 8 — A. walkeh. 9 — Lymnaea stagnalis, 10 — 
Bulimnea megasoma, 11 — Stagnicola palustris. 12 — S. cata- 
scopium, 13 — S. caperaia, 1-4 — S. reflexa, 15 — Fossaria decam- 
pi, 16 — F. exigtia, 17 — F. modicella. 18 — F. parva. 19 — Physa 
gyrina, 20 — P. jennessi, 21 — Aplexa hypnorum. 22 — Ferrissia 
parallela. 23 — F. riciilaris, 24 — Ilclisoma trivolvis, 25 — H. 
pihbryi infracarinatum, 26 — H. corpulentum, 27 — H. cani- 
panulatum, 28 — H. anceps. 29 — Planorbula armigera, 30 — F. 
campestris. 31 — Promenetus exaciwus, 32 — F. umbilicatellus, 
33 — .Armigpr crista. 34 — Cyraulus parvus, 35 — G. circtim- 
striatus. 36 — G. deflectus. 

e.xclusive for some parameters. Thus, with respect to the 
variables e.xamined, the majority of the species studied 
could theoretically occur together in the same eri\ iron- 
meiit, near the lower end of the concentration scale for 
inorganic parameters. For example 25 of the species could 
occur through the entire range of 100-200 mg liter for 
total dissolved solids, and an additional eight could occur 
in jiortions thereof (Pip, 1986). 

.Mean niche positions may be regarded as the types of 
environments for which each species was most charac- 
teristic. Stagnicola reflexa. Helisorna corpulentum. Bu- 
limnea megasoma. Campeloma decisun}. //. campan- 
ulatum. Amnicola walkeri. and Marstonia decepta were 
classified in clusters 1 and 2 in both figures 1 and 2. These 
species showed low mean values for inorganic concen- 
trations and narrow tolerance ranges; they may be re- 
garded as specialists for environments with low values 
of these factors. Since the latter environments in the study 
area are most common on the Precambrian Shield, it is 
not surprising that these species are also limited to, or 
most frequent in. Shield waters. 



It is interesting that some species showed marked dif- 
ferences in terms of their relative positions in the den- 
drograms when mean niche positions and niche con- 
gruencies were compared. For e.xample Lymnaea 
stagnalis. Stagnicola palustris. Cyraulus deflectus. and 
Helisoma anceps showed wide niches, but their central 
tendencies of occurrence were much more moderate. 
Indeed G. deflectus tended to occur most often at the 
low end of its ecological range, perhaps because it does 
not compete w ell w ith the nimierous other species which 
frequent higher inorganic concentrations. 

Species with similar mean niche positions, contained 
in the same clusters in figure 1, would be expected to 
compete with each other more frequently than with 
species in neighboring clusters, because they would be 
likely to occur in the same water bodies more often. 
Thus, although each species occupies a unique ecological 
niche (Hutchinson, 1957), the clusters of adaptively sim- 
ilar species could be viewed as functional groups (e.g., 
Stanley, 1979) which, on average, respond to certain 
aspects of the environment in similar ways. However, 
species w ithin the same clusters in figure 1 often showed 
differences in niche congruency in figure 2. For example, 
the species comprising cluster 2 in figure 1 show ed similar 
mean niche positions located at lower inorganic concen- 
trations, but of these species, Marstonia decepta was 
included in cluster 1 in figure 2, Ferrissia rivularis and 
Cincinnatia cincinnatiensis in cluster 3, Helisoma pih- 
bryi infracarinatum in cluster 4, and Cyraulus deflectus 
in cluster 5. Such differences in the ecological ranges of 
species w ith similar central tendencies of occurrence may 
ha\e been ad\antageous in that the reduced o\erlap al- 
le\ iated competition (e.g.. Hughes, 1980). since the pro- 
portion of cases where the tw o species would not be able 
to occur together was increased. 

The most common species in the study area [i.e.. Physa 
gyrina (6I.9'~( of sites sampled), Lymnaea stagnalis 
(60.7'( ). Helisoma trivolvis (45.3%), Stagnicola palustris 
(40.5%), Cyraulus parvus (39.5%)] (Pip, 19S7) showed 
broad tolerance ranges for the variables examined, and 
can occupy a wide xariety of habitats. These species 
would be expected to compete w ith each other frequent- 
ly, since large portions of their niches were congruent. 
However, they occurred most commonly at different 
mean positions w ithin their niches. Ot these five species, 
w hicli clustered in two adjacent groups in terms of niche 
congruenc) (figure 2), onl\ L. stagnalis and H. trivolvis 
occurred in the same cluster in terms of mean niche 
position (figure 1). 

Nonetheless, because ot the large amoimt of overlap, 
species with similar niches and broad ranges still often 
coexisted in the same water bodies. For example, the five 
most common species above were significantly associated 
w ith each other in the stuck area (Pip, 1978). Such species 
with large proportions ot niche oxerlap may minimize 
competition where they coexist by partitioning the hab- 
itat. For example G. parvus, a small species, feeds pri- 
mariK on periphy ton, w hile the larger of the common 
species also ingest macrophyte tissue, and ma\ favor 



E. Pip, 1988 



Page 71 



separate macrophyte species within the same water bod\ 
(Pip & Stewart, 1976). Interspecific differences in diet 
have also been reported by Reavell (1980). Other types 
of habitat partitioning may occur in certain situations, 
for example with respect to depth (Lacoursiere ct al., 
1975), temperature (Boag, 1981), and turbulence (Calow, 
1973). 

The proportionally greater niche differences between 
species found at low inorganic concentrations were re- 
flected b\ the lack of significant association between the 
species in clusters 1 and 2 in figure 2 at the same sites 
within the study area [e.xcept for M. decepta and H. 
campanulatum (Pip, 1978)]. Greater niche differences 
in unproducti\e environments ma\' have been associated 
with more intense competition (Emlen, 1973) than in 
nutrient-rich situations, where niches among many species 
were similar, and a number of interspecific associations 
were observed. Eutrophic waters show greater produc- 
tivity of algae and macroph\ tes, w hich constitute much 
of the food of freshwater gastropods (Reavell, 1980). 
Food has been proposed as an important factor regulating 
gastropod abundance and distribution [e.g., Dillon & 
Benfield, 1982). Species richness of gastropod commu- 
nities in the study area is positiveK correlated with tro- 
phic state (Pip, 1987). This correlation derives from the 
tendencies of many species to occur more often at higher 
values of phosphorus (Pip, 1978, 1986), despite the ob- 
servation that most of the species can also tolerate low- 
values of this factor. 

Competition has been put forward as an important 
factor governing gastropod distribution (e.g., Lassen, 
197.5). Species which are taxonomically and morpholog- 
ically similar {i.e., "sibling species") (Aho et al., 1981) 
often were not grouped within the same clusters with 
respect to niche similarity (e.g., the species of each Stag- 
nicola, Fossaria, Helisorna) (figures 2 and 3), indicating 
intrageneric ecological differentiation, w hile other con- 
geners (Gyraulus circumstriatus and G. deflectus) were 
grouped together. Aho et al. (1981) found a similar lack 
of consistency in the distributions of gastropods in Fin- 
nish lakes and suggested that competition is only one of 
a number of factors that contribute towards distribution. 

Conversely, most of the groupings w hich emerged in 
figures 2 and 3 contained a wide representation of dif- 
ferent taxonomic and morphological entities. Accord- 
ingly, while the niches within each group were similar 
with respect to the parameters examined, species behav- 
ior and ecological requirements were likeK to be differ- 
ent, decreasing competition. Furthermore, differences 
with respect to other parameters ma\ also have been 
important, for example t\ pe of water bod\ and bottom 
substrate (Pip, 1986), t\pe of vegetation (Pip, 1978), or 
other, unmonitored, chemical and physical variables. 

Principal component analysis yielded a representation 
of the species in a hyperspace, of w hich the first two 
dimensions accounted for more than three-quarters of 
the variance. While the first dimension was clearly re- 
lated to the types of chemical environments in which 
the species most often occurred, the groupings in figure 



3 also reflected the geographical distributions and types 
of habitats frequented by the respective species. Species 
in groups A and B were present largely in Precambrian 
Shield w aters characterized by low total dissolved solids, 
and were absent or rare west of the Shield boundary. 
Species in group C frequented ponds and small lakes 
(Pip, 1986) with intermediate water chemistry values. 
Group D occurred frequently in lakes. Group E species 
occurred in a very w ide variety of water chemistry types, 
and differed w ith respect to preferences for water body 
type, in some cases lacking discernable preferences en- 
tirely. Species in group F occurred in the most extreme 
ranges of water chemistry, but were found most fre- 
quently in ponds (Pip, 1986). 

While the niche means and widths utilized here are 
those observed for the study area as a whole, ecological 
ranges may vary for a given species in different geo- 
graphical regions (Pip, 198.5), and are likeh different for 
individual populations. The importance of particular pa- 
rameters ma\ also vary among habitats with different 
chemical and physical characteristics (Pip, 1987). Thus 
the exact position of the niche may be a plastic attribute 
that can vary to some extent with situation as well as 
time. 

LITERATURE CITED 

Aho, J. 1966. Ecological basis of the distribution of the littoral 
freshwater molluscs in the vicinity of Tampere, South Fin- 
land. Annales Zoological Fennici 3:287-322. 

Aho, J, E. Ranta, and J. Vuorinen. 1981. Species composition 
of freshwater snail communities in lakes of southern and 
western Finland. Annales Zoological Fennici 18:2.33-241. 

American Public Health Association. 1971. Standard methods 
for the examination of water and wastewater. American 
Public Health .Association, New York, 874 p. 

Boag, D. .\. 1981. Differential depth distribution among 
freshwater pulmonale snails subjected to cold tempera- 
tures. Canadian Journal of Zoology 59:733-737. 

Boycott, C. 1936. The habitats of the freshwater molfuscs in 
Britain. Journal of Animal Ecology 5:118-186. 

Calow, P. 1973. Gastropod associations w ithin Malham Tarn, 
Yorkshire. Freshwater Biology 3:521-534. 

Clifford, H. T. and VV. Stephenson. 1975. An introduction to 
numerical classification. Academic Press, New York, 229 p. 

Dillon, R. T., Jr. and E. F. Benfield. 1982. Distributions of 
pulmonate snails in the New River of Virginia and North 
Carolina, U.S.A.: interaction between alkalinity and stream 
drainage area. Freshwater Biology 12:179-186. 

Dussart, G. B. J. 1976. The ecology of freshwater molluscs in 
north west England in relation to water themistry. Journal 
of Molluscan Studies 42:181-198. 

Emlen, J. M. 1973. Ecology: an evolutionarx approach. .\d- 
dison- Wesley Co., Reading, \i.\, 493 p. 

Harman, W. N. and C. O. 15erg. 1971. The freshwater snails 
of central New York with illustrated ke\s to the genera 
and species. Search: Cornell L niversit\ Agricultural Ex- 
perimental Station 1:1-68. 

Hubendick, B. 1947. Die Verbreitungsverhaltnisse der lim- 
nischen Gastropoden in Sudschweden. Zoologische Bid- 
rucke Uppsaliense 24:419-.559. 

Hughes, R. N. 1980. Strategies for survival of aquatic organ- 



Page 72 



THE NAUTILUS, Vol. 102, No. 2 



isms. In: Barnes, R. S. K. and K. H. Mann (eds.). Funda- 
mentals of aquatic ecosystems. Blackwell Scientific Pub- 
lications, Oxford, p. 162-184. 

Hutchinson, G. E. 19.57. Concluding remarks. C^old Spring 
Harbor Sym[X)sium. Quantitative Biology 22-41.5-427 

Lacoursiere, E., G N'aiiiancourt, and R. Goiilure 1975. Re- 
lation entre les plantes aquatiques et les gasteropodes (Mol- 
lusca. Gastropoda) dans la region de la centrale nucleaire 
Gentilly 1 (Quebec). Canadian Journal of Zoology 53:1868- 
1874. 

Lassen, H. H. 1975. The diversity of freshwater snails in view 
of the equilibrium theory of island biogeography. Oeco- 
logia 19:1-8. 

Okland, J. 1979. Distribution of en\ ironmental factors and 
fresh-water snails (Gastropoda) in .Norway: use of Euro- 
pean invertebrate surve\ principles Malacologia 18:211- 
222. 

Okland, J. and k. A. Okland. 1980. Acidification threatens 
trout diet. Research in Norway 1980, p. 21-27. 

Pip, E. 1978. A survey of the ecolog\ and composition of 
submerged aquatic snail-plant communities, Canadian 
Journal of Zoology 56:2263-2279. 

Pip, E. 1985. The ecolog\ of freshwater gastropods on the 



southwestern edge of the Precambrian Shield. Canadian 

Field-Naturalist 99:76-85. 
Pip. E. 1986. The ecology of freshwater gastropods in the 

central Canadian region. Nautilus 100:56-66. 
Pip. E. 1987. Species richness of freshwater gastropod com- 
munities in central North America Journal of MoUuscan 

Studies 53:16.3-170. 
Pip, E. and J M Stewart. 1976 The dynamics of two aquatic 

plant-snail associations. Canadian Journal of Zoolog\ 54: 

1192-1205. 
Reavell, P. E. 1980. A study of the diets of some British 

freshwater gastropods. Journal of Conchology 30:253-271. 
Sneath, P. H. A. and R. R. Sokal. 1973. Numerical taxonomy. 

\V. H. Freeman and Co., San Francisco, 573 p. 
Stanley. S. M. 1979. Macroevolution, pattern and process. VV. 

H. Freeman and Co., San Francisco, 332 p. 
Tatsuoka. M. M. 1971. Multivariate analysis: techniques for 

educational and psychological research. John \\ ile\ & Sons, 

New York, 310 p.' 
Wishart. D. 1969. Mode analysis: a generalization of nearest 

neighbour which reduces chaining effects. In: Cole, .\. J. 

(ed). Numerical taxonomy. Academic Press, London, p. 

282-308. 



THE NAUTILUS 102(2):73-77, 1988 



Page 73 



Factors Affecting the Distribution of Sphaeriid Bivalves in 
Britannia Bay of the Ottawa River 



B. \^ . Kilgour 
G. L. Mackie 

Departmt'iit of Zoology 
Universit) of Guelph 
Guelpli, Ontario NIG 2W1 
Canada 



ABSTRACT 

Distribution of Sphaeriidae in Britannia Bay of the Ottawa 
River was not homogeneous and was primarily affected by 
sediment particle size and depth. Total sphaeriid diversity is 
maximal at particle sizes near 0.18 mm. Total sphaeriid density 
and abundance of the genus Pisidiurn increase with decreasing 
water depth. Distribution of species within the genera Pisidiurn 
and Musculium is variable. These data indicate that compar- 
isons of pisidiid distribution should consider at least depth, and 
particle size in the sampling design. 



INTRODUCTION 

The importance of sphaeriids as part of the aquatic com- 
munity has lead to a number of studies dealing with the 
distributional patterns of this group (Healey , 1978; Mack- 
ie et ai. 1980). Most studies, liowever, ha\e dealt with 
distribution patterns among a group of lakes, rivers, or 
sections of a river. Usually it is assumed that variation 
among sites is greater than the variation within sites. 
However, data in Avolizi (1976) and Holopainen (1979) 
suggest that the abundance of some species can be vari- 
able within a single site, but little work has been done 
to determine the extent of this variation. 

Britannia Bay , a section of the Ottawa River just above 
Ottawa and Hull, Canada, is relatively unpolluted, and 
is known to contain a diverse sphaeriid fauna ( 15 species), 
including members of Musculium. Pisidiurn, and 
Sphaerium (Mackie, 1971). The purpose of this study 
was to determine if the depth distribution of sphaeriids 
in Britannia Bay is homogeneous, and if distribution is 
a function of the sediment particle size, organic matter 
content, algal biomass, and depth. 

MATERIALS AND METHODS 

During mid-.\ugust. 1985. benthos and sediment samples 
were taken at eight depths (0.25. 0.35, 0.50, 1.0, 2.0, 3.0. 
4.0, and 6.0 m) along a transect in Britannia Bay of the 
Ottawa Ri\er (figure 1). 

At each depth, five Ekman grab (15 x 15 cm, 0.32 



mm mesh screen on top) samples were taken, except at 
0.36 m where four samples were taken. A sediment sam- 
ple was taken at each depth from which three subsamples 
(approximately 100 ml) were taken. Two for determi- 
nation of percent organic matter content and one for 
geometric mean particle size. 

From each benthic sample, all sphaeriids and algae 
(entangled masses of Lyngbya sp. and Vaucheria sp.) 
were hand picked. Sphaeriids were counted and iden- 
tified to species. Algae was dried at 70 °C and weighed 
to the nearest 1.0 mg. Total sphaeriid diversity (Shannon 
& Weaver, 1949) and density (N/m-), and individual 
species abundances were calculated for each benthic 
sample. 

The organic content of the sediments was estimated 
by ashing two of the sediment samples from each depth 
and expressing the result as percent loss on ignition. Geo- 
metric mean particle size was determined with the third 
sediment sample from each depth using the methods 
outlined by Lotspeitch and Everest (1981). 

Stepwise multiple regressions (Ostle & Mansing, 1975) 
were performed to determine which environmental fac- 
tors significantly affected total sphaeriid diversity and 
density, and the abundances of individual genera and 
species. Only those variables found to be independent 
by simple correlations (table 1), were used coincidentally 
in the regression models. 

Box plots (McNeil, 1977) determined that all data, 
except diversity values, were most normally distributed 
when the depth means were transformed by logjQ. All 
simple correlations and stepwise multiple regressions were 
calculated using the normalized data. 

RESULTS 

Depth means of algal biomass, and ?? organic matter 
content are presented in figures 2 and 3. Both parameters 
increased with increasing water depth. Trends in sedi- 
ment particle size are given in figure 4. Compared to 
other depths, sediments were coarser at the 1.0, 2.0. and 
6.0 m depths. 

Twelve sphaeriid species were found in Britannia Bay 



Page 74 



THE NAUTILUS, Vol. 102, No. 2 




laoa- 



Figure 1. Map of the Ottawa River, and an enlarged map of 
Britannia Bay above Ottawa indicating location of tfie transect. 
Depth contours on enlarged map are in metres. 



with P. casertanum the most abundant species. Mean 
abundances at each depth for each species are presented 
in table 2. 

Diversit) values were highest at the shallow depths 
(0.2.5-0.50 in) and at .'3.0 and 4.0 m (figure 5). Stepwise 
multiple regression indicated that diversity increased with 
decreasing particle size (table 3). Figure 6 indicates a 
unimodal distribution oi diversity in relation to geo- 
metric mean particle size. Maximum diversilN occurred 
near a particle size of 0.18 mm. 

Regression analysis indicated that total sphaeriid den- 
sity significantly increased w ith decreasing water depth 



Table I. C^nrrelatinns between some environmental variables. 
Sigiiilitant rorrclaliniis [P < O.O.'j) are indicated 1)\ asterisks 
(•) 



.Mgal 
biomass 



Organic 
matter 



Depth 



Organic matter 
Depth 
Particle size 



0.787* 
0.839* 
f) Vyl 



0.863* 
-0,043 



80D- 



<00- 



o.so 1.0 2.a 
Depth (in) 



«.0 



6.0 



Figure 2. Mean algal biomass at each depth along the transect 
with 95^< confidence limits indicated b\ vertical lines. 



(table 3). Density at 0.25 m was less than that at 0.36 
and 0.5 m (figure 7). 

Abundance of the genus Fisidium significantK in- 
creased w ith decreasing water depth (table 3). A similar 
relationship was found for P. variabile. Abundance of P. 
casertanum significantK increased with finer sediments 
and lower organic matter content. None of the other 
seven Pisidium species were correlated with the en\i- 
ronmental variables. 

Abundance of the genus Musculiniu did imt correlate 
with any of the variables tested, .\bundance ot A/, trans- 
versum increased significantK with decreasing water 
depth (table 3). Abundance of A/, secitris correlated pos- 
itively with algal biomass. 

No relationship was found tor the genus Sphaerium 
which was represented b\ onK one species, S. striati- 
nuin. 



at 2- 




322 



0.25 0.36 0.50 1.0 2.0 3.0 t. 6.0 

Depth (m) 

Figure 3. Mean organic matter content of the sediments at 
each depth along the transect with 95^t confidence limits in- 
dicated l>\ \frtital lines. 



B. W. Kilgour and G. L. Mackie, 1988 



Page 75 




Figure 4. Geometric mean particle size at each depth along 
the transect. 



2.5- 



1.5- 



0.50 l.D 

Depth (m) 



6.0 



Figure 5. Mean diversity values at each depth along the tran- 
sect with 95^0 confidence limits indicated bv vertical lines. 



DISCUSSION 

In Britannia Bay, total sphaeriid diversity significantK- in- 
creases with finer sediments. The finer sediments at 3.0 
and 4.0 m, which correspond with high diversity values, 
contradict the positive correlation which exists between 
particle size and depth in rivers (Hamill, 1975). The sand 
bar between the 3.0 and 4.0 m sites (figure 1) may have 
affected the distribution of sediments in the same manner 
that P'olke and Ward (1957) describe for a sand bar in 
the Brazos River. 

The unimodal distribution of diversity in relation to 
particle size (figure 6) indicates that diversity is maximal 
at a particle size near 0.18 mm, and decreases with finer 
or coarser material. Finer substrates enhance burrowing 



(Rogers, 1976), production (Hamill, 1975), growth and 
reproduction (Mackie & Qadri, 1978), and may improve 
survival in young clams (Gale, 1976). Fine substrates, 
however, may reduce oxygen a\ailability (Meier-Brook, 
1969; Hartnoll, 1983). The particle size at which diversity 
is maximized may represent the optimal grain size for 
efficient burrowing and /or oxygen/ nutrient availability'. 

Total sphaeriid density increases in shallow water in 
Britannia Bay. From the literature, it appears that re- 
source availability and utilization may be affected by 
depth. Bacteria and phytoplankton (especially diatoms), 
which are food resources for sphaeriids (Holopainen, 
1985), are generally more abundant in shallow water (0- 
2.0 m) than in deep water (> 5.0 m) (Hargrave, 1970). 

The drop in total sphaeriid density at 0.25 m which 



Table 2. Mean abundance (No. /m-) of individual species at each depth along the transect. 











Depth (m) 




























Total 




















no. 


Species 


0.25 


0,36 


0.5 


1.0 


2.0 


3,0 


4.0 


6.0 


sampled 


Musculium 




















M. securis (Prime, 18.32) 


17.8 





8.9 





88.9 


80.0 


115.5 


35.6 


39 


M. transversum (Say, 1829) 


444.4 


266.6 


26.6 


8.9 


17.8 











80 


Pisiditim 




















P. casertatuim (Poli, 1795) 


942.1 


1,833.2 


1,173.2 


35.6 


17.8 


142.2 


106.7 





437 


P. duhitim (Sav, 1816) 


17.8 


11.1 


26.7 





8.9 





8.9 





8 


P. fcrrugineum (Prime, 1852) 





22.2 


44.4 








8.9 








8 


P. hcnsloivanum (Sheppard, 1825) 





188.9 


168.9 











8.9 





37 


P. lilljeborgi (Esmark and Hoyer, 1886) 


124.4 


600.0 


1,262.1 


8.9 


8.9 


35,6 


88.9 


35.6 


230 


P. nitidiim (Jenvns, 1832) 


231.1 


944.4 


1,528.7 








17,8 


26.7 


26.7 


291 


P. punctatum (Sterki, 1895) 


133.3 


422.2 


160.0 








8.9 


17.8 


8.9 


75 


P. lahabile (Prime, 1852) 


106.7 


100.0 


195,5 


8.9 














44 


P. ualkcri (Sterki, 1895) 





11.1 




















1 


Sphacriuni 




















S. striatiniim (i.amarck, 1818) 








195.5 


26.7 


35,6 


231.1 


62.2 


17.8 


64 


Total no. sampled 


227 


396 


539 


10 


20 


59 


49 


14 


1,314 





Page 76 



THE NAUTILUS, Vol. 102, No. 2 



2.5- 




.'-*-, 




2- 




A 


\ 




1.5- 




'' \ 


s,. 




1- 




1 1 ^'**«,«„,^^ 




.5- 














5 


1 
.2 


1 
.25 


. 



sooo- 



Geometric Mean Particle Size (mm) 

Figure 6. Relationship between mean diversity and geometric- 
mean particle size with 95/( confidence limits indicated b\ 
vertical lines. 



4500- 

<ooo- 

'^ 3500- 

E 

o 3000- 
>^ 2500- 
i 200O- 

Of 

CD 

c 1500- 
o 

Of 

^ 1000- 
500- 



^ 




0. 50 1.0 2. 
Depth (m) 



3.0 



4.0 



6.0 



Figure 7. Mean densits at each depth along the transect with 
95% confidence limits indicated h\ vertical lines. 



is inconsistent with the regression model, ma\ result from 
wave action and or summer lethal temperatures, which 
can significantly reduce the numbers of sphaeriids in the 
shallower areas in Britannia Ba\ (Mackie, 1971; Mackie 
& Qadri, 1978). 

Regression analysis suggests that the genus Pisidium 
prefers shallower water in Britannia Bay. The relation- 
ship between P. variabile and depth is consistent with 
the distribution of the genus. In contrast, P. casertanum 
prefers finer substrates with lower organic matter con- 
tent. The distribution of P. casertanum suggests that 
there can be variabilitv in the distributional patterns of 
species within a genus. 

That the distribution of the genus Musciiltum is not 
related to an\ oi the variables tested may be due to 
variation in the distribution patterns of the two species 
of Musculiiim. Competition between M. securis and M. 
transversttni in Britannia Bay, as described b\ Mackie 
et al. (1978), ma\ also be responsible for these results. It 
is likely that two species are not sufficient to determine 
a general distribution pattern of a genus. 

Table 3. Significant relations between the dependent \ ariables 
(D) and the independent variables (I), where, R = correlation 
coefficient between 1) and I, a = algal biomass, b = depth, c = 
geometric mean particle size, and d = % organic matter content 
of the sediments 









Significance 


D 


I 


fi (D.I) 


level 


Total sphaeriid diversity 


c 


-0.791 


P < 0.020 


Total sphaeriid density 


b 


-0.720 


P < 0.050 


Musculiiuu scrtnis 


a 


as 19 


P < 0.025 


Mttsculinni trunsrcr^iun 


1, 


-0 940 


P < 0.010 


I'isidiuni 


b 


-0.733 


P < 0.030 


Pisidium cascfidiniin 


c 


-0.711 






d 


-0614 






total R- = 


0.883 


P < 010 


Pisidium vnhahilc 


1. 


(19.39 


P < OOIO 



The positive relationship between abundance of M. 
securis and algal biomass has also been described by 
Mackie and (^adri (1978) who suggested that M. securis 
utilizes the algae Lyngbya and Vaucheria as food re- 
sources in Britannia Ba\. The \ariabilit\ in the distri- 
bution of the two species of Musculium ma\ also be 
related to the food forms with which the clams are as- 
sociated. 

That the abundance of the genus Sphaeriuin and most 
Pisidium species do not correlate with the em ironmental 
variables ma\ be a result of modal relationships with 
these variables, or relationships with variables that were 
not tested. The relationships found centre on the im- 
portance of particle size and depth, w hich have been 
shown b\ Green ( 1971 ) to be important factors affecting 
distribution of sphaeriids among lakes. It is suggested 
that in studies dealing with the distribution of sphaeriids, 
particle size and depth are important factors to be con- 
sidered in the sampling design. 

ACKNOWLEDGEMENTS 

We wish to thank Mr. Clarence Kilgour for help with 
the sampling. Comments from Mr. Chris McCiall and 
two anonymous reviewers improved the manuscript. The 
study was funded b\' the Natural Sciences and Engi- 
neering Research Council of Canada, Grant No. A-9882 
awarded to C; LM. 



LITERATURE CITED 

.\volizi, R J. 1976. Biomass turnover in populations of vivip- 
arous sphaeriid clams: comparisons of grow th. fecundity, 
mortalitv and liiomass production Hvdrobiologia 51: 163- 
168. 

Folke, R. L and W C W ard 1957 Brazos River Bar: a study 
in the significance of grain size parameters. Journal of 
Sedimentary Petrology 27:3-26 

Gale, W. F. 1976. \'ertical distribution and burrowing be- 



B. W. Kilgour and G. L. Mackie, 1988 



Page 77 



haviour of the fingernail clam, Sphaeriuni transvcrsum. 
Malacologia 9:121-125. 

Green, R. H. 1971. .\ multivariate statistical approach to the 
Hutchinsonian niche: bivalve molluscs of central Canada. 
Ecologv 52:543-556. 

Hamili. S. E. 1975. Production of sphaeriid clams and am- 
phipod crustaceans in the Ottawa River near Ottawa-Hull, 
Canada. M.Sc. thesis. University of Ottawa, 98 p. 

Hargrave, B. T. 1970. Distribution, growth, and seasonal 
abundance of Hyallela azteca (Amphipoda) in relation to 
sediment microflora. Journal of the Fisheries Research 
Board of Canada 27:685-699. 

Hartnoll, R. G. 1983. Substratum. In: Earll, R. and D. G. 
Erwin (eds. ). Sublittoral ecology: the ecologv of the sub- 
littoral benthos. Clarendon Press, 0.\ford, p. 97-124. 

Healey, M. C. 1978. Sphaeriid mollusc populations of eight 
lakes near Yellowknife, Northwest Territories. Canadian 
Naturalist 92:242-251. 

Holopainen, I. J. 1979. Population dynamics and production 
ol Pisiditim species (Bivalvia, Sphaeriidae) in the oligo- 
trophic and mesohumic lake Paajarvi, Southern Finland. 
Archiv fiir Hvdrobiologie 54iSuppl ):466-508. 

Holopainen, I. 1985 Feeding biology of Pisidiidae (Bivalvia) 
with special emphasis on functional morphologv of the 
digestive tract Lammi Notes 12:5-9. 

Lotspeitch, R. B and F. H Everest. 1981 A new method for 
reporting and interpreting te.xtural composition of spawn- 
ing gravel United States Forest Service Research Note 
PN\\-369. 



Mackie, G. L. 1971. Some aspects of the distributional ecology 
of macrobenthos in an industrialized portion of the Ottawa 
River near Ottawa and Hull, Canada M.Sc. thesis, Uni- 
versity of Ottawa, 161 p 

Mackie, G. L. and S. U. Qadri. 1978. Effects of substratum 
on growth and reproduction of Musculium securis (Bi- 
valvia: Sphaeriidae). The Nautilus 92:135-144. 

Mackie, G.L.S. U.Qadri, and R. M. Reed. 1978. Significance 
of litter size in Musculiuryi securis (Bivalvia: Sphaeriidae). 
Ecology 59:1069-1074. 

Mackie, G. L., D. S. White, T. W. Zdeba, and N. A. Thomas 
1980. A guide to freshwater mollusks of the Laurentian 
Great Lakes with special emphasis on the genus Pisidium. 
U.S. Environmental Protection Agency, Duluth, MN, 144 p. 

McNeil, D. R. 1977. Interactive data analysis: a practical 
primer. John Wiley and Sons, Toronto, 186 p. 

Meier-Brook, C. 1969. Substrate relations in some Pisidium 
species (Eulamellibranchiata: Sphaeriidae). Malacologia 9: 
121-125. 

Ostle, B. and R. W. Mansing. 1975. Statistics in research: basic 
concepts and techniques for research workers, 3rd ed. Iowa 
State University Press. Ames, 596 p. 

Rogers, G. E. 1976. Vertical burrowing and survival of sphae- 
riid clams under added substrates in Pool 19, Mississippi 
River. Iowa State Journal of Research 51:1-12. 

Shannon, C. E. and W. Weaver. 1949, The mathematical 
theory of communication. University of Illinois Press, Ur- 
bana, 125 p. 



THE NAUTILUS 102(2):78-81, 1988 



Page 78 



Hijpselostonia holimanae New Species, a Pupillid 
Land Snail from Thailand 



Fred G. Thompson 

Florida State Museum 
University of Florida 
Gainesvilie, FL .32611, USA 



Harry G. Lee 

709 Lomax Street 
Jacksonville, FL 32204, 



USA 



ABSTRACT 

Hypselostorna holimanae new species (Gastropoda, Pulmonata, 
Pupillacea, X'ertiginidae) is described from a limestone range 
near Kancliaiiaburi. Thailand Its morphological characteristics 
are distinct to the extent that no close relationship between this 
form and other nicmbers of the genus is apparent. Hypselos- 
torna tubifcrum (Benson, 1856) from Burma is it closest phy- 
logenctic and geographic congener. The two species are alike 
in details ol sculpture and umbilical width, but differ widely 
in shell shape and aperture barriers. Similarities of reduced 
apertural dentition among other species of Hypselostorna are 
cnnsidered to be due to convergence. 



INTRODUCTION 

The land snail fauna of Thailand is \ery poorly known. 
The genus Hypselostorna is wideK distributed from Bur- 
ma through CJamhodia, Vietnam, Mala\a, the Loo C^hoo 
Islands and the Philippine Archipelago. Yet not a single 
species has hitherto been recorded from Thailand. The 
species de.scribed in this paper was collected by Stephen 
C. Holimaii while serving as a Peace Corps volunteer in 
Thailand. During his few opportunities of leisure, Mr. 
Holiman collected mollusk specimens for his mother, 
Mrs. Stanley (Bonnie) Holiman of Jacksonville, Florida, 
who has a.ssemhled a private collection of fair importance 
becau.se of the data that accompanies the specimens. The 
mollusks from Thailand were submitted to us by Mrs. 
Holiman for identification. The new species of Hypse- 
lostorna described in this paper shows strong similarity 
to the generic t\ pc species from Burma, and differs con- 
spicuously troi7i other known species. We are honored 
to name this snail after Mrs. Holiman in recognition of 
her bringing this species to our attention. 

Hypselostorna holimanae new species 
(figures 1-6) 

Description: Shell small, about 2.6-2.9 mm wide and 
2.3-2.6 mm high; about 0.8.5-0.9.5 times as high as w ide. 
Shell turlMii-shaped with a moderateK long conical spire 
(figures l-.'3, holotype). Last whorl conspicuously en- 
larged, and with a distinct peripheral angle and a basal 



angle. Last whorl flat below peripheral angle, and strong- 
1\ shouldered above. Occasional specimens may be weak- 
1\ furrowed below periphery. Last whorl ascending at 
about 10° to longitudinal axis of spire (figures 2, 4). Neck 
of last w horl becoming narrowed behind aperture, and 
extending forward for about ',5 of minor diameter of last 
whorl (figure 3); indented externally over junction of 
angulo-parietal lamella and slightK so over colinnellar 
lamella. Base of shell broadK umbilicate due to lateral 
expansion of last whorl. Umbilicus about 0.40-0.53 times 
minor diameter of last w horl as measured across the basal 
angle. Whorls 4.6-4.9. Protoconch consisting of about 1.5 
whorls that appear smooth under light microscopy; at 
higher magnifications the whorls are sculptured with a 
dense mesh of fine reticulating threads that have an 
underlying spiral arrangement (figure 5). Whorls of te- 
leoconch sculptured with raised spiral threads that are 
nearK uniformly distributed o\er the surface of the shell 
(figure 4). Threads weak liut distinct on spire; most con- 
spicuously developed on last whorl (figure 6). Spiral 
threads interrupted at irregular intervals b\ incremental 
growth striations, which in some specimens ma\' cause 
the spiral sculpture to appear cancellate or beaded on 
the spire. Color dark brow n with a light brow n aperture 
and white lamellae within the aperture. Face of aperture 
translucent w ith fine radiating lirow n lines due to raised 
spiral threads on opposite surface Peristome broadly ex- 
panded and nearK uniformly wide around aperture. Ap- 
erture barrier with four teeth located on inner rim of 
the aperture just behind expanding peristome and ar- 
ranged o|5posite each other in a cross-configuration. .An- 
gulo-parietal lamella rectilinear, torming a single short 
undulating blade; bifid as is typical for genus, with an- 
gular segment smaller and separated from parietal por- 
tion by a weak notch. Palatal and basal plicae short and 
lateralK llattcned; coiiliiied to inner ritu of peristome. 
Columellar lamella tubercular and located on a slight 
callus. 

Measurements in mm for the holot> pe and three para- 
t\ pes (I'F 1 13428) selected to show \ariation follow , The 
minor diameter is the transverse w idth of the bod\ w horl 
posterior to the neck of the aperture. Other measure- 
ments were made of standard parameters. 



F. G. Thompson and H. G. Lee, 1988 



Page 79 




1 






Figures 1-6. Hypselostoma holimanae new species. 1-3. Holotvpe (UF 113427). x 26. 4-6. Parat\pe (UF 113483). 4. x 40. 
5. X 120, 6. X 160. 



Page 80 



THE NAITILUS, Vol. 102, No. 2 



Specimen Height Maj. w. Min \v .\per w. L'mbil Whorls 



Holotype 


■2A 


2.6 


2.4 


1.4 


1.2 


4.7 


Paratvpe 


2.3 


2.7 


2.3 


1.4 


0.9 


4.8 


Paralype 


2..3 


2.6 


2.3 


1.2 


1.0 


4.6 


Paralype 


2.6 


29 


2.5 


1.6 


1.0 


49 



Type locality: Thuiluiid. Kaiicliaiiaburi Pro\ ince, small 
limestone range on the west border of the Kanchanaburi 
Agricultural College, about 15 km w est of Kanchanaburi. 
Holot\pe: IF 113427; collected March 15, 1987 b\ Ste- 
phen Holiman. Paratypes: UF 113428 (12), UF 113483, 
Mahidol Universit\ Malacology Collection, Bangkok (12), 
and the private collections of Harry G. Lee (12) and 
Bonnie Holiman (12). 

The specimens constituting the t\ pe series are recently 
dead shells. They were collected at the height of the dry 
season from leaf-litter in a forested knoll at the top of 
the limestone range. 

COMPARISONS 

Hijpselostoma Benson, 1 856 belongs to a group of genera 
that also includes Boysidia .\n\e\. 1881, Paraboijsidia 
Pilsbry, 1917, Gyliotrachela Tomlin, 1930, Aulacospira 
Moellendorff, 1890, Anauchen Pilsbry, 1917, and Sys- 
tenostoma Bava\ and Dautzenberg, 1909. These genera 
were monographed b\ Pilsbry (1917). More recently, 
Jutting (1950) reviewed the known species of the first 
four genera. Members of this group have protoconch 
sculpture similar to that described above for H. holi- 
manae. The variation of this sculpture w ithin the group 
and its ph\ logenetic significance v\ill be discussed else- 
where (Thompson, in preparation). The classification of 
species within these genera is sometimes problematic, 
and no one who has worked with the group seems to 
have been comfortable with the systematic arrangements 
of previous authors (Pilsbry, 1917; Jutting, 1950, 1961; 
Solem, 1981; Thompson & Dance, 1983). Much of the 
problem centers on the emphasis that is placed on the 
development of the parietal and angular lamella, the 
reduction ot the aperture barrier, the sculpture on the 
teleoconch and the degree to which the aperture is at- 
tached to or free of the previous whorl. Hypselostoma 
hohmanae is placed w ithin Hypselostoma because of the 
fused angulo-parietal lamella, the free aperture and the 
spiral sculpture ot the teleoconch. The angulo-parietal 
lamella is weaker than in most other species of the genus, 
and the aperture barrier in general approaches the con- 
dition that characterizes Anauchen. However, all of the 
species of Anauchen lack any indication of an angular 
lamella and none have raised spiral sculpture on the 
teleoconch. 

Few species of Hypselostoma have thus far been de- 
scribed from the mainland on southeast .\sia. Pilsbrv 
(1917) discussed the species known at that time. More 
recently. Jutting (1950) briefly reviewed the genus and 
described several additional species (1950, 1961, 1962). 
Hypselusluma. as used b\ both Pilsbrv and Jutting, is a 
poKpIn letic assemblage that contains species belonging 
to at least three genera. F"our species (H. terae Tomlin, 



1939, H. megaphona Jutting, 1949, H. elaphis Jutting, 
1949, // pcrigyra Jutting. 1949) were transferred to Boy- 
sidia (Dasypupa} b> Thompson and Dance (1983:109). 
Three others related to H. dayanum Stoliczka, 1871 be- 
long in another genus (Thompson, in preparation). 

The remaining species placed in Hypselostoma differ 
strikingK from //. holimanac. although H. tubiferum 
(Benson, 1856) from Burma appears to be the most closely 
related congener. It has a similarly broad umbilicus and 
similar spiral sculpture on the teleoconch, though not as 
strong. Because of these similarities the two species are 
considered to be more closeK related than either is to 
other known species w ithin the genus. The two differ in 
several conspicuous features. Hypselostoma tubiferum 
is much more depressed, the aperture is turned upw ard 
above the apex of the spire and the aperture barrier 
consists of 6-7 lamellar teeth (see Pilsbrv, 1917 for a 
description and illustrations). Hypselostoma holimanae 
differs from all other mainland species b\ its strong spiral 
sculpture, its reduced aperture barrier with tubercular 
columella lamella, its strong circumumbilical basal keel, 
and its broad umbilicus. Some Philippine species {H. 
sibuyanicum Moellendorff, 1896. H. quadrasi Moellen- 
dorft, 1896, H. roebeteni Moellendorff, 1894, and H. 
latispira Thompson and Auffenberg, 1984) are similar 
to H. holimanae in that the aperture barriers have been 
reduced to denticles, although they are not as weak as 
in //. holimanac. None has sculpture as strongK- devel- 
oped as //. holimanae. none has a basal keel circum- 
scribing the umbilicus, nor are any as widely umbilicate. 
Other differences that separate H. holimanae from these 
Philippine species are the shape of the shell and the 
contour of the whorls. Hypselostoma roebeleni and H. 
latispira are \er\ depressed species in which the up- 
turned aperture reaches almost to the level of the apex 
of the spire. Hypselostoma sibuyanicum and H. quadrasi 
have more slender shells w ith regularh increasing whorls, 
and both are narrovvlv rimate. The combination of all 
of these morphological differences indicate that the sim- 
ilar aperture barriers of the Philippine species and H. 
holimanae have evolved independentlv through tooth 
reduction from more comple.x ancestral conditions, and 
that no close relationship can be inferreil on the basis of 
these barriers. 



ACKNOWLEDGEMENTS 

We wish to thank Mrs. Stanley (Bonnie) Holiman for 
generously donating the holotvpe and part of the para- 
type series to the Morida State Museum and to Mahidol 
University. Stephen Holiman provided us with infor- 
mation about the type localitv. The illustrations com- 
prising figures 1-3 were rendered bv Ms. Wendy B. 
Zomlefer, Staff Illustrator, Florida State Museum. 

LITERATURE CITED 

Jutting, VV. S. S. van Bentliem 1950. The Malavan species 
of Boysidia. Paraboysidia, Hypselostoma, and Gyliotra- 



F. G. Thompson and H. G. Lee, 1988 



Page 81 



chela (Gastropoda. Pulnioiiata, Nertiginidae) with a cat- 
alogue of all the species hitherto described Bulletin of the 
Raffles Museum 21 5-47 

Jutting, W S. S. van Benthem 1961. Additional new species 
and new localities of the family X'ertiginidae and the gen- 
era Oophana and Opisthostoma from Mala>a. Bulletin of 
the Raffles Museum 26:34-48, pis. 8-14. 

Jutting, W. S. S, van Benthem. 1962. Coquilles terrestres 
nouvelles de quelques collines calcaires du Cambodje et 
du Sud X'ietnani. Journal de Conchyliologie 102:3-15. 

Pilsbrv, Henr\ .\. 1916-18. Manual of conchology, Ser. II, 
24. Pupillidae, Gastrocoptinae. Philadelphia, p. i-xii, 1- 
380, pis. 1-49. 



Solem, A. 1981. Small land snails from Northern Australia, 
1: species of Gtjiiotraclicia Tonilin, 1930 (Mollusca, Pul- 
monata, Nertiginidae). Journal ot the Malacological So- 
ciety of Australia 5:87-100 

Thompson, F. G. and K. Auffenberg 1984. Hypselostoma 
latispira, a new pupillid land snail Irom the Philippine 
Islands. Proceedings, Biological Societv of Washington 97: 
86-89. 

Thompson, Fred G. and S. Peter Dance 1983. Non-marine 
mollusks of Borneo, II. Pulmonata: Pupillidae, Clausili- 
idae. III. Prosobranchia: Hydrocenidae, Helicinidae Bul- 
letin Florida State Museum 29:101-152. 



THE NALTILL'S 102(2):82-87, 1988 



Page 82 



Eledone gaucha, a New Species of Eledonid Octopod 
(Cephalopoda: Octopodidae) from Southern Brazil 



Manuel Haimovici 

Dt-partamento de Oceaiiografia 

Fuiida^ao Universidadf de 

Rio Grande 

Cx. Postal 47-1, Rio Crande RS 96.200 

Brazil 



ABSTRACT 

A lieu species of Eledone is described from tlie southwestern 
Atiaiitif at depths of 60 to 160 m, off Rio (Jrande do Sul, Brazil. 
The characters that distinguish this species from the other species 
of the genus are presented, as well as a morphometric com- 
parison with the sympatric Eledone massyae Voss, 1964. 



INTRODUCTION 

Several cephalopods were collected during a survey of 
the demersal resources of the inner shelf of Rio Grande 
do Sul between Solidao (30°40'S) and Chui (34°20'S) 
at depths to 100 m (figure 1) by the R/V "Atlantico Sul" 
of Funda^ao Universidade de Rio Grande (FURG). Hai- 
movici and Andriguetto (1986) stated that two species 
of the octopod genus Eledone were found. One of them, 
E. rnassijae, was described by \ oss (1964) and the second 
was a new species. Both sympatric species possess the 
generic character of papillae at the tips of the non- 
hectocot) iized arms of the males. Morphological analysis 
presented here as well as ijiochemicai dilferences fouml 
by Levi et al. (1985) separate these two similar species. 

MATERIALS AND METHODS 

All specimens studied were killed with fresh water, fixed 
in 10% formalin for 24 hr and preserved in 70% ethanol. 
Measurements were taken in millimeters, and all mea- 
surements and indices used are among tho.se described 
by Roper and Voss (1983). The drawings of £. gaucha 
are by Jose Angel Alvarez Perez. The types are deposited 
in the Museu Oceanografico de Rio Grande (MORG), 
Museu Nacional de Rio de Janeiro (MNRJ), Museu de 
Zoologia da L ni\ersidade de Sao Paulo (MZUSP), Museo 
Nacional de Historia Natural, Liruguay (MNHN), Museo 
de Ciencias Naturales de La Plata, Argentina (MCNLP), 
University of Miami Mollusks Laboratory (UMML) and 
National Museum ot Natural History, Smithsonian In- 
stitution (USNM). 



This work has been partially supported by a grant of 
the Conselho Nacional de Pesquisas Cientificas e Tec- 
nologicas (CNPq), Proc. 403293-83 and the surve\ fi- 
nanced bv the Comissao Interministerial de Recursos do 
Mar (CIRMj. 

Eledone gaucha new species 
(figures 2-14, table 1) 

Material examined: Holot\ pe; 6 32.5 mm ML, K \' 
Atlantico Sul, cruise 13/83, S'ta. 37, 32°58'S, 5ri9'W, 56 
m, trawl, 17 Nov. 1983, MORG 23544. Parat\pes: 7 6 
27-41 mm ML and 5 9 32-34 mm ML, R \' Atlantico 
Sul, cruise 13/83, Sta. 37, 32°58'S, 51°19'W, 56 m, trawl, 
MORG 23544; 2 <5 32-34 mm ML, R/\' Atlantico Sul, 
cruise 13/83, Sta. 39, 32°50'S, 50°45'W, 87 m, trawl, 18 
Nov. 1983, MORG 23545: 1 S 21 mm ML. R \" Atlantico 
Sul, cruise 13/83, Sta. 2, 31°46'W, 100 m, 9 Nov. 1983, 
MORG 23846; 1 9 33 nun ML, R/\' Atlantico Sul, cruise 
10/83, Sta. 51, 33°43'S, 52°13'W, 60 m, trawl. 29 Aug. 
1983, MORG 23547; 4 9 28-33 mm ML, R/\' Atlantico 
Sul, cruise 10/83, Sta 57, 33°13'S, 51°25'\\'. trawl, 60 m, 
30 Aug. 1983, MORG 23548; 1 <J 33 mm ML and 1 9 44 
mm ML, R/V Atlantico Sul, cruise 10/83, Sta. 53, 33°52'S, 
5r55'W, 52 m, trawl, 29 Aug. 1983, UMML 32.2066; 
1 6 34 mm ML and 1 9 42 mm ML, R \' Atlantico Sul, 
cruise 10/83, Sta. 53, 33°52'S, 5r55'W, 52 m, trawl, 
29 Aug. 1983, USNM 816613; 1 <5 26 mm ML and 1 9 
29.5 mm ML, R/V Atlantico Sul, cruise 10/83, Sta. 51, 
33°43'S, 52''13'W, 74 m, trawl, 29 Aug. 1983, MZUSP 
25242; 1 S 32 mm ML, R/\' Atlantico Sul, cruise 10/83, 
Sta. 55, 32°52'S, 51°43'\V, 140 m, trawl, 30 Aug. 1983, 
MGNLP 4681; 1 9 29.5 mm ML, R/V Atlantico Sul, 
cruise 10/83, Sta. 57, SS'IS'S, 51''25'W, 60 m, trawl, 30 
Aug. 1983. MC:NLP 4682; 1 <? 30 mm ML and 1 9 38 
mm ML, R/ \' Atlantico Sul, cruise 10/83, Sta. 53, 32°52'S, 
51°55'W, 52 m, trawl, 29 Aug. 1983, MNHN 14762, 
MNHN 14763; 1 S 35 mm ML and 1 9 42 inm ML, R/V 
Atlantico Sul, cruise 3/83, Sta. 42, 33°02'S, 51°30' \V, 130 
m, trawl, 17 June 1980, MNRJ 5626, MNRJ 5627. 



M. Haimovici, 1988 



Page 83 



DESCRIPTION 

Animal small, maximum observed mantle length 65 mm 
(figures 2, 3). Mantle firm, not very thick, ovoid and 
elongated (MVVl 3: 61.0; 2: 59.2) separated from head 
by small constriction. Body surface smooth with some 
papillae on the dorsal mantle and head. Head narrow er 
than mantle (HWl S: 36.2; 9: 37.3); eyes slightly protu- 
berant. One supraocular cirrus. Funnel long (FLI S: 44.6; 
9: 45.4) with anterior half free (FFuLI 6: 21.6; 5: 21.1); 
funnel organ (figure 4) W shaped. 

Arms long and rather slim. Arms length order 
1 >2>3>4 in most specimens \\ ith dorsal arms markedly 
longer than others. All arms longer in males (except hec- 
tocotylized arm) than in females (ALl I to IV S: 273.2- 
239.8-134.9-208.1; 9: 250.0-213.8-200.6-191.0). 

Suckers small, uniserial, well separated and deeply set 
into arms. Suckers somewhat crowded near tips of the 
arms on females. Two rows of minute fleshy papillae on 
all non-hectocotylized arms of males (figure 10). Number 
of suckers on basal half of the first right arms varies from 
17 to 23 (ASC 6: 20.1; 9: 19.4), suckers slightly larger on 
all arms of males (lASI S: 7.7; 9; 6.5). 

Web extends over half the length of arms and de- 
creases from dorsal to ventral surface; web formula most 
trequentlv A:B:C:D:E. Web indices similar in both sexes; 
24.4-24.1-21. 6-18.6-15.6 for males and 24.4-23.9-21.1- 
18.3-14.9 for females. 

Third right arm in males hectocotylized (figure 9) 
(HcAI: 58.9). Ligula small (LLI: 8.8), without differen- 
tiated calimus (figure 9); spermatophore grove deep. 

Gill count in external hemibranch from 7 to 10, most 
trequentlv 8 in males and 9 in females (Gilc S: 8.2; 9: 
8.9). 

Males reproductive system with no special figures (fig- 
ure 11). Penis long and tubular (PLI: 23.4) with a rather 
short diverticulum (PdLI: 7.8). Spermatophores undif- 
ferentiated (figure 13) from 12 to 20 mm (SpLI: 45; 
SpLWT: 1.56). Number of spermatophores in a sample 
of 40 mature males from 7 to 92 (mean 32.1). 

The proximal o\ iduct long, the oviductal glands small, 
the distal oviducts shorter and somewhat stouter (figure 
12). Intraovaric eggs oval (figure 14); maximum lengths 
of apparently mature eggs approximately 8 mm. Num- 
ber of developing eggs in a sample of 42 maturing fe- 
males ranged 10-55 (mean 30.2). 

Buccal mass well developed, with small anterior sali- 
vary glands and larger posterior salivary glands. Esoph- 
agus connects to developed crop leading to muscular 
stomach and smaller spiral caecum united by two ducts 
to the large digestixe gland. Intestine thin and leads to 
the anus adjacent to ink sac opening. Ink sac superficialK 
embedded in the digestive gland (figure 7). Radula with 
a tricuspid rachidean tooth, three lateral teeth and a 
marginal plate (figure 8). 

Color of living animals changed from brow n to almost 
white dorsally always remaining clear ventrally. Color 
of specimens preserved in alcohol purplish gray dorsally 
and pale \ellow ventralK'. Inner surface of the arms, 
mouth, and ventral mantle with few chromatophores. 



29° 




















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54° 53° 52° 51° 50° 

Figure 1. Sample localities of Eledone gaucha new species. 

Type locality: 32°58'S, 51°19'W, south Rio Grande, Bra- 
zil in 56 m. 

Etymology: The name gaucha refers to the coastal planes 
of Argentina, Uruguay, and southern Brazil and its peo- 
ple. 

Distribution: Eledone gaucha is knov\n onK from off 
Rio Grande do Sul between Solidao (30°40'S) and Chui 
(34°20'S) (figure 1). 

DISCUSSION 

The new species belongs in the genus Eledone because 
of a single row of suckers, the heteromorphic arms in 
the males, with the non-hectocot\ lized arms having the 
suckers at their tips modified into fleshy papillae or lam- 
inae, and the hectocot\ lus without a differentiated cal- 
imus. These characters distinguish Eledone from related 
genera Pareledone, Vosseledone, Graneledone and oth- 
ers (Palacio, 1978). 

The genus Eledone occurs on the Atlantic continental 
shelves of South America. Africa, and Europe and in the 
Mediterranean Sea. It includes six described species: E. 
moschata (Lamarck, 1798) and £. cirrhosa (Lamarck, 



Page 84 



THE NAUTILUS, Vol. 102, No. 2 






;-. V" :■■ r;" ■■■:1'. . ' Vj 




-'T^ 



■r \ mm .... 







!5 
I,, 



10 



M. Haimovici, 1988 



Page 85 



Table 1. Ranges and means of measurements and 
Eledune gaticlui new species, from southern Brazil 



indites of 10 males and 10 females each of Eledone massyae V'oss, 1964 and 







Eledone mass, 


yae \ dss 


, 1964 






Eledune gaucha new species 








Males 






Females 






Males 






Females 






Lower 




Upper 


Lower 




Upper 


Lower 




Upper 


Lower 




Upper 


Index 


limit 


Mean 


limit 


limit 


Mean 


limit 


limit 


Mean 


limit 


limit 


Mean 


limit 


Total length (TL) 


148.0 


176.4 


207.0 


195.0 


218.2 


236.0 


81.0 


123.2 


137.0 


100.0 


131.9 


178.0 


Mantle length (ML) 


45.0 


54.0 


63.0 


60.0 


65.3 


71.0 


21.0 


31.6 


41.0 


28.0 


35.5 


50.0 


Mantle width index (MWI) 


65.1 


76.2 


88.2 


75.8 


79.3 


88.9 


47.6 


61.0 


71.9 


40.0 


59.2 


70.0 


Head width index (HWI) 


40.4 


47.4 


56.5 


38.2 


41.9 


45.0 


34.2 


37.5 


40.0 


28.0 


36.2 


43.6 


1° right arm length index 


























(1 ALI) 


184.0 


201 4 


216.0 


185.0 


209.9 


247.0 


244.0 


275.2 


322.0 


227.0 


250.0 


284.0 


2° right arm length index 


























(II .ALII 


195.0 


203.3 


235.0 


190.0 


217.3 


248.0 


193.0 


239.8 


281.0 


162.0 


213.8 


242.0 


3° right arm length index 


























(III ALI) 


155.0 


199.1 


225.0 


185.0 


214.1 


2.52.0 


85.0 


134.9 


159.0 


155.0 


200.6 


2.36.0 


4° right arm length index 


























(IV ALI) 


191.0 


204.0 


224.0 


186.0 


215.5 


243.0 


134.0 


208.1 


256.0 


162.0 


191.0 


213.0 


Arm formula (AF) 




4:2:1:3 






2:4:3:1 






1:2:3:4 






1:2:3:4 




A web depth index (A WDI) 


19.1 


24.3 


28.5 


19.8 


22.5 


27.4 


16.8 


24.2 


30.9 


18.9 


24.4 


28.6 


B web depth index (B WDI) 


18.2 


24.5 


27.5 


21.0 


23.8 


29.6 


19.0 


24.1 


30 1 


18.9 


23.9 


25.8 


C web depth index [C WDI) 


22.9 


25.9 


28.3 


21.2 


24.1 


28.9 


18.8 


21.6 


28.9 


17.6 


21.1 


23.7 


D web depth index (D WDI) 


22.7 


24.8 


28.6 


20.9 


24.3 


27.6 


14.6 


18.6 


23.0 


15.8 


18.3 


20.5 


E web depth index (E WDI) 


13.4 


20.6 


25.3 


18.6 


20.8 


22.9 


11.5 


15.6 


21.6 


12.3 


14.9 


17.4 


Web formula (WE) 


C;D:B:A:E 


D:C:B:A:E 


A:B:C:D:E 


A:B:C:D:E 


Gill lamellae count (GiLC) 


8/10 


9.4/9.2 


11,7 


9/10 


9.7/9.6 


8/10 


7/9 


8.2/8.5 


9/9 


8/8 


8.9/8.5 


10/10 


Funnel length index (ELI) 


37.8 


41.9 


45.9 


40.8 


44.2 


48.2 


36.6 


44.6 


51.9 


32.5 


45.4 


51.7 


Free funnel length index 


























(FFuLI) 


17.5 


23.2 


29.5 


20.9 


23.8 


27.9 


14.6 


21.6 


31.0 


10.0 


21.1 


30.3 


Arm sucker count (ASC) 


16 


18.1 


20 


16 


17.8 


20 


17 


19.4 


22 


18 


20.1 


23 


Arm sucker index I (I ASI) 


6.3 


i . i 


9.1 


7.0 


7.9 


8.7 


4.7 


7.7 


9.3 


6.0 


6.5 


8.0 


Arm sucker index II (II ASI) 


6.3 


8.0 


9.1 


7.0 


8.2 


9.6 


4.7 


7.3 


9.3 


5.0 


6.4 


8.0 


Arm sucker index III 


























(III ASI) 


5.5 


7.4 


8.9 


7.0 


7.9 


8.5 


4.7 


7.6 


9.6 


5.0 


6.0 


7.5 


Arm sucker index I\' 


























(IV ASI) 


5.5 


7.2 


8.9 


6.7 


7.7 


8.4 


4.7 


6.7 


7.8 


3.0 


5.1 


6.7 


Penis length index (PLI) 


19.3 


28.1 


39.3 








19.5 


23.4 


28.1 








Penis diverticulum length index 


























(PdLI) 


6.6 


15.0 


22.2 








4.9 


7.8 


10.5 








Spermatophore length index 


























(SpLD 


20.9 


31.6 


37.7 








38.3 


45.0 


50.0 








Spermatophore w idth index 


























(SpWI) 


1.8 


2.5 


3.1 








1.2 


1.6 


2.1 








Hectocotvlized arm index 


























(HcAI)' 


59.0 


69.2 


81.4 








41.7 


58.9 


67.4 








Ligula length index (LLI) 


5.3 


80 


97 








49 


8,8 


12.5 









1798) both from the NE Atlantic and Mediterranean, £. 
thijsanophora Voss, 1962 and E. caparti Adam, 1950 
from the SE .\tlantic, and E. massyae Voss, 1964 and £. 
gaucha trom the SW .Atlantic. 

Eledone gaucha seems to be a relatively abundant 
species off Rio Grande do Sul. The possible reasons why 
it has not been recognized to date are its small size, its 



similarity to E. massyae, and the scarcity of scientific 
cephalopod collections by research vessels in this area. 
Due to its small size it is not retained in the cod ends of 
commercial trawlers and e\en in the R \' Atlantico Sul 
surveys it most often was found entangled in the wings 
of the net. Palacio (1977) reviewed several museum col- 
lections of Argentina, Uruguay, and Brazil and found 



Figures 2-12. Anatomical features of Eledone gaucha new species. 2. 3. Dorsal (2) and lateral (3) views of holot>pe (MORG 
23544, 32 mm ML). 4. Funnel organ. 5. Upper mandible. 6. Lower mandible. 7. Digestive tract. 8. Radula. 9. Hectocot) lized 
arm tip 10. \on-hectocot\ lized arm tip of male 1 1. Male reproductive organs. 12. Female reproductive organs. 



Page 86 



THE NAUTILUS, \ol. 102, No. 2 





Figures 13, 14. Rpprocliictive products of Eledone gaucha 
new species. 13. Sperinatophore. 14. Egg. 



onl\ one eledoiiid. E. inaasyae. A survey of the MORG 
collection by the author showed one specimen of E. gau- 
cha (MORG 15'341) lornierK classified as E. massijae. It 
is expected that reviews in other collections w ill expand 
the range of £. gaucha. 

The sympatric species E. nias.syae and E. gaucha ini- 
tially look similar but many differences may be seen in 
a more detailed stud\'. In ortler to compare both species 
morphologicalK, the same indices were calculated for 
10 males and 10 females of E. massyae {collected in the 
same survey) which were fixed, preserved, and measured 
in the same way as the new species (table 1). Eledone 
gaucha is smaller antl has a narrower mantle and head. 
The arms are thinner, longer, and decrease in size while 
in £. massyae all arms are approximately the same length. 
The hectocotvlized arm is shorter and the web depth 
decreases troni the dorsal to the ventral surface in E. 
gaucha. while in E. massyae the web is shorter only 
between the ventral arms. The funnel organ is W shaped 
in E. gaucha, w shaped in £. massyae. The number of 
inner and outer gill lamellae is one unit lower and the 
arms sucker count two units higher in E. gaucha. .Arm 
sucker indices are similar in males of both species, but 
in females they are smaller in E. gaucha. Perhaps the 
best single diagnostic character to distinguish mature 
males of both species is the spermatophore, which is 



shorter and much thinner in £. gaucha. ExternalK, the 
best distinctive character is the arm length pattern. 

Eledunc caparti was described b\ .Adam (1950) based 
on five specimens, two males and three females collected 
in the equatorial west .Africa at depth ranging from 60 
to 170 m. No figures or tables were included in the 
original description. The decreasing arm length and web 
depth of £. caparti are similar to those of the new species. 
However, £. caparti does not have a supraocular cirrus, 
has enlarged suckers at the base of the lateral arms of 
the males, and the number ot suckers on the dorsal arms 
is almost double that in E. gaucha. The radula of £. 
caparti has an A2 seriation and the spermatophore is 
insufficiently described for comparisons. 

Eledone thysanophora was described by \'oss (1962) 
based on a single male specimen collected in a tide pool 
in western South Africa. The morphometric description 
is short, but the number of papillae on the tips of the 
non-hectocotylized arms and the structure of the sper- 
matophore, w ith the inner w all ot the horn portion lined 
with teeth, differentiate £. thysanophora irom the new 
species. 

Summary descriptions of £. cirrhosa (Lamarck, 1798) 
and £. moschata (Lamarck, 1798) are presented in Roper 
et al. (1984), and the species are compared b\ Rees 
(,1956). Both species can be distinguished from £. gaucha 
by several characters. Eledone cirrhosa has moderately 
short arms, a ridge along the mantle, non-hectocot\lized 
arms of males w ith a single row of compressed sucker- 
like cirri, and spermatophores with spines. Eledone mos- 
chata has subequal arms, 11 to 12 filaments on the outer 
hemibranch of the gills, big, sausage-shaped eggs 15 mm 
long, and a characteristic musk odor. 



ACKNOWLEDGEMENTS 

J. M. Andriguetto Filho and J. A. Alvarez Perez helped 
with collection of the specimens, G. L. Voss and an anon- 
ymous reviewer improved dramaticalK the final version, 
and E. G. Rios stimulated m\ interest in mollusks. To all 
of them m\' sincere gratituile. 



LITERATI RE CITED 

.Adam, W. 1950. Notes sur les cephalopodes .\.\11. Deux nou- 
velles especes de la cote africaine occidentale. Bulletin 
Institut Ro\ al des Sciences Naturelles de Beigique 2(i(45): 
1-9. 

Haimovici, M. and J. M .Andriguetto Fo. 1986. Celalopodes 
costeiros capturados na pesca de arrasto do iitoral sul do 
Brasil .Archives de Biologia e Tecnologia. Parana 29(3): 
473-495. 

Levi, J. A., M. Haimovici. and M. B. Concei^ao. 1985. Car- 
acteriza9ao eletofortica de dois morfotipos do genero Ele- 
done (Cephalopoda: Octopodidae). Resumes do XII Con- 
gresso Brasileiro de Zooiogia. (lampinas. 27 Jan to 1 Feb. 
1985:39. 

Palacio, J. F. 1977, .A stud\ of the coastal ceplialopods from 
Brazil with reference to Brazilian zoogeography. Ph.D. 
thesis, University of Miami, 311 p. 



M. Haimovici, 1988 



Page 87 



Palacio, J. F. 1978. Vosseledorie charrua. a new Patagonian 
ceplialopod (Octopodidae) with notes on related genera, 
bulletin of Marine Science 28(2):282-296. 

Rees, W J. 1956. Notes on the European species ol F.lcdone 
with special reference to eggs and larvae. Bulletin British 
Museum of Natural Histor\ .3(6):283-292, pis. 9-10, 

Roper, C. F. E., M. J. Sweene>, and C. E. Nauen. 1984. FAO 
species catalogue, Vol. 3. Cephalopods of the world. FAO 
Fisheries Synopsis No. 125, 3:227. 



Roper, C. F. E. and G. L. Voss. 1983. GuideHnes for taxonomic 
descriptions of cephalopod species. Memoirs of the Na- 
tional Museum of Victoria 4449-63. 

Voss, G. L. 1962. Soutli African cephalopods. Transactions of 
the Royal Societ)' of South Africa 36(Part 4):245-272. 

Voss, G. L. 1964. A note on .some cephalopods from Brazil 
with a description of a new species of octopod: Eledone 
massyae. Bulletin of Marine Science of the Gulf and Ca- 
ribbean 14:511-516. 



THE NAUTILUS 102(2);88, 1988 



Page 88 



News and Notices 



.54TH ANNUAL MEETING OF THE AMERICAN 
MALACOLOGICAL UNION 
CHARLESTON. SOUTH CAROLINA 
RADISSON FRANCIS MARION HOTEL 
June 19-24, 1988 

The 54th annual meeting of the American Malacological 
Liiioii \\ ill be held June 19-24, 19SS in (Charleston, South 
Carolina. Charleston is a historical cit\, many parts of 
which have been beautifully restored, as has the Radisson 
Francis Marion Hotel, which is located downtown, with- 
in walking distance of many restaurants, shops and other 
attractions. Charleston is easily accessible both by air and 
b\ interstate highway. 

Three symposia are planned: Applications of Nucleic 
Acid Techniques to the Study of Molluscan Evolution, 
convened b> Dr. M.G. Harasewych, Department of In- 
vertebrate Zoology, National Museum of Natural His- 
tory, Smithsonian Institution; S\ stematics and Evolution 
of Non-marine Mollusks, convened by Dr. Robert 
Hershler, Department of Invertebrate Zoology, National 
Museum of Natural History, Smithsonian Institution; and 
History of Malacolog\, convened by Dr. W. Backhuys, 
E.J. Brill, Inc Leyden, The Netherlands. 

In addition to these ssmposia, contributed papers and 
poster presentation, scheduled events w ill include a tour 
of historic Charleston, guided field trips to terrestrial and 
marine molluscan communities, an auction to benefit the 
s\ inposium fund, and a bantiuet. 

For further information, please contact: 

Richard E. Petit 

President, AMU 

P.O. Box 30 

North Mvrtle Beach, SC 29582 USA 

Telephone (803) 249-1454 



16TH CONCHOLOGISTS OF AMERICA 

CONVENTION 

FORT MYERS, FLORIDA 

SHERATON HARBOR HOUSE HOTEL 

July 11-15, 1988 

The 16th convention of the Conchologists of ,\merica is 
scheduled for Jul\ 11-15, 1988. This con\ention will be 
headquartered in the Sheraton Harbor House Hotel in 
Fort M\ers, Florida, and hosted b\ the Southwest Florida 
Concliologist Societ\ . Con\ention acti\ ities are centered 
around informative, shell-related programs, fascinating 
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to the Sarasota Fossil Pits is scheduled for Saturda\ , Jul\- 
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Pre-registration forms and packets for the Convention, 
Hotel and Field Trip reservations are available from: 

Gene Herbert 

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or 

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2265 West Gulf Drive # 240E 
Sanibel, FL 33957 
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THE NAUTILUS 



Volume 102, Number 3 
August 29, 1988 
ISSN 0028-1344 

A quarterly devoted 
to malacology. 



Marine Biological Laboratory . 
LIBRARY 

SEP 6 1988 i 

Woods Hole, Mass. 




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



ASSOCIATE EDITOR 
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.■\merican Malacologists, Inc. 
P O Box 2255 
Melbourne, FL 32902 



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Department of Mollusks 
Delaware Museum of 
Natural Histor\ 
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Wilmington, DE 19807 

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Department of 

Living Invertebrates 

The .\merican Museum of 

Natural History 

New York, NY 10024 

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Suffield. CT 06078 

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Division of Mollusks 
National Museum of 
Natural History 
Smithsonian Institution 
Washington, DC; 20560 

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Division of Mollusks 
National Museum of 
Natural History 
Smithsonian Institution 
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Museum of Comparative Zoology 
Harvard I'niversity 
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% Department of Mollusks 
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THE €9 N An^ LJicaiXio/ i L U S 



CONTENTS 



j LIBRARY 

SEP 6 1988 



Volume 102, Number 3 
August 29, 1988 
ISSN 0028-1344 



Woods Hole, Mass. 



Timothy M. Askew 



A new species of pleurotomariid gastropod from the 
western Atlantic 



89 



IM. G. Harasewych 
Shirley A. Pomponi 
Timothy M. Askew 



Spongivory in pleurotomariid gastropods . 



92 



James H. McLean 
Ricardo Silva Absalao 
Renato Luiz 

dos Santos Cruz 



A new species of Macrarene (Turbinidae: Liotiinae) from 
Brazil 



99 



Geerat J. Vermeij 
Timothy M. Collins 



Nerita fortidentata, a new gastropod from the Neogene of 
Panama, with comments on the fossil record of Nerita in 
tropical America 



102 



Anthony D^Attilio 
Barbara W. Myers 



A new species of Favartia from the eastern Pacific 
(Gastropoda: Muricidae) 



106 



William K. Emerson 
Salter E. Sage III 



Conus baccatus G. B. Sowerby III, 1877: A Panamic 
faunal constituent 



110 



Kenneth J. Boss 



References to Molluscan Taxa Introduced by Linnaeus in 
the Systema Naturae (1758, 1767) 



115 



Mark E. Gordon 



Frederick Benjamin Isely: Biographical Sketch and 
malacological contributions 



123 



William F. Adams 
Andrew G. Gerberich 



Rediscovery of Planorbella magnifica (Pilsbry) in 
southeastern North Carolina 



125 



Donald M. McKinstry 



Bite by Octopus joubini: a case report 127 



Oscar J. Polaco 
Wolfgang Mendl 



Occurrence of mites in Mexican land snails 129 



Richard E. Petit 



Axelella, new name for Olssonella Petit, 1970, a 
preoccupied taxon (Mollusca: Cancellariacea) 



130 



THE NAUTILUS 102(3):89-91, 1988 



Page 89 



A New Species of Pleurotomariid Gastropod 
from the Western Atlantic 



Timothy M. Askew 

Harbor Branch Oceanographic 
Institution, Inc. 
5600 Old Dixie Highway 
Fort Pierce, FL 34946, USA 



ABSTRACT 

Perotrochus charlestonensis, a new species of pleurotomariid, 
is described from off the coast of South Carolina. This is the 
12th Recent pleurotomariid taxon to be described from the 
western Atlantic. Its habitat is described, environmental data 
are provided, and comparisons made with closely related con- 
geners. 

Key words: Gastropoda; pleurotomariid; Perotrochus; slit shells; 
western Atlantic; JOHNSON-SE,\-LINK 



INTRODUCTION 

Since the discovery of the first living species of the pre- 
dominant!) Mesozoic and Paleozoic family Pieuroto- 
mariidae in the western Atlantic over a century ago 
(Fischer & Bernard!, 1856), 24 Recent species and sub- 
species have been described, usually on the basis of one 
or a very few specimens. The habitat of these animals, 
generally steep-wailed, hard substrates at depths in e.xcess 
of 100 meters, accounts for their infrequent collection 
by such methods as trawling, dredging, and grab sam- 
pling, and, therefore, for the paucity of data on the 
biology and distribution of most species. Since the pub- 
lication of a review of the Recent pleurotomariids that 
included six species from the West Indies (Bayer, 1966), 
three species (Bayer, 1967; Rios & Mathews, 1968; Leme 
& Penna, 1969) and two subspecies (Okutani & Goto, 
1983, 1985) have been described from the western At- 
lantic. 

Another new species of pleurotomariid, described 
herein, was collected while conducting fish population 
studies approximately 90 nautical miles east of Charles- 
ton, South Carolina, utilizing the submersible JOHN- 
SON-SEA-LINK I (Harbor Branch Oceanographic In- 
stitution, Inc., Fort Pierce, Florida). Bottom topography 
at the stud\ area was extremely rugged, making sampling 
by any other means difficult. 

SYSTEMATICS 

Family Pleurotomariidae Swainson, 1840 
Genus Perotrochus P. Fischer, 1885 



Perotrochus charlestonensis new species 

(figure 1) 

Description: Shell (figure 1) moderately large (maxi- 
mum diameter 87.4 mm, minimum diameter 80.1 mm, 
height 73.0 mm), broadly turbiniform, very thin, fragile; 
spire angle 89°, spire slightly convex in profile; proto- 
conch of 1.0 whorls, translucent, glassy; transition to te- 
leoconch marked by axial costae, with selenizone ap- 
parent by second postnuclear whorl; teleoconch of 8'/3 
whorls; early whorls nearly flat-sided, becoming pro- 
gressively more inflated; selenizone near suture in early 
whorls, shifting to slightly below mid-whorl by fifth post- 
nuclear whorl; anal slit depth at upper margin 89°, at 
lower margin 57°; anal slit width 4 mm; suture adpressed; 
periphery rounded; base inflated, convex, non-umbili- 
cate; nacreous umbilical callus extending '/6 the distance 
from axis to periphery; spiral sculpture of 21 uniformly 
sized spiral cords between suture and anal slit, 20 cords 
of variable thickness between anal slit and periphery, 40 
cords along base; selenizone with 0-5 broad cords, num- 
ber increasing with shell size; axial sculpture of weak 
nodes on early whorls (88 on fourth postnuclear whorl), 
forming cancellate sculpture; axial sculpture decreasing, 
sculpture limited to spiral cords by sixth postnuclear whorl 
above selenizone, and seventh postnuclear whorl below 
selenizone; aperture broadly ovate; columellar lip slightly 
thickened, weakly recurved; color creamy white with 
diffuse brownish orange axial streaks and blotches; na- 
creous layer visible through porcellaneous layer, creating 
iridescent hue; color lighter on base than on dorsal sur- 
face; selenizone margins with cream colored lines most 
evident on penultimate and body whorls; aperture na- 
creous, iridescent; operculum multispiral (7 whorls), 
horny, brownish-yellow, translucent; soft parts unknown. 

Type localitv: 90 nautical miles east of Charleston, South 
Carolina (32°43'80"N, 78°05'60"W), in 213 m, R/S 
JOHNSON-SEA-LINK I, dive 1250, August 6, 1982. 

Holotype: USNM 859961, maximum diameter 87.4 mm. 

Etymology: Named after the t\pe localit\ , commonly 
referred to as the Charleston Lumps. 



THE NAUTILUS, Vol. 102, No. 3 




Figure 1. Perotrochus charlestunensis new species. Apertural, right lateral, apical, and basal views of the holotype (USNM 859961 ), 
off Charleston, South Carolina (32°43'80"N, 78°05'60"W), in 213 m, maximum shell diameter 87,4 mm. 



Ecology: This species is known only from the type lo- 
cality, an area of extremely rugged terrain where the 
bottom topograph}' consists of steep, large hills and val- 
leys. Topographical features (figure 2) consist of a pave- 
ment of relithified phosphorite and fibrous concretionary 
apatite composed of calcium phosphate and other min- 
erals (Manheim et ai, 1980). This pavement, which ranges 
in thickness from 10 cm to almost a meter, has been 
undermined in some areas, causing large pieces to break 
off and fall down-slope forming rubble and boulder zones. 
Hills range in height from several meters to about 30 
meters. Valleys contain sand composed primarily of brown 



to black phosphorite. This area was formed during the 
middle Tertiary and has remained stable since the Mio- 
cene (Baturin, 1982). 

Marine life in the vicinity indicates an area of high 
productivity resulting from warmer Gulf Stream waters. 
Large snowy grouper [Epinephehts niveatus (Valen- 
ciennes, 1828)] and blue-lined tile fish [Caulolatilus mi- 
crops (Goode & Bean, 1878)] are abundant in the area 
together with many species of small, deep-reef fish, which 
generally inhabit rocky terrain. Common invertebrates 
include basket and brittle stars, sea urchins, solitary and 
colonial anemones, solitar\ corals, arrow, spider, and gal- 



T. M. Askew, 1988 



Page 91 



RCXKSOULOER LECX3E 




HOCK RUBBLE 



'-a.-.. 



i^^, SAND PEBBLE 



BARREN SAND 



Figure 2. Cross-section of bottom topography at collection 
site. Hills range in height from 3 to 30 m. 

atheid crabs, barrel and encrusting sponges, and hy- 
droids. A pink featherlike hydroid covers many of the 
broken boulders along the ridge tops. Prevalent gastro- 
pods include Perotrochus amabilis (Baser, 1963), Cal- 
liostoma sayana (Dall, 1889), Stenorhytis pernobilis 
(Fischer & Bernardi, 1857), Aurinia gouldiana (Dall, 
1887), and Pterynotus phaneus (Dall, 1889). 

Remarks: Perotrochus charlestonensis is a member of 
the species complex consisting of P. midas Bayer, 1966, 
P. pyramiis Bayer, 1967, P. africanus (Tomlin, 1948), 
P. teremachii (Kuroda, 1955), P. tangaroana Bouchet & 
Metivier, 1982, and an undescribed species from off 
northwestern Australia (Group B, Bayer, 1966:745). All 
are characterized by having large, thin shells with in- 
flated whorls and proportionally large, broadly ovate ap- 
ertures. This new species most closely resembles P. af- 
ricana and Perotrochus sp. (Bayer, 1966; fig. 29) from 
Japan, but differs from these taxa in having a thinner 
shell w ith more inflated whorls, and a more convex profile 
of the spire. Perotrochus africanus has a more stepped 
spire, a more strongly recurved and thicker columella, 
and a broader umbilical callus (Vi distance from axis to 
periphery). Of the western Atlantic species, P. charles- 
tonensis is most similar to P. pyramus. but is more than 
twice the size, and is much higher-spired. Perotrochus 
charlestonensis also somewhat resembles Perotrochus 
midas, but lacks its characteristic flat, blunt spire and 
angular periphery. Perotrochus charlestonensis occurs 
in shallower water (213 m) than either P. pyramus (420- 
648 m) or P. midas (600-770 m). 

ACKNOWLEDGEMENTS 

I thank the following persons, without whose help and 
guidance this paper would not be possible: Dr. Richard 
Cooper, University of Connecticut; Dr. Roger Theroux 
and Joseph L zmann. National Marine Fisheries Services, 
Woods Hole, Massachusetts; Dr. Walter Nelson and El- 



mer Gutherz, National Marine Fisheries Service, Pas- 
cagoula, Mississippi; and Tom Smoyer, Harbor Branch 
Oceanographic Institution photographer, for his excel- 
lent photographs. This is Harbor Branch Oceanographic 
Institution Contribution No. 642. 

LITERATURE CITED 

Baturin, G. N. 1982. Phosphorites on the sea floor — origin, 
composition and distribution. Developments in Sedimen- 
tology 33:111-112. 

Bayer, F. M. 1963. A new pleurotomariid gastropod trawled 
in the Straits of Florida by R/V Gerda. Bulletin of Marine 
Science of the Gulf and Caribbean 13(3):488-492. 

Bayer, F. M. 1966. New pleurotomariid gastropods from the 
western Atlantic, with a summary of the Recent species. 
Bulletin of Marine Science 15(4):737-796. 

Bayer, F. M. 1967. Another new western Adantic pleuroto- 
marian gastropod. Bulletin of Marine Science 17(2):389- 
397. 

Bouchet, P. and B. Metivier. 1982. Living Pleurotomariidae 
(Mollusca: Gastropoda) from the South Pacific. New Zea- 
land Journal of Zoology 9:309-318. 

Dall, VV. H. 1887. [A letter containing notes on Antillean 
mollusks.] Conchologist's Exchange 2(1):9-10. 

Dall, W. H. 1889. Reports on the results of dredging ... in 
the Gulf of Mexico (1877-1878) and in the Caribbean Sea 
(1879-1880), by the U.S. Coast Survey steamer "Blake" 
. . . XXIX. Report on the Mollusca. Part II. — Gastropoda 
and Scaphopoda. Bulletin of the Museum of Comparative 
Zoology, Harvard 18:1-492, pis. 10-40. 

Fischer, P. 1885. Manuel de conchyliologie et de paleonto- 
logie conchyliologique. Histoire naturelle des mollusques 
vivants et fossiles. Fascicule 9:785-896. Libraire F. Savy, 
Paris, xxiv -I- 1369 p., 23 pis. (1887). 

Fischer, P. and A. C. Bernardi. 1856. Description dun pleu- 
rotomaire vivant. Journal de Conchvliologie 5:160-166, 
pi. 5. 

Fischer, P. and A. C. Bernardi. 1857. Descriptions d'especes 
nouvelles. Journal de Conchyliologie 5:292-300, pis, 8, 9. 

Kuroda, T. 1955. A new Pleurotomaria from Japan with a 
note on a specimen of P. rumphii Schepman collected 
from Taiwan. Venus 18(4):21 1-221, pis. 8, 9, 

Leme, J. P. L. and L. Penna. 1969. Ocorrencia de Mikado- 
trochus no Brasil com descri^ao de uma nova especie. 
Papeis Avulsos de Zoologia (Sao Paulo) 22(21 ):225-230. 

Manheim, F. T., R. M. Pratt, and P. F. McFarlin. 1980. Com- 
position and origin of phosphorite deposits of the Blake 
Plateau. The Society of Economic Paleontologists and Min- 
eralogists, Special Publication No. 29:117-137. 

Okutani, T. and Y. Goto. 1983. A new subspecies of Adanson's 
slit shell from Bermuda. Venus 42(4):305-311. 

Okutani, T. and Y. Goto. 1985. A new subspecies of Pero- 
trochus quoyanus from Bermuda. Venus 44(1):27-31. 

Rios, E. C. and H. R. Mathews. 1968. Nova especie de Pleu- 
rotomariidae do Brasil (Mollusca: Gastropoda). Arquivos 
Estaceo de Biologia Marinha da Universidade Federal do 
Ceara 8(l):65-68. 

Tomlin, J. R. le B. 1948. A new species of Pleurotomaria. 
Journal of Conchology 23:2, pi. 1. 



THE NAUTILUS 102(3):92-9S, 1988 



Page 92 



Spongivory in Pleurotomariid Gastropods 



M. G. Harasewych 

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



Shirley A. Pomponi 
Timothy M. Askew 

Harbor Branch Oceanographic 
Institution, Inc. 
5600 Old Dixie Highway 
Ft. Pierce, FL 34946, USA 



ABSTRACT 

Direct in situ observations of feeding together with analyses 
of the gut contents of Perutruchus ruidas and P. amabilis in- 
dicate that these species feed predominantK and selectively on 
sponge tissue. Foraminiferal and diatom tests previously re- 
ported in the gut contents of pleurotomariids are derived from 
planktonic sediment coating the surfaces of sponges and do not 
significantK contribute to the nutrition of these gastropods. The 
family Pleurotomariidae represents an adapti\e radiation to 
spongivory that most likely occurred after the divergence of 
the herbivorous Scissurellidae and Haliotidae from the pleu- 
rotomariid progenitor in the late Paleozoic, but prior to the 
appearance of umbilicate pleurotomariids with deep anal ca- 
nals in the Upper Jurassic. 

Key words: Pleurotomariidae; diet; sponges; spongivory; Pero- 
trochus; Mikadotrochus, Entemnutrochus. 



INTRODUCTION 

Since the discovery of the first living species of the family 
Pleurotomariidae in the mid-nineteenth century, nu- 
merous papers have been published on various aspects 
of the biology and anatomy of these, the most primitive 
living gastropods (e.g., Bouvier & Fischer, 1899, 1902; 
Woodward, 1901; Fretter, 1964, 1966; Yonge, 1973; 
Hickman, 1984a, b). As pleurotomariids are predomi- 
nantly restricted to hard substrates and bathyal depths 
in the Recent fauna, sampling has pro\ed difficult and 
most published observations are based on limited ma- 
terial, often poorly preserved. 

The common occurrence of sponge spicules, forami- 
niferal tests, and diatoms in the alimentary systems and 
fecal pellets of pleurotomariids has been variously in- 
terpreted as being indicative of a spongivorous diet con- 
sisting of a single species of sponge (Woodward, 1901: 
252); a diet consisting principally of sponges (Thiele, 



1935:1129; Hyman, 1967:.360; Yonge k Thompson, 1976: 
52); a diet of encrusting invertebrates, predominantly 
sponges (Hickman, 1984a:29); or detrital feeding and 
vegetarian diet (Fretter & Graham, 1976:1). The pres- 
ence of sponge spicules in the gut contents, however, is 
not necessarily evidence of spongivory. Although mem- 
bers of the pleurotomariacean family Haliotidae are 
known herbivores (Leighton, 1961; Leighton & Boo- 
lootian, 1963; Shepherd. 1973; Fretter & Graham, 1976: 
6), Leighton and Boolootian (1963:229) reported sponge 
spicules, comprising 1-2% of the gut content \olume, in 
25-50/c of the specimens of Haliotis cracherodii they 
examined. Similarly, Herbert (1987:289) identified sponge 
spicules, foraminiferans, annelid setae, and crustacean 
remains in the alimentary systems of members of the 
Solariellinae (Trochidae), a group adapted to feeding on 
superficial and interstitial detritus. Three species of pleu- 
rotomariids maintained in aquaria have been reported 
to feed on a wide variety of foods, including starfish, 
bivalve meat (Arakawa et al., 1978) and sliced, raw fish 
(Matsumoto et al, 1972; Sekido et al, 1976). 

The increasing use of research submersibles in deep 
sea investigations has made possible in situ observations 
on the feeding of two species of pleurotomariids. These 
observations, supported by gut content analyses of col- 
lected voucher material, form the basis of this report. 

MATERIALS AND METHODS 

During the course of numerous di\es aboard the research 
submersible JOHNSON-SEA-LINK-II throughout the 
northern and central Bahamas, li\ ing Perotrochus midas 
Bayer, 1966, were observed on 16 occasions at depths 
ranging 670 to 853 meters. Observations were noted and 
specimens photographed when practical, using a BEN- 
THOS 35 mm camera, with 85 mm lens. Two of the 



Figures 1, 2. Perotrochus midas Bayer, 1966, and Strongylophora hartmani \'an Soest, 1981, in situ. 1. Perotrochus midas feed- 
ing on Strongylophora hartmani. JSL-II dive 1501, #2, west end of Island at Ciouldings Cay, New Providence Island, Bahamas, 
25°00'00"N, 77°34'06"\V. in 766 m. October 20, 1987. Specimen not collected Note area of sponge consumed by snail. 2. Perotrochus 
midas and Strongylophora hartmani along steep wall, JSL-II dive 1505, Chub Cay, Berry Islands, Bahamas, 25°22'22"N, 77°50'25"W, 
in 777 m. October 23, 1987. Specimen not collected. Note layer of planktonic sediment. 



M. G. Harasewvch et al, 1988 



Page 93 




Page 94 



THE NAUTILUS, Vol. 102, No. 3 



specimens were collected, one together with its prey, 
using the hydraulic arm and clamshell scoop. These spec- 
imens were fixed in formalin and preserved in 70% eth- 
anol. 

Fifty-six specimens of Perotrochus arnabilis (Bayer, 
1963) were observed and collected during four dives 
aboard the research submersible Nekton Delta on the 
"Charleston Lumps", an area of rough bottom topog- 
raphy approximately 90 miles east of Charleston, South 
Carolina (32°43'78"-32°44'90"N, 78°05'68"-78°06'00"\V) 
(Askew, 1988:91), at depths ranging from 200 to 230 m. 
Samples were collected by suction through a 2 inch (51 
mm) diameter hose. Specimens were frozen on dry ice 
and maintained at — 80°C. 

After identification of the intact sponge on which a 
specimen of P. midas was feeding, a spicule sample for 
SEM examination was prepared by digesting a portion 
of the sponge in 70% nitric acid (HNO5) until only sili- 
ceous material remained. Two specimens of P. midas 
and three specimens of P. amalnlis were dissected, and 
sections of the esophagus between the esophageal valve, 
situated behind the buccal mass (Fretter, 1964:181, fig. 
5, ov; 1966:609, fig. 2, v), and the long sphincter at the 
opening of the stomach (Fretter, 1966:609) were excised. 
Also removed were portions of the rectum (Fretter, 1966: 
fig. 1, r; 1964: fig. 2, r). These sections were teased apart 
and examined under a dissecting microscope, then treat- 
ed with warm diluted bleach (1-2% sodium hypochlorite, 
NaOCl) to dissolve organic material while leaving the 
siliceous and calcified remains. The preparations were 
rinsed in distilled water, filtered through 0.4 ^m Nucle- 
pore membrane filters, and the filters mounted directly 
onto SEM stubs. Samples were coated with carbon and 
gold, and photographed using a Hitachi S-570 scanning 
electron microscope. A transverse section of the intact 
sponge was critical point dried prior to SEM examination. 

The following voucher material is deposited at the 
National Museum of Natural History: 

Perotrochus midas: Specimen 2. USNM 857097, JSL-II 
dive 1501, #2, West end of Island at Gouldings Cay, 
New Providence Island, Bahamas, 25°00'00"N, 
77°34'06"W, in 766 m. October 20, 1987. 

Perotrochus amabilis: Specimens 1, 2, and 3. USNM 
846900, DELTA Dive 560, 129 km due east of Charles- 
ton, South Carolina, USA, 32°43'95"N, 78°05'72"W, in 
198-210 m. May 3, 1987. 



RESULTS 

Of the 16 sightings of Perotrochus midas. this species 
was observed in close proximity to, or actively feeding 



upon, the sponge Strongylophora hartmani Van Soest, 
1980 (class Demospongiae, order Haplosclerida) on six 
occasions (figures 1, 2). Two specimens of P. midas were 
collected, one (specimen 1) together with the sponge on 
which it was feeding. The esophagus of both snails was 
distended and full of sponge tissue. Comparisons of spic- 
ules taken from the esophagus of both specimens of Per- 
otrochus midas (specimen 1, figure 5) with those of Stron- 
gylophora hartmani (figures 3, 4) confirmed the identity 
of the prey species. The rectum of specimen 1 contained 
spicules of S. hartmani (figures 6, 7) as well as spicules 
tentativeK attributed to a species of PachastreUa (figure 
7). The rectal contents of specimen 2 included spicules 
of S. hartmani, but spicules of an unidentified sponge 
belonging to the order Hadromerida (figure 8) comprised 
an estimated 80% of the \olume. The contents of the 
esophagus and rectum from both specimens of P. midas 
consisted almost entirely (estimated >95% by volume) 
of sponge spicules, with foraminiferal and ostracode tests 
accounting for most of the other identifiable remains. 

Of the 53 specimens of Perotrochus amabilis observed 
and collected, the majority were on phosphorite blocks 
ranging in size from several centimeters to over a meter 
in length. Only three specimens were on or near a sponge. 
In all three instances, the sponge, which was small and 
roughly spherical (<6 cm diameter), could not be col- 
lected. Examination of the contents of the esophagus and 
rectum of these three specimens revealed a more het- 
erogeneous assemblage of sponge spicules (about 80%), 
diatoms, and foraminiferal tests (about 20%) (figures 9, 
10). In no instance could more than an estimated 50% 
of the contents of either the esophagus or the rectum be 
attributed to a single species of sponge. 

DISCUSSION 

Direct feeding observations as well as gut content anal- 
yses indicate that Perotrochus midas feeds on the sponge 
Strongylophora hartmani b> rasping large, deep depres- 
sions (>2 cm diameter, >1 cm depth) in its surface 
(figure 1). In each of the two specimens dissected, the 
voluminous esophagus was distended, and contained a 
corresponding volume (2-3 cm') of sponge tissue. Rectal 
contents also consisted almost exclusively of sponge spic- 
ules, though from species belonging to the orders Cho- 
ristida and Hadromerida. The most abundant groups of 
deep water sponges (in terms of biomass) in the tropical 
western Atlantic and Caribbean are the orders Haplo- 
sclerida and Choristida, with the most common species 
often being highly silicified (Pomponi, unpublished ob- 
servations). Although six of the 16 sightings of P. midas 
were on or near a haplosclerid sponge, it is unclear 



Figures 3, 4. Strongylophora fiartmani Van Soest, 1981. 3. Transverse section, outer surface at top (SEM), scale bar = 300 ^lm. 
4. Nitric acid spicule preparation (SEM), scale bar = 200 urn. Figure 5. Perotrochus midas, contents of mid-esophagus of spetimen 
1, showing high concentration of Strongylophora hartmani spicules (SEM), scale bar = 200 ^m. Figures 6, 7. Perotrochus midas, 
rectal contents of specimen 1. 6. Strongylophora hartmani spicules (SEM), scale bar = 200 ^m. 7. Spicules of S. hartmani as well 
as of ?Pachastrella sp. (arrows) (SEM), scale bar = 200 ^m. Figure 8. Perotrochus midas, rectal contents of specimen 2, with 
spicules of unidentified sponge, order Hadromerida (arrows) (SEM), scale bar = 200 ^m. 



M. G. Harasewych et al, 1988 



Page 95 




Page 96 



THE NAUTILUS, Vol. 102, No. 3 




Figures 9, 10. Perotrochus amabilis (Bayer, 1963), contents of mid-esophagus of specimen collected 90 miles due east of Charleston, 
SC, in 210 m. 9. Spicule of an unidentified species, order Poecilosclerida (SEM), scale bar = 20 ^m. 10. Spicules of Strongylophora 
sp. (small arrow) and an unidentified species, order Choristida or Spirophorida (large arrows) (SEM), scale bar = 100 ^m. 



whether prey species are selected, or mereK reflect the 
relative abundance of sponges in a nutrient-poor envi- 
ronment. In either case, available evidence indicates that 
sponges comprise the major component of the diet of P. 
midas, and that the small amounts (<5%) of foraminif- 
eral and ostracode tests as well as other planktonic sed- 
iment found in the esophagus and rectum of this species 
were present on the surface of sponges (figures 1, 2), and 
probably do not contribute significantly to its nutrition. 

Although far more specimens of Perotrochus amabilis 
were observed and collected, fewer were in the proximity 
of a sponge. Due to the smaller size of this species and 
the inability to collect the sponges, no direct observations 
of feeding can be documented. E.xamination of the con- 
tents of the esophagus and rectum of three specimens 
indicate that sponge spicules comprise approximately 
80% of the bleach insoluble mass, the remaining fraction 
consisting mostly of foraminiferal tests and diatom frus- 
tules. Fretter (1964:182) reported similar contents in the 
stomach of this species and concluded that the animal 
was a microphagous scavenger. We suggest that P. ama- 
bilis, like P. midas, feed exclusively on sponges, but, due 
to their smaller size and correspondingly shorter snout, 
these animals are not able to penetrate as deeply into the 
tissues of the sponge, and feed on surface tissues covered 
with deposits of planktonic sediment, thus accounting 
for the higher proportion of diatoms and foraminifera 
in the gut. 

With a single exception (Barnard, 1963), all published 
reports on the gut contents of pleurotomariids (table 1) 
list sponge spicules as a major component. Woodward 
(1901:252) was the first to speculate that the distinctive 
radula, shared by all pleurotomariids, was adapted for 
spongivory, with the hooked teeth "tearing away great 



pieces of the sponge , and the brush teeth used to "rasp 
away some of the flesh from the spicules". The occur- 
rence of brush or filament-tipped teeth, long considered 
unique to Pleurotomariidae, in unrelated, sponge-feed- 
ing mesogastropods of the genus Seila. has led Hickman 
(1984a:35) to conclude that this tooth morpholog\ is a 
functional adaptation to sponge predation and not a phy- 
logenetically constrained, conservative, morphological 
feature. 

Although exceptions have been documented (e.g., Gra- 
ham, 1939; Perron, 1975), archeogastropods are gener- 
all\ considered to be herbivores (Yonge & Thompson. 
1976). This has led several authors (Yonge, 1973; Hick- 
man, 1984a) to imply that carnivory was not the original 
mode of feeding of pleurotomariids. Yonge (1973) fur- 
ther suggested that the change to a carnivorous diet may 
have been associated with the ecological shift of this 
family from shallow water reefs, which it inhabited dur- 
ing the Paleozoic and Mesozoic, to deeper water (>200 
m), hard substrates by the end of the Eocene. 

As members of all three Recent pleurotomariid genera 
have radulae with filament-tipped teeth, it would appear 
that spongivory in Pleurotomariidae predates the diver- 
gence of the genus Entemnotrochus Fischer, 1885, char- 
acterized by a deep, broad umbilicus and an anal slit 
extending nearK 180° back from the aperture, from the 
genera Perotrochus Fischer, 1885, and Mikadotrochus 
Lindholm, 1927, both with shallow (<90°) anal slits and 
without umbilici. Inclusion of the genus Conotomaria 
Cox, 1959, which is also characterized by having shells 
w ith deep anal slits and umbilici, in the same clade as 
Entemnotrochus would date this divergence, and there- 
fore the adaptation to spongivory, prior to the Late Ju- 
rassic (Knight et al., 1960). The restriction of the family 



M. G. Harasewych et ai, 1988 


Page 97 


Table 1. Published reports on the contents of the alimentary system and fecal pellets of pleurotomariid gastropods. 


Species Food Reference 



Mikadotrochxis heyrichi 
Mikadotrochus hirasei 

Perotrochus africanus 
Perotrochus amahilis 

Perotrochus amahilis 
Perotrochus midas 



Sponge spicules, order Poecilosclerida 
Sponge spicules, orders Haplosclerida, 

Poecilosclerida, and Hadromerida 
Amorphous mass with few tiny foraminiferans 
Sponge spicules, foraminiferans, diatoms, and 

algal fragments 
Sponge spicules, foraminiferans, and diatoms 
Sponge spicules, foraminiferans, and diatoms 



Woodward, 1901:252 
Arakawa ei ai. 1978 

(table 1) 
Barnard, 1963:156 
Fretter, 1964:182 

Present study 
Present study 



to bathyal depths at the time of formation of the psy- 
chrosphere, the lower, cooler layer of a two-layer ocean 
(Bruun, 1957; Benson, 1975) was accompanied by a con- 
siderable reduction in diversity, compared to Paleozoic 
and Mesozoic fauna (Woodward, 1885). 

Many sponges contain secondary metabolites that are 
toxic (Green, 1977; Bakus & Thun, 1979). Nevertheless, 
mollusks, echinoderms, fishes, and marine turtles are nat- 
ural predators of sponges, with nudibranchs being among 
the most species-specific spongivores (Sara & Vacelet, 
1973). Halichondria okadai (Kadota, 1922), the preferred 
pre\ of several species of nudibranchs, for e.xample, con- 
tains potent cytotoxic, antifungal, and tumor promoting 
compounds (Tachibana et ai, 1981; Fujiki et ai, 1987). 
Strongylophora hartnmni, the sponge species eaten by 
Perotrochus midas, contains puupehenone, a cytotoxic 
compound (Komoto et al.. 1987). It is possible that in- 
corporation of diet-derived toxic metabolites may confer 
some protection from predators upon pleurotomariids, 
but this hypothesis remains to be tested. 

ACKNOWLEDGEMENTS 

We thank Richard Cooper and Peter Auster of the 
NOAA National Undersea Research Program, University 
of Connecticut, Avery Point, for making submersible 
time available at the Charleston Lumps site. The assis- 
tance of M. Cristina Diaz with the identification of the 
intact sponge, and Susann Braden with the scanning elec- 
tron microscopy is gratefully acknowledged. This is Har- 
bor Branch Oceanographic Institution Contribution No. 
643. 



LITERATURE CITED 

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Engen, J. Clardy, Y. Gopichand, and F. J. Schmitz. 1981. 
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THE NAUTILUS 102(3):99-101, 1988 



Page 99 



A New Species of Macrarene (Turbinidae: Liotiinae) 
from Brazil 



James H. McLean 

Los Angeles County Museum of 

Natural Histor)- 

900 Exposition Boulevard 

Los Angeles, California 90007, USA 



Ricardo Silva Absalao 
Renato Luiz dos Santos Cruz 

Departamento de Zoologia, 
Instituto de Biologia, CCS. 
Universidade Federal do 
Rio de Janeiro 

Ilha do Fundao, Rio de Janeiro, 
Brasil 21940 



ABSTRACT 

The new species Macrarene digitate from the northeast Bra- 
zilian coast represents the first record of this principally eastern 
Pacific genus in the western Atlantic. The species had previ- 
ously been known from juvenile specimens reported as Liotia 
admirabilis E. A. Smith, the holotype of which is not a member 
of the subfamily Liotiinae. 



INTRODUCTION 

The species here described was first recognized as a mem- 
ber of the Brazilian fauna by Rios (1975, 1985), who 
referred it to Liotia admirabilis E. A. Smith, 1890. That 
species was described from the oceanic island of Saint 
Helena. According to the original description. Smith's 
species has a maximum dimension of I'A mm. The 15 
syntypes were examined at the British Museum (Natural 
History) (catalogue numbers 1889.10.1.1554-68) by the 
senior author in 1984 and found to be similar (although 
not clearly referable) to the skeneiform genus Parviturbo 
Pilsbry & McGinty, 1945, which is not a member of the 
Liotiinae. 

The Brazilian species was clearly undescribed, but its 
true generic affinity was not readily apparent because 
the specimens available to Rios, which have been ex- 
amined by McLean (figure 2), were not mature and the 
expression of the mature lip was impossible to determine. 
More recently, two larger specimens have come to light 
and it can now be maintained that the species has the 
characters of the genus Macrarene Hertlein & Strong, 
1951. 

Abbreviations for institutions are as follows: LACM, 
Los Angeles County Museum of Natural History; MORG, 
Museu Oceanografico, Universidade do Rio Grande, R.S., 
IBUFRJ, Instituto de Biologia, Universidade Federal do 
Rio de Janeiro. 



SYSTEMATICS 

Family Turbinidae Rafinesque, 1815 

Subfamily Liotiinae H. & A. Adams, 1854 

Shells of the subfamily are characterized by turbiniform 
to discoidal profiles, nacreous interiors, fine lamellar 
sculpture, intritacalx (calcified periostracum) in most 
genera, circular apertures, and multispiral opercula with 
calcareous beads. Radula like that of members of other 
turbinid subfamilies. 

Although previously treated by most authors as a full 
family, the Liotiinae have recently been ranked as a 
subfamily of Turbinidae by McLean (1987). 

Genus Macrarene Hertlein & Strong, 1951 

Type species (original designation): Liotia californica 
Dall, 1908. Recent, off Baja California, Mexico. 

Macrarene species are characterized by turbinate white 
shells, broad umbilici, and presence of axial ribs and 
spiral cords that form spines at their intersections. Spac- 
ing of the axial ribs increases in the final whorl. In some 
species the ribs then become more closely spaced in the 
final quarter whorl. The final lip is not thickened at 
maturity. 

Some Macrarene species reach relatively large sizes. 
The genus differs from Arene in lacking shell pigmen- 
tation and in having the spacing of the axial sculpture 
increasingly separated as the shell matures. The white- 
shelled genus Liotia is smaller and retains tight spacing 
of the axial sculpture. 

The white-shelled, new world Liotiine genera Mac- 
rarene, Liotia, and Cyclostrema differ as a group from 
those of the Indo-Pacific and Australasian regions in lack- 
ing the thickened mature lips that characterize the gen- 
era Bathtjiiotina Habe, 1961, Liotina Fischer, 1885, 
Dentarene Iredale, 1929, and Austroliotia Cotton, 1948. 



Page 100 



THE NAUTILUS, Vol. 102, No. 3 




Figures 1, 2. Macrarene digitata new species. 1. Holotype, IBUFRJ 1562. x 7.1. 2. Immature specimen, MORG 18.359. x 10.0. 



For further remarks on the Indo-Pacific group see McLean 
(1988). 

There are si.\ previously described species of Macra- 
rene: M. californica (Dall, 1908), M. cookeana (Dall, 
1908), M. diegensis McLean, 1964 (Pliocene); M. far- 
allonensis (A. G. Smith, 1952), M. lepidotera McLean, 
1970, and M. spectahilospina Shasky, 1970. All occur 
offshore in the tropical to temperate eastern Pacific. 

Macrarene digitata new species 
(figures 1, 2) 

Description: Shell small for genus, turbinate, white, 
interior weakly nacreous, ma.ximum diameter 6.7 mm, 
whorls 3.5, aperture only slightK ohliciue, final lip not 
thickened. Whorls circular in outline; final w horl in con- 
tact with previous whorl at tips of axial ribs. Shell surface 
marked by microscopic lamellar growth increments; in- 
tritacalx present. Protoconch diameter about 200 ^m. 
Suture deeply impressetl, first and second whorls rising 
above protoconch, third whorl descending, resulting in 
flat-topped profile for earlv whorls. F"irst teleoconch whorl 
nearly smooth (except for fine lamellar sculpture), second 
with about 20 strong axial ribs and nine, nearly equal 
spiral cords defining deep, rectangular pits. Inter.sections 
of axial and spiral sculpture produce sharpK- projecting 
spines that are slightly upturned adapicalK . Spiral cords 
of final whorl increasing to 12; axial ribs decreasing to 
15. Spines produced on final whorl by interaction of spiral 



and axial sculpture; spines with w eblike interconnections. 
Axial ribs narrow across umbilical wall, forming single 
descending row of sharp-pointed projections along in- 
nermost spiral cord. Operculum and radula unknown. 

Type locality: Off northeast coast of Brazil (03°59'N, 
49°35'W), 100 m, Brazilian Naval Research Vessel Al- 
mirante Saldaiilia. station 1913, May 6, 1968, 1 speci- 
men. 

Type material: Holotype (figure 1), IBUFRJ 1562. 
Height 5.0 mm, diameter 6.7 mm. The holot\pe is a 
dead collected specimen in good condition. Parat\pe, 
LACM 2377, off Cabo San Roque, Rio Grande do Norte, 
Brazil (04°30'S, 50°03'\\'), 146 m, Brazilian Naval Re- 
search Vessel Almirante Saldanha. station 1921, May 8, 
1968 (height 4.3 mm, diameter 6.6 mm). The paratope 
agrees with the holot)pe in size and sculptural details 
but is in subfossil condition with attached sedimentary 
deposits on the base. 

Referred material: MORG 18.359, 2 immature speci- 
mens [height 2.7, diameter 4.3 mm (figure 2); height 1.5, 
diameter 2.6] from Paripueira, Alagoas, Brazil, in beach 
drift collected b\ P. S. Cardoso, December, 1964; MORG 
20.620, 2 immature specimens (height 2.1, diameter 3.7 
mm; height 2.4, diameter 3.6 mm), Fernando de No- 
ronha Island, 6 m, collected b\ L. Barcellos, January, 
1979. 

Etymology: From the Latin digitatus. having fingers. 



J. H. McLean et at., 1988 



Page 101 



Remarks: Macrarene digitata is the smallest species of 
the genus described to date. However, it is not certain 
that the holot\pe is mature. Although this genus does 
not form a thickened lip, maturit) in other members of 
the genus is indicated by closer spacing of the axial ele- 
ments in the final quarter whorl of growth, as indicated 
in the original illustration of M. spectabilospina of Shasky 
(1970: fig. 2). The absence of such closer spacing of the 
axial element suggests that a quarter whorl of additional 
growth (and a substantial increase in diameter) is possible 
for M. digitata. 

Macrarene digitata is unique in the genus in having 
all elements of the spiral sculpture of similar strength, 
rather than having a strongly projecting peripheral 
carination. Such a sculptural distinction is not regarded 
as a generic level character because generic characters 
in the Liotiinae are more reliably based on apertural 
morpholog\', particularly the structure of the final lip. 

The most characteristic feature of this species is the 
fingerlike aspect of the projecting spines. It cannot easily 
be confused with any other member of the Liotiinae. 

ACKNOWLEDGEMENTS 

We are grateful to Prof. E. C. Rios for the loan of the 
referred material of this species. 



LITERATURE CITED 

Dall, W. H. 1908. Reports on the dredging operations off the 
west coast of Central America to the Galapagos, to the 



west coast of Mexico, and in the Gulf of California . . . 
,\1V. The Mollusca and Brachiopoda Bulletin of the Mu- 
seum of Comparative Zoology, Harvard University 43: 
205-487, pis. 1-22. 

Hertlein, L. G. and A. M. Strong. 1951. Eastern Pacific ex- 
peditions of the New York Zoological Society, XLIII. Mol- 
lusks from the west coast ol Mexico and Central America. 
Zoologica, Scientific Contributions of the New York Zoo- 
logical Society 36:67-120, pis. 1-11. 

McLean, J. H. 1964. New species of Recent and fossil west 
American aspidobranch gastropods. The Veliger 7:129- 
133. 

McLean, J. H. 1970. New species of tropical eastern Pacific 
Gastropoda. Malacological Review 2:115-130. 

McLean, J. H. 1987. Angariinae and Liotiinae — the primitive 
living trochaceans. .Annual Report of the Western Society 
of Malacologists 19:16. 

McLean, J. H. 1988. Two new species of Liotiinae (Gastropo- 
da: Turbinidae) from the Philippine Islands. The Veliger 
30:408-411. 

Rios, E. C. 1975. Brazilian marine mollusks iconography. 
Museu Oceanografico, Fundagao Universidade do Rio 
Grande, 331 p., 91 pis. 

Rios, E. C. 1985. Seashells of Brazil. Museu Oceanografico, 
Funda^ao Universidade do Rio Grande, 328 p., 102 pis. 

Shasky, D. R. 1970. New gastropod taxa from tropical western 
America. The Veliger 13:188-195. 

Smith, A. G. 1952. Shells from the bird guano of southeast 
Farallon Island, California, with description of a new species 
of Liotia. Proceedings of the California Academy of Sci- 
ences, Series 4, 27:383-387. 

Smith, E. A. 1890. Report on the marine Mollusca of the 
island of Sta. Helena. Proceedings of the Zoological Society 
of London 1890:247-322, pis. 21-24. 



THE NAUTILUS 102(3): 102-105, 1988 



Page 102 



Nerita fortidentata, a New Gastropod from the Neogene of 
Panama, with Comments on the Fossil Record of 
Nerita in Tropical America 



Geerat J. Vermeij 

Department of Zoology 
University of Maryland 
College Park, MD 20742, USA 



Timothy M. Collins 

Department of Geology and 

Geophysics 

Yale University 

New Haven, CT 06520, USA 



ABSTRACT 

Nerita fortidentata new species is described from the Neogene 
of Bocas del Toro, Panama It is most closely related to the 
Recent Caribbean \. fulgurans Gmelin, 1791. The fossil record 
and biogeography of tropical American Nerita are reviewed. 
At least two lineages of Nerita present in tropical America 
during the Tertiary have become restricted in the Recent fauna 
tn the Infln-Wf^t-Pacific region 



INTRODUCTION 

Neritid gastropods are prominent members of tropical 
rocky intertidai communities. Because ihe fossilization 
potential of tfiese animals is poor, little is known about 
tlie historical development of the genus Serita Linnaeus, 
1758. It was, therefore, of considerable interest to find 
an excellently preserved specimen of an apparently hith- 
erto unrecognized species from the Neogene of tropical 
America. Here we describe the new species and review 
briefly some biogeographically interesting aspects of the 
history of the genus Nerita in tropical America. 

METHODS 

Species of Nerita are distinguished conchologically by 
characters of shape, apertural dentition, external sculp- 
ture, form and sculpture of the parietal callus, and the 
shape and sculpture of the calcareous operculum. Two 
ratios are especially helpful in describing the overall shell 
form of Nerita (X'ermeij, 1973). The first is globosity, G, 
defined as the distance H, between the dorsal surface of 
the body whorl and the center of the parietal callus 
divided by the geometric mean betw een the shell's major 
diameter Dj and minor diameter D2: G = H,/(D,D2)'''. 
The second ratio is the degree of basal excavation. The 
plane of the parietal callus typically lies at an angle to 
the horizontal w hen the shell lies aperture-down on a 
flat surface. The greater the angle, the greater is the 
degree of basal excavation. An approximation of the de- 
gree of basal excavation is given by the ratio E = H2/ 



H,, where Ho is the distance from the horizontal plane 
on w hich the shell rests to the dorsal surface of the body 

whorl. 



SYSTEMATIC DESCRIPTION 

Class Gastropoda 
Subclass Prosobranchia 
Order Neritacea 
Family Neritidae 

Genus Nerita Linnaeus, 1758 

Type species: Nerita peloronta 1758, Recent, tropical 
Western Atlantic. 



Subgenus Theliostyla Morch, 1852 

Type species: Nerita albicilla Linnaeus, 1758, Recent, 
Indo- West-Pacific. 



Nerita (Theliostyla) fortidentata new species 
(figures 1, 2) 

Diagnosis: Shell thick, moderately globose (G = 0.59), 
base little excavated (E = 1.10), apex of spire barely 
raised above rest of shell. Outer lip very thick, inner edge 
with 12 strong teeth; two teeth nearest spire \er\ large 
and protruding, as is third tooth from abapical end of 
lip; columellar lip with tw^o strong centrally placed teeth; 
adapical portion of parietal region with a fold of about 
the same size and strength as adapical tooth, w hich curves 
into aperture; sculpture consisting of 21 regularK -spaced 
flat-topped smooth spiral cords, w hich Dare slightly from 
base to barely overhang incised interspaces about one- 
third the width of ribs; parietal callus small, its surface 
sculptured with about 10 strong ridges that bear up to 
3 large granules each; holotype shows faint radial color 
pattern of alternating continuous and discontinuous pro- 
socline bands of off white and grey-black; operculum 
unknown. 



G. J. Vermeij and T. M. Collins, 1988 



Page 103 




Figure 1. Nerita fortidentata new species, from Bocas del 
Toro, Panama, Holotype, USNM 423644, height 19.7 mm, 
Apertural view, showing thickened outer lip and enlarged teeth 
on adapical portion of outer lip. 

Holotype: United States National Museum number 
423644. Major diameter 21.3 mm, minor diameter 16.9 
mm, Hi 11.2 mm, H^ 12.3 mm, standard shell height 
19.7 mm (apex abraded), standard shell diameter 19.9 
mm, shell thickness at midpoint of outer lip 3.5 mm. 

Type locality: Panama, Province of Bocas del Toro, 
Archipelago of Bocas del Toro, Punta Robalo quadrangle. 
Island of Cayo Agua, eastern side about 400 meters south 
of Punta Nispero on the shoreline in clayey, tuffaceous, 
quartzose, blue-grey siltstones with dense shelly horizons. 
We have followed Woodring (1982) in referring to the 
Late Miocene-Pliocene deposits of the Bocas del Toro 
area as the Limones Formation. The true relationships 
between the Miocene-Pliocene of Bocas del Toro with 
respect to the Costan Rican Limones Formation and the 
Gatun Formation of the Canal Zone have not been elu- 
cidated. Laurel B>bell of the U.S.G.S. (personal com- 
munication) has assigned a preliminary age of Late Mio- 
cene to Early Pliocene to the locality from which the 
holotype of N. fortidentata was collected. This age de- 
termination is based on the presence of the calcareous 
nannofossils Sphenolithiis abies Deflandre, in Deflandre 
and Fert, 1954 (last occurrence middle Pliocene), and 
Discoaster brouweri Tan, 1927 (first occurrence middle 
Miocene). Thomas M. Cronin, U.S.G.S. (personal com- 
munication), examined the ostracode faunas from several 
adjacent localities of the same formation on Cayo Agua. 
He noted a remarkable similarity between the ostracodes 
from these samples and the ostracode fauna described 
by van den Bold (1967) from the type Gatun Formation 
of the Canal Zone. On the basis of these similarities he 
suggests a preliminary age of Late Miocene for this fau- 
na. Harry Dowsett, also of the U.S.G.S. (personal com- 
munication), examined the planktic foraminifera from 
an adjacent localit\ of the same formation on Cayo Agua 
and found an assemblage indicative of planktic zone 
N17-NT8 (Late Miocene-Early Pliocene). The consensus 
at this stage, therefore, is that the beds from which N. 
fortidentata was collected are Late Miocene to Early 
Pliocene in age. 




Figure 2. Nerita fortidentata new species, from Bocas del 
Toro, Panama. Holotype, UNSM 423644, height 19.7 mm. Ab- 
apertural view. 

Remarks: The new species clearly belongs to the sub- 
genus Theliostyla Morch, 1852 (type N. albicilla Lin- 
naeus, 1758), which is characterized by granulate sculp- 
ture on the parietal region, a barely protruding spire, 
and w ell-developed external spiral sculpture. Among the 
four Recent species of this subgenus in tropical America, 
N.fulgurans Gmelin, 1791, bears the closest resemblance 
to A', fortidentata. Measurements of 17 specimens of N. 
fulgurans in the Vermeij collection from the Atlantic 
coasts of Panama, Costa Rica, Venezuela, and Jamaica 
show that this species is less globose (G = 0.54 ± 0.020, 
range 0.50-0.57) and basally much more excavated 
(E = 1.27 ± 0.05, range 1.21-1.39) than is the new 
species. Nerita funiculata Menke, 1851, the Recent east- 
ern Pacific cognate of N. fulgurans, is also less globose 
(G = 0.51 ± 0.04, range 0.42-0.58, based on 15 speci- 
mens in the Vermeij collection from Costa Rica, Panama, 
and Ecuador) and more excavated (E = 1.28 ± 0.15, 
range 1.13-1.60) than is N. fortidentata. Both N. ful- 
gurans and N. funiculata have weaker and more nu- 
merous denticles on the outer lip, weaker and more finely 
granulated ridges on the parietal region, and spiral cords 
that are more numerous and more variable in size on the 
body whorl (18-35 in N. fulgurans, usually more than 
30 in N. funiculata). The spiral cords of N. fulgurans 
show a tendency to bifurcate on the body whorl, whereas 
no such tendencN is seen in iV. fortidentata. The three 
subspecies of N. ascensionensis Gmelin, 1791, from is- 
lands in the tropical south Atlantic (Vermeij, 1970) differ 
from iV. fortidentata by having a nearly smooth parietal 
region and by the very weak dentition on the outer lip. 
The West Indian A', tessellata Gmelin, 1791. differs from 
A', fortidentata b\' having low rounded spiral cords bro- 
ken irregularl) bv high and low areas correlating to the 
characteristic black and white checkered pattern found 
in this species, a finely granulated parietal region of 
relatively large extent, and a weakK denticulated outer 
lip (Russell, 1941). 

Jung (1965) recorded Serita fulgurans from the Mid- 
dle Miocene and Upper Pliocene of Venezuela, but he 
pointed out that his specimens differed from Recent shells 
by having stronger apertural dentition and a less concave 



Page 104 



THE NAUTILUS, Vol. 102, No. 3 



(that is, less excavated) parietal region. Later Jung (1969) 
found a similar shell in the Late Miocene Melajo Clay 
Member of the Springvale Formation of Trinidad. Like 
the Venezuelan material, the shell from Trinidad has 
only 16 ribs, but instead of having 2 centrali\ placed 
columcllar teeth, as in the Venezuelan material, the Trin- 
idad specimen has one upper tooth which curves into 
the aperture (this upper tooth is probably equivalent to 
the parietal fold found on the tvpe of \. fortidentata] 
and 2 somewhat lower denticles. Jung tentati\ely re- 
ferred both lots to N. exuvioides Trechmann, 1935, a 
species described on the basis of one incomplete speci- 
men from the Pliocene of Carriacou in the Grenadines. 
Jung (1971) redescribed this latter specimen as having 
only 12 ribs whose edges overhang the adjacent inter- 
spaces. Vokes (1983) clarified the status of N. exuvioides 
when she described a very strongly ribbed shell with 1 1 
ribs from the Gatun Formation of Panama. This speci- 
men closely resembles the one from Carriacou and is 
clearly referable to S. exuvioides. We believe that Jung's 
(1965, 1969) specimens from Venezuela and Trinidad 
belong neither to N. fulgurans nor to N. exuvioides, but 
instead to our new species, iV. fortidentata. This species 
appears therefore to be intermediate in sculpture be- 
tween N. exuvioides with only 1 1-12 ribs and the Recent 
.v. fulgurans, usually with more than 21 ribs. The re- 
lationship between N. fortidentata and the Late Oli- 
gocene or Early Miocene N. tampaensis Dall, 1892, from 
the Tampa Formation of Florida is unclear. Merita tam- 
paensis, whose granulated parietal region suggests place- 
ment in Theliostyla, is a small species with weakly de- 
veloped apertural dentition and highly \ariable external 
sculpture, some shells being nearly smooth whereas oth- 
ers have fine spiral cords of varying sizes. 

The shape of xV. fortidentata suggests that this species 
inhabited the upper zones of rocky shores. Merita ful- 
gurans, its most similar living relative, is usually found 
in areas of reduced salinity , such as the mouths of harbors 
(Russell, 1941) or protected embayments. Vermeij (1973) 
showed that shells with low globosity, high basal exca- 
vation, weak sculpture, and relatively broad apertures 
with weak dentition are found in middle to lower inter- 
tidal species of Merita; whereas species with a globose, 
little excavated, strongly sculptured shell and a small 
aperture bordered by strong teeth are found at higher 
shore levels. The latter shell form is especially charac- 
teristic of the subgenera Cyrnostyla von Martens, 1887, 
and Ritena Gray, 1858. Of the living and fo.ssil members 
of the subgenus Theliostyla, most of which live in the 
middle zones of the intertidal, M . fortidentata most closely 
approaches species of Ritena. Other fossils collected with 
N. fortidentata, including Oliva, Olivella, Conus, Na- 
tica, Polinices, Strombus, Phalium, Dentalinm. and 
Corlmla, suggest a variety of different environments, 
implying post-mortem transport and mixing of assem- 
blages. 

BIOGEOGRAPHY OF AMERICAN MERITA 

Among the fossil species of Merita that have been de- 
scribed from late Eocene and younger strata in tropical 



.\merica, at least two have close affinities with li\ing 
Indo-\^'est-Paci£ic species. Merita listrota Woodring, 
1973, from the late Eocene (?) Gatuncillo Formation of 
Panama has a finely ribbed shell w ith a peripheral keel, 
fine teeth on the outer lip, 7 teeth on the columellar lip, 
and a sparseK papillate parietal region. Woodring (1973) 
noted the striking similarity' between A', listrota and the 
recent mangrove-associated M. planospira Anton, 1839 
(the type and only known species of the subgenus Ily- 
nerita von Martens, 1887) from the tropical Indo-Pacific. 
He doubted that the tw^o species were closely related, in 
part because M. planospira has 5 rather than 7 columellar 
denticles. We consider the similarities to be so numerous 
that an inference of close relationship seems warranted. 
If A', listrota belongs to the subgenus Ilynerita, as we 
believe it does, that subgenus may be added to the grow- 
ing list of taxa whose distributions became restricted to 
the Indo- West-Pacific during the Tertiary (N'ermeij, 
1986). As Vokes (1983) points out, \. (Theliostyla) exu- 
vioides may represent a second lineage that has become 
restricted (as N. exuvia Linnaeus, 1758) to the Indo- 
West-Pacific. Merita exuvioides differs from the Western 
Pacific M. exuvia chiefly by having 1 1 instead of 14 strong 
overhanging spiral cords on the body whorl. 

The other fossil species of Merita that have been de- 
scribed from late Eocene and younger deposits in tropical 
America do not easih fit with any living members of the 
genus. These are M. hadra Woodring, 1973, from the 
late Eocene (?) Gatuncillo Formation of Panama and N. 
oligopleura Dall and Ochsner, 1928, from the Pleistocene 
of the Galapagos. Merita hadra has very fine spiral threads 
on the bod\' whorl and a smooth parietal region, whereas 
M. oligopleura has 3 broad spiral ribs on the body whorl. 

Although the record of Merita in tropical America is 
very meager, the history of the genus points to multiple 
instances of extinction and geographical restriction. It is 
too earl\ to assess the scope of these changes, but the 
record of Merita suggests that intertidal species have been 
no less affected by events leading to extinction and re- 
striction than have species from the better-sampled fossil 
environments of subtidal bottoms. 

ACKNOWLEDGEMENTS 

T.M.C. gratefully acknowledges a Smithsonian Tropical 
Research Institute short-term fellow ship, directed by Jer- 
emy Jackson, during w hicli large collections of Late Ter- 
tiary fossil mollusks were made. Collecting was made 
more fruitful and pleasant by the expertise and company 
of the other members of the field part\-, A. G. and J. M. 
Coates. We thank Laurel B\ bell, Thomas M. Cronin, and 
Harry Dowsett of the U.S.G.S. for their age determi- 
nations. 

LITERATURE CITED 

Anton, H. E. 1839. Verzeichniss der Conchylien welche sich 
in der Sammlung von Hermann Eduard .^nton befinded. 
Halle, xvi -I- 110 p 

Bold, W. A. van den. 1967. Ostracoda of the Gatun For- 
mation, Panama. Micropaleontology 13(3):306-318. 



G. J. Vermeij and T. M. Collins, 1988 



Page 105 



Dall, W. H. 1892. Contributions to the Tertiary fauna of 
Florida; with especial reference to to the Miocene silex 
beds of Tampa and the Pliocene beds of the Caloosahatchie 
River. Transactions of the Wagner Free Institute of Sci- 
ence 3(2):201-473, pis. 13-22. 

Dall, W. H. and W. H. Ochsner. 1928. Tertiary and Pleis- 
tocene Mollusca from the Galapagos Islands. Proceedings 
of the California Academy of Science 17{4):89-139, pis. 
2-7. 

Gray, J. E. 1858. Observations on the genus Nerita and its 
operculum. Proceedings of the Zoological Society of Lon- 
don 26:92-94. 

Gmelin, J. F. 1791. In: Linnaeus' Systema naturae, Tom. I, 
Pare VI, p. 3021-3909. 

Jung, P. 1965. Miocene Mollusca from the Paraguana Pen- 
insula, Venezuela. Bulletins of American Paleontology 
49(223):389-652. 

Jung, P. 1969. Miocene and Pliocene mollusks from Trinidad. 
Bulletins of American Paleontology 55(247):293-657. 

Jung, P. 1971. Fossil molluscs from Carriacou, West Indies. 
Bulletins of American Paleontology 61(269):147-262. 

Linnaeus, C. 1758. Systema naturae, 10th ed. Stockholm, 
824 p. 

Martens, E. von. 1887-89. Die Gattungen Nerita und Neri- 
topsis. Svstematisches Conchylien-Cabinet von Martini und 
Chemnitz 2(11):1-147. 

Menke, K. T. 1851. Conchylien von Mazatlan, mit kritischen 
Bemerkungen. Zeitschrift fiir Malakozoologie, yr. 8. 

Morch, O. A. L. 1852 Catalogus conchyliorum quae reliquit 
D. Alphonso d Aguirra & Gadea Comes de Yoldi, Copen- 
hagen, 170 p. 

Russel, H. D. 1941. The Recent molluscs of the family Ne- 
ritidae of the western .Atlantic. Bulletin of the Museum of 
Comparative Zoology at Harvard College 88(4):347-404. 



Trechmann, C. T. 1935. The geology and fossils of Carriacou, 
West Indies. The Geological Magazine 72(858):528-555, 
pis. 20-22. 

Vermeij, G. J. 1970. The Nerita ascensionensis species com- 
plex (Gastropoda: Prosobranchia) in the South Atlantic. 
The Veliger 13(2):135-138. 

Vermeij, G. J. 1973. Morphological patterns in high intertidal 
gastropods: adaptive strategies and their limitations. Ma- 
rine Biology 20:319-346. 

Vermeij, G. J. 1984. The systematic position of the neritid 
prosobranch gastropod Nerita polita and related species. 
Proceedings of the Biological Society of Washington 97(4): 
688-692. 

Vermeij, G. J. 1986. Survival during biotic crises: the prop- 
erties and evolutionary significance of refuges. In: D. M. 
Elliott (ed.). Dynamics of extinction. Wiley, New York, p. 
234-246. 

Vokes, E. H. 1983. Nerita exuvioides Trechmann (Mollusca: 
Gastropoda) from the Gatun Formation of Panama. Tu- 
lane Studies in Geology and Paleontology 17(4): 131-134. 

Woodring, W. P. 1973. Geology and paleontology of Canal 
Zone and adjoining parts of Panama: description of Ter- 
tiary Mollusks (additions to gastropods, scaphopods, pe- 
lecypods: Nuculidae to Malleidae). U.S. Geological Survey 
Professional Paper 306-E:453-539. 

Woodring, W, P. 1982. Geology and paleontology of Canal 
Zone and adjoining parts of Panama: description of Ter- 
tiary Mollusks (Pelecypods: Propeamussidae to Cuspida- 
riidae; additions to families covered in P.306-E; additions 
to gastropods; cephalopods). U.S. Geological Survey 
Professional Paper 306-F:541-759. 



THE NAUTILUS 102{3):106-1{)9, 1988 



Page 106 



A New Species of Favartia from the Eastern Pacific 
(Gastropoda: Muricidae) 



Anthony D'Altilio 
Barbara ^ . Myers 

Department of Marine Invertebrates 
San Diego Natural History Museum 
San Diego, CA 92101, USA 



ABSTRACT 



Favartia (Murexiella) shasktji is described from Isla del Coco, 
Costa Rica, and compared with related species from the eastern 
Pacific, This species is known onl\ from this isolated oceanic 
island 



INTRODUCTION 

Isla del Coco (also known as Cocos Island), one of the 
National Parks of Costa Rica, is a small, uninhabited 
island situated approximately 600 kilometers south south- 
west of Puntarenas, Costa Rica, at 5°33'N latitude and 
87°03'W longitude. Isla Cascara and Roca Sucia are two 
of the many islets surrounding Cocos Island. In May, 
1985, Donald R. Shasky of Redlands, California, and 
Kirstie L. Kaiser of La Canada, California, collected five 
specimens of the new species described herein, at depths 
of 50-80 feet (15.2-24.4 m), under dead coral off these 
two islets. In a previous paper (D'Attilio, M>ers & Shasky, 
et al., 1987), a new species of Phyllonotus Swainson, 
1833, was described from the same area. Montoya (1983, 
1984) published a bibliography of Cocos Island molluscan 
faunal studies. 



SYSTEMATICS 

Family Muricidae Rafinesque, 1815 

SubfamiK Muricopsinae Radvvin & D',\ttilio, 1971 

Genus Favartia Jousseaume, 1880 

Type species: Miirex breviculus Sowerby , 1 834, by orig- 
inal designation. 

Subgenus Murexiella Clench & Perez Farfante, 1945 



Type species: Mtirex hidalgoi Crosse, 1869, b>- original 
designation. 



Favartia (Murexiella i shaskyi new species 
(figures 1-10) 

Description: Shell (figures 1, 2) broadly fusiform, spire 
'/3 shell length; protoconch of holot\pe eroded, proto- 
conch of paratype 1 (figure 3) of 3'/2 smooth, lustrous, 
convex whorls, with axial buttresses arising from the te- 
leoconch; teleoconch with 5-6 subangulate whorls; suture 
weakly impressed; whorls gentK sloping from suture to 
shoulder; body whorl % shell length; aperture ovate; in- 
ner lip erect posteriorly; outer lip thin, erect, crenulate, 
reflecting the spiral cords; siphonal canal '3 shell length, 
broad proximalK', tapering distalK', with narrow \entral 
opening, terminally recurved, tube-like; siphonal fascicle 
of 3 fine distal portions of previous canals; body whorl 
with 6 broad varices, penultimate whorl with 8 varices; 
varices broader than intervarical regions; spiral sculpture 
of 6 cords on body whorl, all nearK equal in width, 
packed closely together; 2 additional cords between 
body whorl and siphonal canal, 2 major and 1 minor 
cord on canal; 2 cords per whorl on spire; shoulder of 
body whorl without spiral cords; all cords terminate as 
spines on varices (figures 4, 5), posterior spines recurved, 
anterior spines long, projecting ventrally; spiral cords 
(figures 6, 7) strongly scabrous; scales prominent, fine, 
white, erect, closely packed, disguising the contour of 
cords; cords and scales microscopicalK grooved, incre- 
mentally incised; adapertural sides of varices, especially 
last varix, completeK- scaled; scales arranged in 3 tiers 
(figure 6), with tube-like spines on tier farthest from 
aperture; radula (figure 8) t>picall\ muricopsine, with 
broad, U-shaped basal plate, and strongK projecting cusps. 
Paratypes 1 and 2 with only 4 postnuclear whorls; suture 
more strongly impressed, spire angulate. 



Figures 1, 2. Favartia (Murexiella) shaskyi new species. 1 . Holot> pe, L'SNM 860012, Isla Cascara, Cocos Island, Costa Rica, under 
dead coral slab, in 24.4 m, 23.0 mm long. 2. Paratype 1, SDNHM 91873, Roca Sucia, Cocos Island, Costa Rica, under dead coral 
slab, in 24.4 m, 14.0 mm long. 



A. D'Attilio and B. W. Myers, 1988 



Page 107 




Page 108 



THE NAUTILUS. Vol. 102, \o. 3 




Figures 3-7. Favartia (Murexiella) shaskyi new species, details of shell sculpture 3. Protoconch of parat\pe 1. 4, 5. Adapertural 
(4) and abapertural (5) views of last varix of holotype. 6, 7. Microscuipture along adapertural (6) and abapertural (7) surfaces of 
a spiral cord of the holotype. Figure 8. Rachidian and right lateral teeth from the holotype. 



Color: Base color buff to tan, with diffuse darker band 
encircling body whorl; scales white, aperture white, si- 
phonal canal white. 

Type locality: Isla Cascara, Cocos Island, Costa Rica, 
under dead coral slab, in 24.4 m. 

Holotype: USNM 860012 (figure 1 ), 23.0 mm long, 14.2 
mm wide. 

Paratypes: Paratype 1: SDNHM 91873 (figure 2), 14.0 
mm long. Paratype 2: Collection of D. R. Shasky, 11.8 
mm long, both from Roca Sucia, Cocos Island, Costa 
Rica, under dead coral, in 24.4 m. 



Additional material examined: Tw o specimens, collec- 
tion of K. Kaiser, 19.2 mm long, and 14.7 mm long, Isla 
Cascara, Cocos Island, Costa Rica, under rocks, in 15.2- 
24.4 m. 

Etymology: We are pleased to name this species for 
Donald R. Shasky, M.D., who collected the three type 
specimens, and who generousK donated the holotype 
and parat\pe 1. 

DISCUSSION 

This new species has a distinct morphology that easily 
distinguishes it from all other eastern Pacific species of 



A. D'Attilio and B. W. Myers, 1988 



Page 109 



the subgenus Murexiella. Favartia (Murexiella) lappa 
(Broderip, 1833), a closely related species, differs in hav- 
ing a higher spire, biconic form, and short, stubby, non- 
recurved spines. Favartia (Murexiella) vittata (Broderip, 
1833) also has a more or less biconic shape, but the body 
whorl is more globose, and the spines, although recurved, 
are short and stubby. Favartia (Murexiella) keenae (E. 
H. Yokes, 1970) has somewhat similar sculpture, a larger 
shell with a globose body whorl strongly constricted at 
the base, and a strongly impressed suture. We have care- 
fully studied and compared the 12 additional nominal 
species of Murexiella, none of which are closely related 
to this new species. 

ACKNOWLEDGEMENTS 

We are grateful to Dr. Donald R. Shasky for donating 
the holotype and paratype and Kirstie Kaiser for the loan 
of her two specimens. We thank David K. Mulliner for 
the photography. M. G. Harasewych, EmiK H. Yokes, 
William K. Emerson and Carole M. Hertz reviewed the 
manuscript and made valuable suggestions. 

LITERATURE CITED 

Broderip, W. 1833. Characters of new species of Mollusca 
and Conchifera collected by Mr. Cuming. Proceedings of 
the Zoological Society of London 2:173-179. 



Clench, W. J. and I. Perez Farf ante. 1945. The genus Mt/rei 
in the western Atlantic. Johnsonia l(17):l-58. 

Crosse, H. 1869. Diagnoses Molliiscorum novorum. Journal 
de Conchyiiologie 17:408-410. 

D'Attilio, A., B W. Myers, and D. R. Shasky. 1987. A new 
species of Phyllonotus (Muricidac: Muricinae) from Isla 
del Coco, Costa Rica. The Nautilus 101(4):162-165, 

Jousseaume, F. 1880. Division methodique de la famille des 
Purpurides. Le Naturaliste 42:335-336. 

Montoya, M. 1983. Los Moluscos marines del la Isla del Coco, 
Costa Rica. I. Lista anotada de especies. Brensia 21:325- 
353. 

Montoya, M. 1984. Marine mollusks of Cocos Island, Costa 
Rica. I. Bibliographic compilation of species. Western So- 
ciety of Malacologists Annual Report (1983) 16:33-44, 

Radwin, G. E. and A. D'Attilio. 1971. Muricacean supra- 
specific taxonomy based on shell and radula. Echo 4:55- 
67. 

Rafinesque, C. S. 1815. Analyse de la nature ou tableau du 
univers et des corps organises. Barravecchia, Palermo, p. 
136-149. 

Sowerby, G. B., II. 1834. Conchological illustrations. Murex. 
Sowerby, London, pis. 58-67. 

Swainson, W. 1833. The zoological illustrations, ser. 2, vol. 3, 
pi. 67. 

Yokes, E. H. 1970. The west American species of Murexiella 
(Gastropoda: Muricidae). Veliger 12(3):325-329, pi. 50. 



THE NAUTILUS 102(3):110-114, 1988 



Page 110 



Conus baccatus G. B. Sowerby 111, 1877 
A Panamic Fauna! Constituent 



^ illiam K. Emerson 
Waller E. Sage III 

Department of Invertebrates 
American Museum of Natural History 
New York, NY 10024, USA 



ABSTRACT 

Specimens of a small species of Conus recent!) received from 
Golfo de Chiriqui, Republica de Panama, are confirmed to be 
referable to Conus baccatus Sowerby , 1877, a species originally 
described from an unknown locality. The provenance and iden- 
tity of this taxon had remained uncertain for more than 100 
years until Rockel (198.5a) compared two specimens from Pa- 
cific Panama in his collection with the holotype of Conus bac- 
catus and determined the specimens to be conspecific. We 
agree with Rockel's conclusion that this species has been con- 
fused in the past w ith the pustulate form of the west American 
Conus perplexus Sowerby, 1857, which is a somewhat similar, 
but distinct species. For Conus perplexus, a lectotype is selected 
and the type locality is restricted to the Golfo de Panama. For 
Conus baccatus. the Golfo de Chiriqui is designated the type 
locality. 



INTRODUCTION 

In 1877, G. B. So\verb%- III briefly described and illus- 
trated in color Conus baccatus from an unknown locality. 
Apparently based on a single specimen in the collection 
of Dr. Prevost of Alen^on, France, the figured specimen 
later was owned by J. C. Melvili (Sowerby, 1887:251) 
and subsec}uently by J. R. Le B. Tomlin (1937:217). The 
vast MeK ill-Tomlin Collection eventual!) w as deposited, 
along with this type, in the National Museum of Wales, 
Cardiff. Coomans et al. (1982:4) examined the holotype 
and provisionally accepted Conus baccatus as a valid 
species. They also rejected the Cjuestionable placement 
by Walls (1979:726) of this ta.xon in the synonymy of C. 
mindanus Hwass in Bruguiere, 1792, from the western 



Atlantic. \'ink (1984:356) subsequently noted certain 
similarities of the holotype of C. baccatus with C. selenae 
Van Mol, Tursch, and Kempf, 1967, a Brazilian taxon 
which we consider referable to the C. jaspidcus (Jmelin, 
1791, species complex. 

Through the generosit)- of Carol Skoglund and Robert 
Koch, we have examined a series of well-preserved spec- 
imens from Golfo de Chiriqui. Panama that are referable 
to Coijus baccatus. Our findings confirm Rockel's (1985a: 
29) conclusion that this taxon is a valid Pacific Panamic 
species, which for many years has been masquerading 
in collections as a pustulose form of Conns perplexus 
Sowerb), 1857. 

A specimen apparently referable to C. baccatus was 
illustrated by Kiener (1846:56, 57, pi. 83, fig. 2) as "Conus 
albomaculatus. Reeve ", in reference to Sow.[erby] (1833, 
Conus fig. 2, which was named C. bicolor h\ Sowerby 
in 1833 and renamed C. albomaculatus h\ Sowerb) in 
1841). Kiener "s figured specimen, measuring 22 mm in 
height, was attributed to the Dupont Collection from an 
unknown locality. This specimen cannot be located in 
the Museum National d Histoire Naturelle, Paris, and 
the fate of the Dupont collection is not known (P. Bou- 
chet, in lift.. March 4, 1987). Reeve (1849:3) considered 
Kiener 's illustrations to represent the western Atlantic 
species C. mindanus Hwass in Bruguiere, 1792. He also 
stated that "C. albimaculalus [sic] [Sowerb) ] has more 
resemblance with very young [pustulose] specimens of 
C. lithoglyphus [.sir] [Hwass in Bruguiere, 1792] ', a con- 
clusion accepted by Coomans et al. (1979:97) and Rocke! 
(1985b, nr. 22). Sowerb)- "s figured specimen of C. albo- 
maculatus has not been located. It is not in the Britisli 
Museum (Natural History) (K, M. Way, in litt.. May 11, 



Figures 1-7. Conus baccatus Sowerby, 1877. 1, 2. Holotype, NMW no. 1955.158.29; x 2 (courtesy of R. M Filmer) 3. 4.. Off 
Isla Coiba, Panama, AMNH no, 221871a; x 2. 5. Off Isla Parida, Panama, AMNH no. 173698a; x 5 6, 7. Off Isla Coiba, Panama. 
AMNH no. 221871; x 2, Figures 8-16. Conus perplexus Sowerby, 1857, 8. 9. Lectotype, BM(NH) no. 1978118; x 2. 10. Pla\a 
de los Angeles, Bahia Tenacalita, Jalisco, Mexico, dredged, mud bottom, Aug., 1975, AMNH no. 221873a, ex C and P. Skoglund; 
X 5. 11, 12. Paralectotype, BM(NH) no. 1978118a; x 2 13, 14. Paralectotype, BM(NH) no. 1978118b; x 2. 15. 16. "West 
Panama" (note the distorted spire), AMNH no. 212547; x 2 Figures 17. 18. Conus puncticulatus form papillosus Kiener, 1849, 
Cartagena, Colombia, Iseach, 1977, ex J. M. Bijur Coll., AMNH no. 225979; x 2. Figure 19. Conus yemanjae \aii Mol et al., 
1967, paratype, pi. 8, fig. lb, Fortaleza (Ceara), Brazil, ex pisce; x 2. 



W. K. Emerson and W. E. Sage III, 1988 



Page 111 











Page 112 



THE NAUTILUS, Vol. 102, No. 3 



1987) nor is it in the National Museum of Wales (A. 
Trevv, in litt.. Ma\, 1987). Therefore, the taxonomic 
status of C. alboviaculatus Sowerby, 1841, cannot be 
determined from the available data. 

Although C. baccatus was not formally named by Sow- 
erb\ ill until 1877, specimens were apparentl\ available 
in European collections by the nii(l-I9th century, as sug- 
gested by Kiener's figured specimen from the Dupont 
collection. There is also a specimen of C. baccatus labeled 
""Panama" in the F. A. Constable Collection (AMNH 
47740), which dates from the latter part of the 19th 
century and was received by the American Museum of 
Natural History from the estate of Louise Constable in 
1929. 

Abbreviations for institutions used in text; AMNH = 
.\merican Museum of Natural History; BM(NH) = Brit- 
ish Museum (Natural History); Los Angeles County Mu- 
seum of Natural History = LACMNH; NMW = National 
Museum of Wales. 

SYSTEMATIC ACCOUNT 

Conus baccatus Sowerby, 1877 
(figures 1-7) 

? "Conus albomaculatus. Reeve"', Kiener, 1846:56, 57, pi. 83, 
fig. 2 (apertura! and dorsal views; as "Conus albimaculatus 
[sic] Sow.[erby] ), from unknown locaiit) , Dupont collec- 
tion. Not C. bicolor Sowerby, 1833: pt 24, fig. 2, renamed 
without explanation C. albomaculatus Sowerby, 1841. 

Conus baccatus Sowerby, 1877:753, 754, pi. 75, fig. 5, from 
unknown locality; Tryon, 1883: ser. 1, vol. 6, p. 22, pi. 6, 
fig. 92 [copy of original fig.], "Habitat unknown "; Sowerby, 
1887: vol. 5, p. 251, Conus pi. 29, fig. 660 [copy of original 
fig.], "Habitat unknown. Coll. Meiviii"; Tomlin, 1937:217, 
"Hab. ?, Type in coll. Tomlin"; Coomans et a/., 1982:4, 
fig. 197, apertural and dorsal views of the holotype; Vink, 
1984:354-358, pi. 20, fig, 1, dorsal view of holotype; Rock- 
el, 1985a:29, dorsal view of holotype and dorsal and aper- 
tural views of two recently collected specimens from "Pe- 
rida Island ", Pacific Panama (figured in color). 

Original description (in part): "Shell short, rather swol- 
len, very minutely decussated, with regular rows of con- 
spicuous granules, whitish, with large orange blotches 
arranged in three bands; spire acute, short, whorls con- 
cave, nearly smooth, last whorl biangulated. . . . Apart 
from its somewhat stunted form, the delicacy of its mark- 
ings and rows of gem-like granules, it is remarkable for 
the double angle at the top of the body whorl. Long. 23, 
lat. maj. 15 mill " [Holotype actually measures 22.2 by 
14.2 mm.) 

Type depository: Holotype in the National Museum of 
Wales, no. 1955.158.29, fide Trew (1982:3); here illus- 
trated (figures 1, 2). 

Type locality: Off Isla Parida, Golfo de Chiriqui, Re- 
piiblica de Panama, here designated. 

Range: Known only from Pacific Panama. 

Material examined: ,\MNH Collection: From Golfo de 
Chiritpii, Pacific Panama: 6 specimens, no. 173698 and 
8 specimens, no. 211778, off Isla Parida (8''05'N, 82°20'W), 



in 5.5-9 m, sand bottom. May, 1972, ex R. E. Hubert; 2 
specimens, no. 210849, off Isla Parida, sand, in 5.5 m, 
March, 1974, ex R. E. Hubert; 3 specimens, no. 225981, 
off Isla de Canal de Afuera, in 55 m, ex J. Ernest; 2 
specimens, no. 221871, off Isla Coiba, in 24-40 m, March 
28, 1986, ex C. and P. Skoglund; 2 specimens, no. 221872, 
off S.E. Isla Rancheria, in 2-7.5 m, March 27, 1986, ex 
C. and P. Skoglund; 2 specimens, no. 223269, off Isla 
Rancheria (7°38'N, 81°44'W), in 3-9 m, coral sand and 
debris bottom, February 24, 1987, ex R. Koch. From 
"Panama"; 1 specimen, no. 47740, ex F. A. Constable 
Coll.; 3 specimens, no. 201566, ex T. H. and V. B. Mun- 
yan Coll. 

Remarks: Specimens of Conus baccatus superficially 
resemble small examples of the pustulose form of Conus 
perplexus. There are several discrete differences in shell 
morphology which serve to distinguish the two taxa. 1, 
Nuclear whorls: In C. baccatus, I'i whorls, nucleus short, 
terminating in a broad, well-defined mammillated pro- 
cess. In C. perplexus, 2'/2 whorls, nucleus elongate, nar- 
row, terminating in a needle-like projection (cf. figure 5 
with figure 10). 2, Shoulder margin of bod\ whorl: In C. 
baccatus, oblique, twofold-angled (i.e., biangulated). In 
C. perplexus. monoangulated (cf. figures 6, 7 with figures 
8, 9). 3, Spiral granulations; In C baccatus, rows of pus- 
tules more widely spaced, some rows weakK developed. 
In C. perplexus, rows consistently spaced, nearly uni- 
formly developed (cf. figures 6, 7 with figures 15, 16). 
4, Color pattern: In C. baccatus, spiral bands of broken 
blotches, orange in faded specimens, reddish-broun in 
fresh specimens. In C. perplexus, narrow spiral lines with 
dark brown dots (cf. figures 1, 2 and 3, 4 with figures 8, 
9). Additionally, the periostracum in C. baccatus is yel- 
lowish tan, whereas in C. perplexus it is a brownish tan. 
The largest specimen of C. baccatus examined (AMNH 
221871a; figures 6, 7) is 26.9 mm in height, compared 
to a maximum of 37.8 mm for C. perplexus (AMNH 
206684) from Isla Cebaco, Golfo de Montijo, Panama. 
Hanna (1963:39) reports a specimen of the latter taxon 
from Isla San Jose, Golfo de Panama, with a height of 
41.5 mm. Both of these taxa have shells with a distinct 
posterior notch on the lip (cf. figure 5 with figure 10), 
whereas C. xirnenes Gray, 1839, C. nmhogani Reeve, 
1843, and C. tornatus Sowerb\, 1833, which also have 
been confused with C. perplexus, have a narrow anal 
notch (see Wolfson, 1962: figs. 6a-d; Tucker 1985: fig. 
c; and Chancy, 1987: figs. 2, 3). All of these nominal 
species have populations sympatric with those of C. bac- 
catus, which is known only from Pacific Panamic waters. 
Conus perplexus, with which C. baccatus is most likely 
to be confused, ranges from the northern part of the Gulf 
of California, south to Mancora, Peru, 4°05'S, 81°03.5'W 
(LACMNH no. 70-101). Specimens of C. perplexus in 
which the interrupted spiral lines of brown dashes or dots 
are raised into prominent pustules are more commonly 
found in the southern populations, from Panama south- 
ward to Peru (figures 15, 16). .\ similar pustulose form 
also occurs in populations of the western .\tlantic C 
puncticulatus Hwass in Bruguiere, 1792, which is a twin 



W. K. Emerson and W. E. Sage III, 1988 



Page 113 



species of C. perplexus. This form was named C. pa- 
pillosus Kiener, 1849 (figures 17, 18). 

On the basis of shell morphology, C. baccatus appears 
to be most closely related to members of the western 
Atlantic C. jaspideus species complex, especially the pus- 
tulose, short-spired Brazilian forms which were given the 
names C. yemanjae (figure 19) and C. selenae by Van 
Mol et al. (1967: pi. 8); also figured by Vink (1984: pi. 
20). 

Conus perplexus Sowerby (1857:20, 1858: pi. 200, fig. 
324) was briefly described in Latin, followed by the com- 
ment; "This shell perplexes me, because there is a variety 
of C. puncticulalus [Hwass in Bruguiere, 1792, a Carib- 
bean twin species] which it nearly resembles. This, how- 
ever, is quite smooth, and more angular' . A single spec- 
imen was figured by Sowerby in a dorsal view, with the 
statement: "Gulf of California, West Columbia [sic], 
Cum.[ing]". Tomlin (1937:290) recorded the presence of 
three "types" in the British Museum (Naural History), 
which were kindh' loaned to us by Ms. K. M. Way 
[BM(NH) 19781 18^ here illustrated, figures 8. 9, 11-14]. 
The original labels confirm these specimens as syntypes, 
although none of them can be referred with certainty to 
Sowerby s figured specimen. The drawing may represent 
a composite. The three syntypes are somewhat faded, 
but are otherwise well-preserved. We have selected as 
the lectotype (figures 8, 9) the syntype that most closely 
resembles the original figure. The lectot\pe is slightly 
larger in height (26.8 vs. 26.1 mm) than the specimen 
depicted in the original illustration. 

The original labels accompanying the syntype lot of 
Conus perplexus list the Gulf of California and West 
Columbia [.sic], which are the same localities cited by 
Sowerby (1858, caption to Conus, pi. 14). Hugh Cuming 
is not believed to have collected in Mexican waters on 
his voyage to the west coast of South America, 1828-30. 
He is known, however, to have made extensive collections 
in the "Gulf of Panama, the Pearl Islands, and the Gulf 
of Chiriqui", as well as visits to Costa Rica, Nicaragua, 
and Honduras (Dance, 1986:114). We here restrict the 
type locality of C. perplexus to the Golfo de Panama, as 
the Isthmus of Panama was part of Colombia at that 
time. 

In summary, Conus baccatus G. B. Sowerby III, 1877, 
is recognized as a Panamic faunal constituent and as a 
twin species of C. jaspideus Gmelin, 1791, from the 
western Atlantic, especially the pustulose form named 
C. yemanjae Van Mol et al., 1967. Conus baccatus is 
compared with pustulose specimens of the sympatric C. 
perplexus G. B. Sowerby II, 1857, with which it has been 
confused, as well as with pustulose specimens of C. punc- 
ticulalus Hwass in Bruguiere, 1792, a Caribbean twin 
species of C. perplexus. A lectotype is selected and the 
t\pe locality is restricted for C. perplexus. A type locality 
is designated for C. baccatus. 

ACKNOWLEDGEMENTS 

We are indebted to Carol Skoglund and Bob Koch, Phoe- 
nix, Arizona, for sending us comparative material, for 



their observations on the differences in the nuclear whorls 
of Conus perplexus and the specimens which were found 
to be referable to C. baccatus. Their interest and coop- 
eration, past and present, are greatly appreciated. We 
acknowledge with thanks the assistance of Philippe 
Bouchet, Museum National d'Histoire Naturelle, Paris; 
Helen DuShane. Whittier, California; James Ernest, Bal- 
boa, Repiiblica de Panama; R. M. Filmer, Chobham, 
England; Robert A. Foster, Santa Barbara, California; 
Barbara Good, San Diego, California; James H. McLean, 
Los Angeles County Museum of Natural History; P. Gra- 
ham Oliver and Alison Trew, National Museum of Wales, 
Cardiff; Donald R. Shasky, Redlands, California; Kay C. 
Vaught, Scottsdale, Arizona; and K. M. Way, British Mu- 
seum (Natural History), London, for the loan of speci- 
mens and/or photographs, or for providing information. 
We also are indebted to Alan Kohn, University of Wash- 
ington for providing data on the complicated nomen- 
clatural history of the taxa Conus bicolor and C. albo- 
maculatus. Stephanie Crooms, AMNH, word-processed 
the manuscript. 

We owe a special debt of gratitude to Henry W. Cha- 
ney, Santa Barbara Museum of Natural History; James 
H. McLean; and James Nybakken, Moss Landing Marine 
Laboratories, for critically reading drafts of the manu- 
script. 

LITERATURE CITED 

Born, I. von. 1778. Index rerum naturalium Musei Cesarei 
Vindobonensis. Pars prima, Testacea. Vienna, xlii + 458 

p., 1 pl. 

Bruguiere, J,-G. 1792. Cone. In: Encyciopedie methodique. 
Historie Naturelle des Vers 1:586-757, illus. 

Chaney, H. W. 1987. A comparative study of two similar 
Panamic cones: Conus ximenes and Conus mahogani. The 
Veliger 29(4):428-436, 6 figs. 

Coomans, H. E., R. G. Moolenbeek, and E. Wils. 1979. Al- 
phabetical revision of the (sub)species in Recent Conidae 
2. adansoni io atbuquerquei. Basteria 4;3(5-6):81-105, figs. 
26-50. 

Coomans, H. E., R. G. Moolenbeek, and E. Wils. 1982. Al- 
phabetical revision of the (sub)species in Recent Conidae. 
5. baccatus to byssinus, including Conus brettinghami 
nomen novum. Basteria 46(l-4):3-67, figs. 172-292. 

Dance, S. P. 1986. A history of shell collecting. E. J. Brill, 
Leiden, xv -I- 265 pp., illus. 

Gmelin, J. F. 1791. Systema naturae per regna tria naturae, 
13th ed. 1(6), cl. 6, Vermes. Leipzig, p. .3021-3910. 

Gray, J. E. 18.39. Molluscous animals and their shells. In: F. 
W. Beechey. The zoology of Capt Beechey's Voyage . . . 
to the Pacific and Behring's Straits in His Majesty's ship 
Blossom. . . . London, p. i-xii -I- 103-155, pis. 33-44. 

Hanna, G D. 1963. West American mollusks of the genus 
Conus — II. Occasional Papers of the California Academy 
of Sciences 35:103 p., 4 text figs., 11 pis. 

Kiener, L.-C. 1846[-48], 1849[-.50]. Species general et ico- 
nographie des coquilles vivantes. Vol. 2, Conus. Paris, p. 
1-379, 111 pis. [Conus alhomaculatus, 1846, pp. 56, 57, 
pl. 83, fig. 2; C. papillosus. 1849, p. 271, pl; 72, fig. 4] 

Mol, J.-J. Van, B. Tursch, and M. Kempf. 1967. Resultats 
scientifiques des campagnes de la 'Calypso' 32. Campagne 



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de la Calypso au large des cotes atlantiques de I'Amerique 
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Reeve, L. 1843. Conchologia iconica, \ ol 1 London Conus. 
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Reeve, L. [184S-]1849. Conchologia iconica, \'ol. .5. Conus, 
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Rtickel, D. 1985a. Conus Imccatus Sowerby III, 1877, a for- 
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Rockel, D. 198.5b. Die Familie Conidae, nr. 22 [Contis lito- 
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Shaw, H. O. N. 1909. On the dates of issue of Sowerby s 
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Sowerby, G. B., II. 1857-58. Thesaurus conchyliorum. Vol. 
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Sowerby, G. B., III. 1877. Descriptions of six new species of 
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Dr Prevost. Proceedings of the Zoological Society of Lon- 
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Sowerby, G.B., III. 1887. /n.G. B. Sowerby II (ed.). Thesaurus 
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genus Conus, Vol. 5, pt. 44 London, p 249-279, pis. 507- 
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Tomlin, J R le B. 1937. Catalogue of Recent and fossil cones. 
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Trew, A. 1982. The .Melvill-Tomlin Collection, part 10, Co- 
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Tryon, G. W., Jr. 1883[-84]. Manual of concholog>, Ser. 1, 
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Tucker, J. K. 1985. Conus ximenes Gra\, 1839, Conus ma- 
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THE NAUTILUS 102(3):115-122, 1988 



Page 115 



References to Molluscan Taxa Introduced by Linnaeus in the 
Systema Naturae (1758, 1767) 



Kenneth J. Boss 

Miiseuiii of Comparative Zoology 
Harvard liniversity 
Cambridge, MA 02138, USA 



ABSTRACT 

Bibliographic documentation is provided for the more than 90 
published resources utilized by Linnaeus as citations for mol- 
lusks in the 10th and 12th editions of his Systema Naturae. 



INTRODUCTION 

Linnaeus's 10th edition of the Systema Naturae of 1758 
is the cornerstone of our system of binominal nomencla- 
ture in zoology. Many species of mollusks were described 
therein, and these have been the subject of individual 
investigations such as revisions of particular taxa as well 
as studies of the specimens themselves (Hanley, 1855; 
Dodge, 1952-59; Dance, 1967). 

Linnaeus also attempted to apply the binominal meth- 
od to the references that accompanied his treatment of 
the species. If he cited a reference that previously men- 
tioned or figured the species under discussion, he usually 
gave an abbreviated citation, which frequently embod- 
ied a shortened name of the author and that of the in- 
dividual work. Many of these are readily apparent to 
someone with a knowledge of the pre-Linnaean litera- 
ture; thus, "Gault. test." may be easily recognized as 
being a reference to Gaultieri's Index Testarum Con- 
chyliorum of 1742; others are more enigmatic. 

Some years ago I thought that it might be interesting 
to decipher these puzzling references in Linnaeus's Sijs- 
tema, both the 10th (1758) and 12th (1767) editions (the 
nth is a reprint of the 10th), much like Wheeler (1979) 
has done for fishes. However, in correspondence with 
the late Professor A. Myra Keen, I found that both she 
and Dr. S. S. Berry also had an interest in this problem. 
Professor Keen sent me a short draft of some 30 titles 
that she had connected with Linnaeus's abbreviations, 
and I supplied her with several that she had been unable 
to find. Eventually, she published a narrative account of 
certain authors whom Linnaeus had cited (see Keen, 
1983a, b). In these papers, she noted that Linnaeus used 
about 40 different references in his sections on mollusks. 
However, when the different citations are tallied for both 
soft bodied and shelled mollusks [including therefore 
Linnaeus's categories with their genera, Vermes: Intes- 



tina: Teredo, Vermes: MoUusca: Clio, Limax, Doris, Te- 
thijs, Scyllaea, and Sepia; and Vermes: Testacea: (all 
genera with the exception of Lepas and Serpula though 
references to the two molluscan Serpula are included)], 
the total exceeds 90. Dance (1967) also provided a short 
list of volumes personally owned by Linnaeus which were 
utilized in various editions of the Systema. 

Unknown to any of us were the studies of Professor 
John L. Heller, Emeritus, of the Department of Classics 
of the University of Illinois on various botanical works 
of Linnaeus (see Heller, 1983, for a collection of these 
articles). He provided extensive bibliographic references 
to similarly abbreviated citations by Linnaeus in some 
of his botanical works such as the Species Plantarum 
and the Hortus Cliffortianus. He also explicated certain 
particularly equivocal abbreviations used by Linnaeus in 
referring to various insects (Heller, 1961) and planned 
to finish a complete bibliographical guide to zoological 
works cited by Linnaeus under the proposed title Bib- 
liotheca Zoologica Limmeana (Heller, 1968, 1979, 1983); 
this work is currently in press (Heller, personal com- 
munication) but will contain more than a malacologist 
need know. 

Although I am separately preparing a more extensive 
treatment of the pre-Linnaean malacological citations 
including comments on subsequent editions, translations 
into other languages as well as annotations of the holdings 
of the Harvard College Library system (Boss, in prepa- 
ration), I present below the bibliographic abbreviations 
utilized by Linnaeus in ihe Systema Naturae (1758, 1767) 
and the full citation for each reference. Of all the ref- 
erences, only one remains undeciphered: Linnaeus (1758: 
708, 1767:1161) referred to "Tessin. epist. 1 n. 28 Cym- 
bium" under Argonauta argo. This is presumed to have 
been a letter of Count Carl Gustaf Tessin, on whose 
collection Linnaeus worked. Despite a thorough check- 
ing of apposite sources (i.e., items in the Literature Cited 
as well as subsequent editions of the Systema and several 
classical cephalopod authorities such as Chemnitz, Ke- 
ferstein, Conrad, Naef, Robson, Tryon, and Ferussac- 
Orbigny), this reference could not be located. 

There are several bibliographic sources for pre-Lin- 
naean works, and sometimes there are discrepancies in 



Page 116 



THE NAUTILUS, Vol. 102, No. 3 



citations of the sources. These may arise due to different 
bibliographic methods, through error, or due to the vari- 
ability in the copies of pre-Linnaean works, which were 
not always issued in uniform copies I have relied prin- 
cipally on the Catalogue of the Books . . . British Mu- 
seum .... Soulsby (1933), Engelmann (1846), Heimann 
(1957), Nissen (1969), and the resources of \arious Har- 
vard libraries. The birth and death dates of authors have 
been included, mainly as listed in the British Museum 
Catalogue; these do not always concur with such dates 
given by other authors, principally Dance (1986) or Nis- 
sen (1969). I have utilized my own method of punctua- 
tion and have spelled out such words as "engraved" and 
"portraits" when describing the texts; I have followed 
standard procedure for the usage of brackets, etc., as 
noted in the Catalogue of the Books . . . British Museum. 
Further, the names of publishers or printers were added 
when 1 knew them. With few exceptions, which are listed 
as "not seen", examination oi the original publication 
was possible. 



ABBREVIATED CITATIONS UTILIZED 
BY LINNAEUS FOR MOLLUSKS 

Act. angl. 301, p. 2051 (see Breyn, 1705) 

Act. helv. 4. p. 212, t.9, f.21, 22 (see Hofer, 1760) 

Act. helv. 5. p. 283, n.4, t.3, f.25, 26 or n.5 t.3 f. 27, 28 

(see Schlotterbeccius, 1762) 
.\ct. paris. 1710, p. 463 (see Reaumur, 1710) 
Act. paris. 1711, t.3, f.4,5 (see Reaumur, 1711a) 
Act. paris. 1711, p. 199, t.6, f.5.7. or 9 (see Reaumur, 

1711b) 
Act. paris. 1712, p. 163 (see Reaumur, 1712) 
Act. Petropol. 7, p. 321, t.U, 12 (see Koelreuter, 1761) 
Act. Upsal. 2. p. 560, t.l52, f.4. idem or f.a. (see Bromell, 

1729) 
Adans. sen. or seneg. (see Adanson, 1757) 
Aldr. exs. or exsang. or exangu. (see Aldrovandi, 1606) 
Aldr. ins. (see Aldrovandi, 1602) 
Amoen. acad. (see Amoenitates Academiae) 
Amoen. acad. 1, p.325 (see Balk, 1746) 
Amoen. acad. 3: 256 (see Odhelius, 1754) 
Argenv. conch, (see Dezallier d'Argenville, 1742) 
Barr. ic. or rar. or Barrel, icon, or rar. (see Barrelierus, 

1714) 
Baster subs, (see Raster, 1759-65) 
Bell. aqu. or aquat. or Rellon. aqu. (see Belon, 1553) 
Blank, ins. (see Blankaart, 1688) 
Bocc. observ. (see Boccone, 1674) 
Bohads. mar. (see Bohadsch, 1761) 
Ronan. kirch. (see Buonanni, 1709) 
Bonan. recr. or recreat. or Bonann. recr. (see Buonanni, 

1681, 1684) 
Brad, natur. or Bradl. nat. or Bradl. natur. (see Bradley, 

1721) 
Breyn. polyth. (see Breynius, 1732) 
Brown, jam. (see Browne, 1756) 
Bruckm. cent. 2 epist. (see Brueckmann, 1743) 
Calceol. mus. (see Cerutus and Chiocco, 1622) 



Chin. Lagerstr. (see Odhelius, 1754) 

Column, aqu. or aquat. (see Colonna, 1606) 

Column, phytob. (see Colonna, 1592) 

(Column, purp. (see Colonna, 1616) 

Crew. mus. (see Grew, 1681) 

E.N.C. (see Francus de Frankenau, 1727) 

Edw. av. (see Edwards, 1758-64) 

Ellis cor. or corall. (see Ellis, 1755) 

En. svec. (see Linnaeus, 1746) 

Faun. svec. or En. svec. (see Linnaeus, 1746) 

Frisch ins. (see Frisch, 1730) 

Gesn. aqu. or aquat. (see Gesner, 1551-87) 

Ginam. adr. (see Ginnani, 1755-57) 

Ginan. or Ginan. adr. (see Ginanni, 1755-57) 

Gmelin act. petrop. vol. 3, p. 246 (see Gmelin, 1729) 

Gnalt. test, (see Gualtieri, 1742) 

Grev. mus. or Grew. mus. (see Grew, 1681) 

Gron. lap. (see Gronovius, 1740) 

Gualt. test, (see Gualtieri, 1742) 

Hasselq. or Hasselqv. itin. (see Hasselquist, 1757) 

Hasselqv. act. ups. 1750. p. 33. (see Hasselquist, 1750) 

Hevde .Ant. Anatome mvtulorum 1683 oct (see Heide, 
1683) 

Imperat. nat. (see Imperato, 1599) 

It. gotl. (see Linnaeus, 1747) 

It. oel. or oeland (see Linnaeus, 1745) 

It. wgot. or wgoth. or wogth. (see Linnaeus, 1747) 

Johnst. aquat. or exsangu. (see Jonstonus, 1650-53) 

Jonst. aquat. (see Jonstonus, 1650-53) 

Kaehl. or Kaehler act. Stockhb. 1754. p. 144, t.3, f.A-E 
or A-F (see Kahler, 1754) 

Kirch, mus. (see Buonanni, 1709) 

Klein or Klen. ostr. (see Klein, 1753) 

Kratz. or Kratzenst. Regenf. (see Regenfuss, 1758) 

Labat. itin. (see Labat, 1722) 

Lederm. micr. (see Ledermueller, 1760-61) 

Lewenh. arcan. (see Leeuwenhoek, 1695) 

List. angl. (see Lister, 1678) 

List. angl. app. [appendices] (see Lister, 1685, under Lis- 
ter, 1678) 

List, conch, or hist, (see Lister, 1685-92[-97]) 

List. exer. 2 or exerc. 2 or exercit. 2 (see Lister, 1695) 

List, exercit. anat. 1 (see Lister, 1694) 

List, exercit. anat. 2 (see Lister, 1695) 

M.L.U. or Mus. L. U. (see Linnaeus, 1764) 

Mus. Ad. Fr. (see Linnaeus, 1754) 

Mus. Tess. or Tessin (see Linnaeus, 1753) 

Needham microsc. (see Needham, 1745) 

Olear. mus. (see Olearius, 1674) 

Osb. iter, or Osbeck. iter, (see Osbeck, 1757) 

Pet. or Petiv. amb. or amboin. (see Petiver, 1713) 

Pet. or Petiv. gaz. (see Petiver, 1764) 

Pet. mus. (see Petiver, 1695) 

Plane, conch, (see Plancus, 1739) 

Reaum. act. paris. 1712, p. 163 (see Reaumur, 1712) 

Regenf. conch, (see Regenfuss, 1758) 

Roes. ins. or in.sect. (see Roesel von Rosenhof, 1746-61) 

Rond. aqu. or aquat. (see Rondeletius, 1554-55) 

Rond. or Rondel, pise, or test, (see Rondeletius, 1554- 
55) 



K. J. Boss, 1988 



Page 117 



Rumph. mus. (see Rumpf, 1705, 1711, 1741) 

Scheuch. Diluv. (see Scheuchzer, J. J., 1716) 

Scheuch. helv. (see Scheuchzer, j., 1708) 

Seb. or Seba mus. (see Seba, 1734-65) 

Sellii historia Teredinis. Traject. or Sellii monogr. ultra 

or Sellii Tered. (see Sellius, 1733) 
Sloan, jam. (see Sloane, 1696) 
Stobaei Diss, epist. Lund, (see Stobaeus, 1732) 
Strom, sondm. 173 (see Strom, 1762) 
Swamm. or Swammerd. bibl. (see Swammerdam, 1737- 

38) 
Tessin. epist. 1, n. 28 (not found; apparently a letter to 

Linnaeus) 
Tournef. iter, (see Tournefort, 1717) 
Tulp. obs. (see Tulpius, 1739) 
Vallisn. nat. (see Valiisnieri, 1721) 
Wolff, hass. (see Wolfart, 1719) 
Worm. mus. (see Worm, 1655) 



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ACKNOWLEDGEMENTS 

I wish to thank Dr. Elizabeth A. Shaw% Bibliographer 
and Research Taxonomist in the Gray Herbarium, Mrs. 
Geraldine C. Kaye of the Fariow Herbarium Library, 
Ms. Barbara Callahan of the Harvard University Her- 
barium, and Eva Jonas, Dana Fisher, Roxane Coombs, 
and Mina Brand of the library of the Museum of Com- 
parative Zoology. Mr. Richard I. Johnson also made avail- 
able rare malacological volumes, and valuable criticisms 
were offered by Mr. Richard E. Petit and two anonymous 
reviewers. 



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schienen sind. Erster Band. Bijcherkunde. Hiilfsmittel, all- 
gemeine Schriften, vergleichende Anatomic und 
Physiologic. Zoologie. Palaeontologie. Verlag von Wilhelm 
Engelmann, Leipzig, p. vii, 786. 

Hanley, S>lvanus C. T. 1855. Ipsa Linnaei Conchy lia. Wil- 
liams and Norgate. London. 556 p.. pis. I-V. 

Heimann. Willy. 1957. .\ catalogue of the works of Linnaeus. 
Issued in commemoration of the 250th anniversary of the 
birthday of Carolus Linnaeus 1707-1778. Sandbergs An- 
tikvariatsforteckning. Nr. 12 Varen. 1957. Sandbergs Bok- 
handel, Stockholm. 179 p. 

Heller. John L. 1961. List. Loqu ' and List Mut.'. two puz- 
zling Linnaean abbreviations Proceedings of the Linnean 
Society of London 173:61-64, pis. 1-4. 

Heller. John L. 1968. Zoological sources of Linnaeus. Journal 
of the Society for the Bibliographv of Natural History 5(1): 
76-77. 

Heller, John L. 1979. Bibliotheca Zoologica Linnaeana. Sven- 
ska Linnesallskapets arsskrift. argang 1978. Yearbook of 
the Swedish Linnaeus Society. Commemorative Volume. 
Uppsala, p. 240-264. 

Heller, John L. 1983. Studies in Linnaean method and no- 
menclature. Marburger Schriften zur Medizingeschichte, 
(Verlag Peter Lang, Frankfurt am Main, Bern, New York. ) 
7(IX):328 p. 

Keen. Angeline Mvra. 1983a. (January). On Linnaeus" book- 
shelf. The Festivus (San Diego Shell Club) 15(1):5-15. 24 
figs. 

Keen. Angeline Myra. 1983b (August 30). On Linnaeus' book- 
shelf [abstract]. .Annual Report of the Western Society of 
Malacologists 15:15. 

Linnaeus. Carolus [1707-78]. 1758. Caroli Linnaei . . . Sys- 
tema Naturae per Regna tria Naturae, secundum classes, 
ordines, genera, species, cum characteribus. differentiis, 
synonymis, locis . . . Editio decima reformata. Impensis 
Direct. Laurentii Salvii: Holmiae, Tom. 1. Animalia. p. 
[iv], 823. 

Linnaeus. Carolus [1707-78]. 1767. Caroli a Linne . . . Sys- 
tema Naturae per Regna tria Naturae, secundum classes, 
ordines. genera, species, cum characteribus. differentiis, 
synonymis, locis . . . Editio duodecima, reformata. Impen- 



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THE NAUTILUS, Vol. 102, No. 3 



sis Direct. Laurentii Salvii: Holmiae, Tom. 1 Regnum 
Animale. Pars II. Classis V-VI, Nomina Generica. Nomina 
specierum propria: Nomina trivialia. Nomina trivialia 
Papiliomim and Phalaenarum Synonyma Termini Artis. 
Appendi.x Svnonvniorum. Addenda. Errata, p. 533-1327 
[36]. 

Nissen, Glaus. 1969. Die Zoologische Buchillustration. Ihre 
Bibliographie und Geschichte. Band I. Bibliographie. An- 
ton Hiersemann, Stuttgart, p. 666. 

Soulsby, B. H. 1933. A catalogue of the works of Linnaeus 



(and publications more immediately relating thereto) pre- 
served in the libraries of the British Museum (Bloomsbury) 
and the British Museum (Natural History) (South Ken- 
sington). Second Edition Trustees of the British Museum, 
London, p. xi, 246; .\ddenda and Corrigenda, 68 p., 7 
plates. 
Wheeler, A. 1979. The sources of Linnaeus knowledge of 
fishes. Svenska Linnesallskapets arsskrift. argang 1978. 
Yearbook of the Swedish Linnaeus Society. Commemo- 
rative Volume. Uppsala, p. 156-211. 



THE NAUTILUS 102(3):123-124, 1988 



Page 123 



Frederick Benjamin Isely: Biographical Sketch and 
Malacological Contributions 



Mark E. Gordon 

Department of Zoology 
University of Arkansas 
Fayettevi'lle, AR 72701, USA 



Frederick B. Isely was known primarily for his pioneer- 
ing research in orthopteran ecology (mainly Acrididae 
and Tettigoniidae) and as an educator/administrator. 
However, he contributed significantly to the basic knowl- 
edge of unionacean biology and ecology. Isely began his 
studies with freshwater mussels in the Chikaskia River 
of north-central Oklahoma during 1906. These initial 
observations soon expanded to include aspects of union- 
acean ecology, growth, migration, and distributional sur- 
vey of mussels within the Arkansas and Red river basins 
of eastern Oklahoma. In these endeavors, he was finan- 
cially aided through an appointment as a scientific as- 
sistant with the U.S. Bureau of Fisheries during the sum- 
mers of 1910-13. As a result, five manuscripts were 
published between 1911 and 1931. Isely's innovative con- 
cepts for ecological investigations are reflected by his 
discussion in the 1914 growth paper of the use and value 
of replicate samples and of the problems encountered 
using mark/recapture techniques. One such problem was 
predation by small bo\s interested in collecting the brass 
tags used to mark the mussels. The distributional survey, 
although funded by and conducted for the U.S. Bureau 
of Fisheries, was delayed in publication due to govern- 
ment reassessment of funding priorities — World War I. 
This manuscript was published independently by Isely 
after the war. His last malacological publication resulted 
from the recovery of one of the "lost" growth study 
specimens 15 years following the completion of that in- 
vestigation. Further evidence for the value of replication 
in research. 

Frederick was born June 20, 1873 of Swiss parents 
(Christian Isely and Elise Dubach) at Spring Grove farm 
near Fairview, Brown County, Kansas. His early edu- 
cation was gained locally, culminating with his gradu- 
ation in 1894 from Hiawatha Academy (Hiawatha, Kan- 
sas). He entered Fairmount College (now Wichita State 
University) in its founding year 1895. He was quite active 
in college activities (e.g., debate team, class president 
1895-99) and sports (e.g., track, football) and was award- 
ed a B.S. in 1899, the first 4-year graduating class of the 
college. In 1909, he earned an M.S. from the University 
of Chicago and continued that summer with further study 
at the Marine Biological Laboratory. Later academic 



endeavors were pursued during the summers of 1929 
and 1931 at the University of Chicago and University of 
Colorado, respectively. 

His teaching career began with an appointment at 
Franklin Rural School, Brown County, Kansas (1894- 
95). Following his graduation from Fairmount College, 




Figure 1. Frederick B. Isely during his Trinity University 
years. Reproduced from a photograph given to the Department 
of Biology, Trinity University by Mrs. F. B. Isely (courtesy of 
H. D. Murray). 



Page 124 



THE NAUTILUS, Vol. 102, No. 3 



he assumed the principalship of Central School, Hia- 
watha (1899-1901) followed by stints as teacher of bi- 
ology at Wichita High School (1901-06) and University 
Preparatory School, Tonka wa, Oklahoma (1906-12). 
Equipped \s ith his M.S., he served as professor of biolog\' 
at Central College, Fayette, Missouri (1912-20) with so- 
journs during the summers of 1915-17 as instructor of 
biology at the L'niversity of Missouri. His administrative 
career was initiated with an appointment as dean and 
professor of biology at Ckilver-Stockton College, Canton, 
Missouri (1920-22) followed by his assumption of the 
same roles at Texas Women's College, Fort Worth (1922- 
31). He returned to a strictK teaching position as pro- 
fessor of biology at Trinity Universit\, Wa.xahachie and 
later San Antonio, Texas (1931-47) where he developed 
the majority of his orthopteran research. He also served 
as department chairman from 1931 to 1946, retiring from 
active teaching in 1946. In recognition of his achieve- 
ments. Trinity University awarded him the Sc.D. honoris 
causa upon his retirement. 

Isely married Mary E. Nickerson of Clearwater, Kan- 
sas, on May 8, 1901. They had four children and he 
appears to have influenced his famiK , as he did his stu- 
dents; biologists continue in the famih to date. In spite 
of a heavy teaching load, administrative duties, and re- 
search, he was active in community organizations 
throughout his life and still managed time for personal 
pleasures {e.g.. camping, gardening, keeping up with the 
latest episodes of Joe Palooka and Little Orphan Annie). 
Isely was a well-loved and enthusiastic instructor in ad- 
dition to being an active and respected scientist. He 
received grants-in-aid for his orthopteran research from 
the National Research Council (1935, 1936) and the 
American Philosophical Society (1937, 1941). He was a 
fellow and founding member of the Oklahoma Academy 
of Science (secretar\ 1901-12) and the North Texas Bi- 
ological Society (president 1924-27); a fellow of AAAS 
and the Texas Academy of Science (president 1938); and 
a member of the Society of American Zoologists, Eco- 
logical Society of America, Entomological Society of 
America, Kansas Academy of Science, Texas Entomo- 
logical Society, and Texas State Teachers Association. 
Frederick B. Isely died December 30, 1947 still active 
in research. Additional information may be acquired 
from two memoria (Alexander, 1949; Geiser, 1949). 

Recently, a remnant of Isely s malacological collection 
was rediscovered at the Stovall Museum, University of 
Oklahoma (Shepard, 1982). About 450 specimens of 
unionids, representing 37 species remain. Some inade- 



quately labeled lots may also be referable to Isely. Thirty- 
nine specimens including seven species represent vouch- 
ers for the 1914 growth paper. Fourteen detached brass 
tags are also present. The remainder of his collection is 
from the 1925 distribution sur\e\. Little else of Isely's 
malacological material, other than some papers in the 
archives of the Biology Department at Trinity University 
(H. D. Murra\, personal communication), remain. 

A small malacological collection of his brother, Dwight 
IseK , was redisco\ered at Wichita State L'niversity (Met- 
calf and Distler, 1984). Further information pertaining 
to D. Isely may be obtained in Miner (1976) and Metcalf 
and Distler (1984). 

I thank the following people for their kind assistance 
in providing data for this note: Mrs. Katherine Isely 
McGuire and Drs. Douglas G. Alexander, Harle\ P. 
Brown, Donald A. Distler, Duane Isely, Charles G. Lin- 
coln (deceased), Harold D. Murra\-, Robert W. Pennak, 
William D. Shepard, and Gary D. Snell. 

LITERATURE CITED 

Alexander, G. 1949. Frederick B. Isely (1873-1947), Ento- 
mology News 60:29-30. 

Geiser, S. W. 1949. Proceedings of the forty-fifth annual 
meeting of the American Society of Zoologists: 3. Memorial 
resolutions, Frederick B. Isely 1873-1947. Anatomical 
Record 103:233-234. 

Metcalf, A. L. and D. A. Distler. 1984. Gastropods collected 
from eastern Oklahoma by Dwight Isely in 1911. The 
Nautilus 98:135-137. 

Miner, F. 1976, Dwight Isely 1887-1974 Journal of Economic 
Entomology 69:289-290. 

Shepard, W. D. 1982. Rediscovery of a portion of the Isely 
unionid collection. The Nautilus 96:8. 

MALACOLOGICAL BIBLIOGRAPHY OF 
FREDERICK B. ISELY 

1911. Preliminary note of the ecolog> of the early juvenile life 
of the Unionidae. Biological Bulletin 20:77-80. 

1914. Experimental stud> of the growth and migration of fresh- 
water mussels. U.S. Bureau of Fisheries Document no. 
792, 26 p. 

1914. Mussel streams of eastern Oklahoma. U.S. Bureau of 
Fisheries Economic Circular no. 9, 6 p 

1925. The fresh-water mussel fauna of eastern Oklahoma. Pro- 
ceedings of the Oklahoma Academy of Science 4:43- 
118. 

1931. A fifteen vear growth record in fresh-water mussels. 
Ecology 12:616-619, 



THE NAUTILUS 102(3): 125-126, 1988 



Page 125 



Rediscovery of Planorhella magnifica (Pilsbry) in 
Southeastern North Carohna 



William F. Adams 

Environmental Resources Branch 
U.S. Armv Corps of Engineers 
P.O. Bo.x 1890 
Wilmington, NC 28402, USA 



Andrew G. Gerberich 

Division of Fishes 

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



Early in this century, Planorbis magnificus was de- 
scribed by Pilsbry (1903) from the lower Cape Fear River 
region of North Carolina. The type localitv for the species 
was given simply as "lower Cape Fear River". Bartsch 
(1908) correctly surmised that the species is an inhabitant 
of lentic environments and found it living in Greenfield 
Lake, a manmade impoundment lying to the south of 
what were then the city limits of Wilmington, NC, as 
well as in Orton Pond, then called Sprunt's Pond, ap- 
proximately 16 km south of Wilmington. Baker (1945) 
figured and studied portions of the anatomy of specimens 
collected b\' Bartsch and concluded that P. magnificus 
should be reassigned to Planorhella. This ta.xon repre- 
sents the largest known planorbid. 

Planorhella magnifica has not been reported since the 
accounts b\ Pilsbry and Bartsch and has been considered 
extinct by some (Opler, 1976; Imlay, 1977; Palmer, 1985). 
Fuller (1977) hypothesized that the species may still sur- 
vive in Orton Pond, a manmade pond similar to Green- 
field Lake in age and physiography, based on an obser- 
vation of large planorbid egg masses there by J. P. E. 
Morrison. 

The Greenfield Lake watershed has been almost totally 
developed since Planorhella magnifica was described, 
and now undergoes intensive management for control of 
nuisance algae. Management measures include appli- 
cation of algicides during the growing season and oc- 
casional drawdown during winter months. Fuller (1977) 
mentioned fruitless attempts by himself and others to 
find Planorhella magnifica in Greenfield Lake. Our in- 
vestigations of that lake indicate that it no longer provides 
a suitable habitat for this species. 

During July 1986, Planorhella magnifica was seren- 
dipitously rediscovered in Orton Pond, Brunswick Coun- 
ty, NC, while collecting water samples. Three living 
specimens were obtained in 20 minutes. Four empty 
shells were also collected from the drift line on the shore. 
During another water-sampling trip in January 1987, an 
attempt was made to collect live specimens for captive 
propagation. This effort was unsuccessful as much of the 
aquatic vegetation had died back. However, two more 
empty shells were found washed up on the shore. 

Our samples of Planorhella magnifica vary consid- 



erably in size. The maximum shell diameters of the spec- 
imens collected alive are 35.5, 21.5, and 16.3 mm. Vouch- 
er specimens from our collection have been deposited in 
the collections of the North Carolina State Museum of 
Natural History (NCSM P468-P471) and the National 
Museum of Natural History (USNM 857935). Because of 
restricted access, additional trips to Orton Pond have not 
been made. 

Live specimens of Planorhella magnifica were found 
on the stems and undersides of the floating leaves of 
Spatterdock, Niiphar liiteum (Sibthorp & Smith, 1809), 
and Fragrant Waterlily, Nymphaea odorata (Aiton, 1789). 
Water depth where living specimens were collected was 
approximately 1 meter, and the bottom substrate was 
organic. Orton Pond closely matches Bartsch's (1908) 
description of habitat suitable to P. magnifica in Green- 
field Lake. 

Like Greenfield Lake, Orton Pond is a manmade im- 
poundment. Both were created early in the last century 
to serve as a fresh-water source for rice agriculture. Orton 
Pond exemplifies a type of lentic waterbody unique in 
southeastern North Carolina. Although blackwater lakes 
and ponds in the region typically are acidic, Orton Pond's 
waters have a circumneutral pH, ranging from 6.2 
through 7.9 (Smock and Lenat, 1978). We suspect that 
Orton Pond may have a direct connection to waters of 
the Castle Hayne aquifer, an Eocene limestone deposit. 
The molluscan fauna of Lake Waccammaw (Columbus 
County, NC), the only other sizable circumneutral-pH 
waterbody in the region, has been investigated frequent- 
ly (Fuller et al, 1976; Porter, 1985), but Planorhella 
magnifica has not been found in that system. Lake Wac- 
camavv differs from Orton Pond in being much larger, 
a natural waterbody, and is not a blackwater system. 

The owners of Orton Pond undertake little manage- 
ment of the pond and manage the lands that surround 
it for timber and wildlife. With a continuation of this 
type of management, the near future for Planorhella 
magnifica seems secure. However, not all of the pond s 
watershed is protected in this way. Therefore, potentially 
adverse developments could occur quite some distance 
from the pond. 

Because of the apparent uniqueness of the waters of 



Page 126 



THE NAUTILUS, Vol. 102, No. 3 




Figure 1 . Shells of Planorbella magnifica collected alive from 
Orton Pond, Brunswick County, North Carolina, July 1986 
(apprcximately natural size). 



Orton Pond and absence of Planorbella magnifica in 
Greenfield Lake and Lake W'accamaw, it appears that 
the species is extremely restricted in distribution and can 
be found today only in Orton Pond. We believe that any 
changes in the quality of the surface waters in Orton 
Pond's drainage basin or in its underlying groundwater 
regime may have a deleterious effect on P. magnifica. 
Due to the uncertainty of the future of the pond, an in- 
depth study of the autecology of the species is needed 
so that potential conservation measures can be developed 
and implemented should they prove necessary. 



LITERATURE CITED 

Baker, F. C. 1945. The molluscan family Planorbidae. Uni- 
versity of Illinois Press, Urbana, p. 1-530. 

Bartsch, P. 1908. Notes on the fresh-water mollusk Planorbis 
magnificus and descriptions of two new forms of the same 
genus from the southern states. Proceedings of the United 
States National Museum 33:697-700. 

Fuller, S. L. H. 1977 Freshwater and terrestrial mollusks. /n: 
Cooper. J. E . S S Robinson, and J B Funderburg (eds. ). 
Endangered and threatened plants and animals of North 
Carolina. North Carolina State Museum of Natural His- 
tory, Raleigh, p. 143-194. 

Fuller, S. L. H., M. J. Imlay, and J. D. Williams. 1976. En- 
dangered or threatened fresh-water mussels (Mollusca: 
Bivalvia; Unionidae) of the Waccamaw River basin of the 
Carolinas. Association of Southeastern Biologists Bulletin 
23:60. 

Imlay, M. J. 1977. Competing for survival Water Spectrum 
9:7-14. 

Opler, P. A. 1976. The parade of passing species: a survey of 
extinctions in the U.S. Science Teacher 43(9):30-34. 

Palmer, S, 1985. Some extinct molluscs of the U.S.A. .Atala 
13(l):l-7. 

Pilsbry, H. A. 1903. The greatest American Planorlns. The 
Nautilus 17:75-76. 

Porter, H. J. 1985. Rare and endangered fauna of Lake Wac- 
camaw, North Carolina watershed s\ stem, \'ols. 1 and 2. 
North Carolina Endangered Species Restoration: Job Title 
No. VI-7. North Carolina Wildlife Resources Commission, 
Raleigh, p. 1-358. 

Smock, L. A. and D. R. Lenat. 1978. Preliminary stud\ of 
the effects of salt water intrusion on the macrobenthic 
community of Orton Pond, North Carolina. U.S. Army 
Engineer District, Wilmington, p. 1-48. 



THE NAUTILUS 102(3):127-128, 1988 



Page 127 



Bite by Octopus joubini: A Case Report 



Donald M. McKinstry 

Division of Science, Engineering, and 

Technology 

The Pennsylvania State University 

at Erie 

The Behrend College 

Erie, PA 16563-1200, USA 



While octopuses are capable of delivering toxic bites, 
human case reports are scarce (Halstead, 1978). This 
report describes a bite by Joubin's octopus (Octopus jou- 
bini Robson, 1929) in a 12-year-old boy. The incident 
occurred on Sanibel Island, Florida, on January 2, 1987 
at about 0900. A storm had deposited a variety of marine 
animals on shore from the Gulf of Mexico. Not realizing 
that octopuses are venomous, the boy, Patrick Reed 
McKinstry, picked up a small specimen and was prompt- 
ly bitten on the dorsal surface of the right thumb prox- 
imal to the interphalangeal joint. He pulled the octopus 
away from his thumb within 10 seconds. The actual bite, 
described as a "sharp pinch", was quickly followed by 
a burning sensation of greater intensit% than a bee-sting. 
A tiny red spot surrounded by a white circular area 6 
mm in diameter was then noticed. Bleeding was minimal, 
a few small drops. Over the next se\ eral hours the thumb 
became moderately swollen and felt sore, stiff, and, at 
times, somewhat numb. These symptoms, to a lesser de- 
gree, extended to the wrist. Medical attention was not 
sought because only local effects, which gradually di- 
minished by day one, were noticed. By day three, the 
thumb was slightly swollen, reddened, and sore, and a 
tiny hole was noticed at the site. Over the next several 
days, the thumb became itchy and was treated with 3% 
hydrogen peroxide solution. This was the first treatment 
of the bite. By day eight, the swelling and redness had 
disappeared but some soreness in the metacarpopha- 
langeal joint was experienced. The hole had enlarged to 
2 mm in diameter and displayed a slightly irregular 
border. During the period up to day 12, the hole became 
darker and a red circular area of up to 7 mm in diameter 
developed around it. Itching continued but soreness in 
the joint area decreased. On day 12, a small amount of 
clear yellow fluid oozed from the hole. The area was 
then treated with hydrogen peroxide solution and anti- 
biotic ointment. The signs and symptoms gradually di- 
minished, and by day 32 the hole was replaced by a 
slightly indented circular scar 1 mm in diameter, sur- 
rounded by a slightly darkened firm area 6 mm in di- 
ameter. By da\- 100, onK' the indented scar was evident 
and the wound was considered completeK healed. .A 



shiny, hard circular scar, 1 mm in diameter, was still 
evident by 1 year. While in excellent health, Patrick is 
allergic to hymenopteran venoms and to penicillin. How- 
ever, this was his first exposure to octopus venom, thus, 
an allergic reaction to explain the bite effects was dis- 
counted. Secondary bacterial infection could have con- 
tributed to the effects experienced, however, signs of 
infection, e.g., pus, lymphangitis, and fever, were not 
observed. 

The octopus died after about 2 hours in a pail of sea 
water and was preserved in 70% isopropyl alcohol. It was 
identified as a nearly reproductively mature male spec- 
imen of O. joubini by Dr. Ronald Toll. This species, the 
smallest of the western Atlantic octopuses, ranges from 
southern Florida to the Bahamas and the West Indies. 
Specimens are often washed ashore by storms on the Gulf 
coast of Florida (Meinkoth, 1981). The dorsal surface of 
the specimen is deep tan, while the arms and ventral 
surface are light tan. The mantle is 3.2 cm in length. I 
have donated this specimen to Dr. Toll s collection. 

Two accounts of envenomation following the bite of 
O. joubini were found. In the first (Anonymous, 1965), 
a physician was bitten on the finger while handling a 
specimen washed ashore on Sanibel Island, Florida. He 
experienced a "sharp" bite with a little bleeding, quickly 
followed by '"the most excruciating pain which rapidly 
became almost intolerable." Within a few hours, the 
finger became very swollen, hard, and deep red. By the 
next day, the inflammation and pain had extended to 
the hand. Later, some white discharge was noted from 
the bite wound. The swelling and hardness gradually 
subsided over a period of 2 months. The only systemic 
effect was a mildly elevated temperature. He noted that 
"there was never any evidence of bacterial invasion or 
infection, only an extreme degree of cellulitis. ' 

The second case (Wittich, 1968), occurred on a re- 
search vessel off Egmont Key, Florida. A Florida Board 
of Conservation worker was bitten on the back of the 
hand while sorting marine specimens. In this case, '"a 
sharp, piercing sensation " was felt, and a "severe pain" 
extended to the upper arm. The bite wound bled slightK 
and almost instantK became surrounded by a "'pure white 



Page 128 



THE NAUTILUS, Vol. 102, No. 3 



welt of about 25 mm diameter." In the following hour, 
the victim experienced pain and sw elling around the bite, 
accompanied by nausea, headache, and fever. Improve- 
ment gradually occurred, but after 3 days the area was 
still sensitive and swollen. The swelling persisted, and 
some itching and a serous discharge from the wound 
occurred. After 1 month, a 2 mm wound surrounded by 
a 6 mm red periphery was present. The wound then 
healed quickK. 

In the case I report, the bite effects were not as severe 
as those noted above even though the victim was a child 
with less tissue mass to dilute the venom. However, the 
amount of venom delivered could have been minimal. 

ACKNOWLEDGEMENTS 

I am grateful to the following for their helpful assistance 
and suggestions: Ronald Toll, The L'niversity of the South, 



Sewanee, TN; Sherman A. Minton, Jr , Indiana Univer- 
sit\ , School of Medicine, Indianapolis, IN; Harry N. Cun- 
ningham, Jr., Kath\' Mauro, and Robert Rose, all of The 
Pennsylvania State University at Erie, The Behrend Col- 
lege, Erie, PA. A special thanks is due Patrick R. Mc- 
Kinstrv for his account of the octopus bite. 

LITERATURE CITED 

.\nonymous. 1965. Florida octopus bite. Sea Secrets 9(7):5. 
Halstead, B. W 1978. Poisonous and venomous marine 

animals of the world The Darwin Press, Princeton, NJ, 

1043 p. 
Meinkoth, N. A. 1981. The Audubon Society field guide to 

North American seashore creatures. .\if red .\. Knopf, New 

York, NY, 799 p. 
Wittich, A. C. 1968. Account of an octopus bite. Florida 

Academy of Science Quarterly Journal 29l4):265-266. 



THE NAUTILUS 102(3):129, 1988 



Page 129 



Occurrence of Mites in Mexican Land Snails 



0§car J. Polaco 

Departamento de Prehistoria 
Institute Nacional de Antropologia 

e Historia 
Moneda #16 
Mexico 06060, Mexico 



Wolfgang Mendl 

Lab. Acarologia 
Departamento Zoologia 
Escuela Nacional de 
Ciencias Biologieas 
Apartado Postal 42186 
Mexico 11340, Mexico 



To date there has only been a single report (Baker, 1945) 
of mites infecting the mantle cavity of a land snail from 
Mexico (Helix pomatia Linne, 1758, an introduced 
species). The mite was identified as Riccardoella oude- 
mansi (Thor, 1932), however, the mite shown by Baker 
(1945) is similar to the figure published by Fain and Van 
Goethem (1986) for Riccardoella limacum (Schrank, 
1776) (Prostigmata: Ereynetidae). This mite has been 
studied by several European authors (Turk & Phillips, 
1946; Baker, 1970) and reported as cosmopolitan. 

In this note, we report findings of mites parasitizing 
the mantle cavities of two species of land snails of the 
famiK Bulimulidae and one species referable to the fam- 
ily Helicidae. The bulimulid Bidimulus unicolor (Sow- 
erby, 1833), collected in Tomas Garrido, in the state of 
Quintana Roo, was parasitized by Riccardoella limacum. 
with one protonymph and one deutonvmph collected 
from nearly 50 specimens of the snail. Of 79 specimens 
of Rhabdotus alternatus (Say, 1830), another bulimulid 
collected in Xicotencatl in the state of Tamaulipas, that 
were examined, one contained three mites of the genus 
Boydaia (Ereynetidae) and another contained a single 
specimen of Eupodes voxencollinus (Sig Thor, 1934). 
This is the first report of the last two mite species on 
land snails. 

Examination of other land snail species of the families 
Helicidae, Achatinidae, Succinidae, Oleanicidae, and 
Polygyridae revealed no evidence of parasitism by mites, 



with the exception of a single specimen of Helix aspersa 
(MiJller, 1774), an introduced species collected in Mexico 
City, which was infected with two specimens of E. vox- 
encollinus. As no mites were found in any Mexican land 
snails from the above families, the family Bulimulidae 
remains the only new record of a host for mites. 

The examined snails are deposited in the Mollusk Col- 
lection of the Departamento de Prehistoria, I.N.A.H., 
and the mites in the Acarology Collection of the Labo- 
ratorio de Acarologia of the E.N.C.B. 

We acknowledge two anonymous reviewers for their 
critical comments to improve this paper. 



LITERATURE CITED 

Baker, E. W. 1945. Five mites of the family Ereynetidae 
from Mexico. Journal of the Washington Academy of Sci- 
ences .35:16-19. 

Baker, R. A. 1970, The food of Riccardoella limacuni (Schrank) 
(Acari: Trombidiformes) and its relationships with pul- 
monate mollusks. Journal of Natural History 4:521-530. 

Fain, A. and J. L, VanGoethem. 1986. Les acariens du genre 
Riccardoella Berlese, 1923, parasites du poumon de Mol- 
lusques gasteropodes pulmones terrestres. Acarologia 27(2): 
125-140. 

Turk, F. A. and S. M. Phillips. 1946. A monograph of the 
slug mite Riccardoella limacum (Schrank). Proceedings of 
the Zoological Society of London 115:448-472. 



THE NAUTILUS 102{3):130, 1988 



Page 130 



Axelella, New Name for Olssonella Petit, 1970, a Preoccupied 
Taxon (Mollusca: Cancellariacea) 



Richard E. Petit 

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



The cancellariid genus Olssonella Petit, 1970, was pro- 
posed for a rather compact and well-defined group of 
species from the Later Tertiary and Recent faunas of 
the Americas. An Eastern .Atlantic species was subse- 
quently placed in this genus by Bouchet and Waren 
(1985;263), and the morpholog\ of the type species was 
published b\ Harasew\ch and Petit (1984). The ta.xon 
has been used b\ a number oi other authors since it was 
proposed. Unfortunately, it has just been noted that 
Olssonella is preoccupied, and a replacement name is 
here proposed. 

Axelella, new name for Olssonella Petit, 1970:83. 
Not Olssonella Glibert & \'an de Poel, 1967:121. 

Type species, by original designation of Olssonella 
Petit, 1970, Cancellaria smithii Dall, 1888, Recent, 
Western .\tlantic. 

This new name honors the late A.xel \. Olsson, as did 
the name it replaces, and the name which makes the 
replacement necessary. 

Appreciation is expressed to Dr. Philippe Bouchet, 



Museum National d'Histoire Naturelle, Paris, for calling 
the prior use of Olssonella to m\ attention. 

LITERATURE CITED 

Bouchet, P. and A. Waren. 1985. Revision of the Northeast 
Atlantic bathval and ab\ssal Neogastropoda excluding 
Turridae (Mollusca: Gastropoda). Bollettino Malacologico, 
Supplemento 1:121-296 

Dall, W. H. 1888. Mollusks. In: Alexander Agassiz, Three 
Cruises of the United States Coast and Geodetic Survey 
Steamer Blake 2:62-75. 

Glibert, M. and L. Van de Poel. 1967, Les Bivalvia fossiles 
du Cenozoique etranger des collections de I'lnstitut Royal 
des Sciences Naturelles de Belgique. \". Oligodontina. In- 
stitut Royal des Sciences Naturelles de Belgique, Memoires 
(2nd ser.') 83:1-152. 

Harasew ych, M. G. and R. E. Petit. 1984 Notes on the mor- 
phology of Olssonella smithii (Gastropoda: Cancellari- 
idae). The Nautilus 98(l):37-44. 

Petit, R. E. 1970. Notes on Cancellariidae (Mollusca: Gastrop- 
oda) — II. Tulane Studies in Geologv and Paleontology 8(2): 
83-88. 



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Volume 102, Number 4 
December 21, 1988 
ISSN 0028-1344 

A quarterly devoted 
to malacology. 



Marine Biological Laboratory 
LIBRARY 

JAN 3 1989 



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EDITOR-IN-CHIEF 
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National Museum of 
Natural History 
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National Museum of 
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THEt^NAUTILUS 



CONTENTS 



Volume 102, Number 4 

December 21, 1988 

ISSN 0028-1344 



Nidia S. Romer 
Donald R. Moore 



A new species of Alvania (Rissoidae) from the West Indian 
region 



131 



Charles N. D'Asaro 


Micromorphology of neogastropod egg capsules 


134 


Philippe Bouehet 


Two new species of Metula (Gastropoda: Buccinidae) with 
a description of the radula of the genus 


149 


Kurt Auffenberg 
Harry G. Lee 


A new species of intertidal Terebra from Brazil 


154 


Douglas C. Smith 


Notes on the biology and morphology of Margaritifera 

hemheli (Conrad 1838) (Unionacea: Marearitiferidae) 


159 








Alan R. Kabat 
Richard E. Petit 


The two printings of J. F. Gmelin's Systema Naturae. 13th 
Edition (1788-96) 


164 



Marine Biological Laboratory 
LIBRARY 

JAN 3 1989 



Woods Hole, Mass. 



THE NAUTILUS 102(4):131-133, 1988 



Page 131 



A New Species of Alvania (Rissoidae) from the 
West Indian Region 



Nidia S. Ronier 
Donald R. Moore 

Rosenstit'l School of Marine 
and Atmospheric Science 
l'niversit> of Miami 
4600 Bickenbacker Causeway 
Miami, FL 33149, USA 



ABSTRACT 

Alvania (Alvania) colombiana new species was found from 45 
to 261 m (24 to 143 fms) depths off the Caribbean coast of 
Colombia to South Florida, the Gulf of Mexico, and Puerto 
Rico. Alvania colombiana differs from Rissoa xanthias (Wat- 
son, 1886), in having a protoconch with a single spiral zig-zag 
line of tiny nodules just above the suture, and by its smaller 
shell size. Alvania colombiana also differs from Rissoa precip- 
itata (Dall, 1889), by its protoconch and size. 



INTRODUCTION 

Exploration of the deep sea a century ago was the equiv- 
alent of the space age today. The famous "Challenger" 
expedition sampled two offshore stations in the tropical 
western Atlantic in which small mollusks were numerous. 
These were described by Watson (1886). Among the 
Rissoidae were two species, Rissoa pyrrhias (Watson, 
1886) and Rissoa xanthias (Watson, 1886), that were 
similar in size and shape. Dall (1889), reporting on the 
"Blake" material, described two more species of Rissoa, 
Rissoa acuticostata and Rissoa precipitata. Rissoa acu- 
ticostata was similar to Rissoa xanthias. and finally Dall 
(1927) acknowledged that with more specimens it was 
impossible to separate the material into two species. Ris- 
soa precipitata, on the other hand, is known principally 
from the original description and illustration. 

There is another small western Atlantic rissoid which 
at first glance appears similar to the illustration of Rissoa 
precipitata. However, there are a number of features 
that differentiate them. Size is one: the small species has 
an average adult size of 1.2 mm while Rissoa precipitata 
(holotype) has an adult size of 3.92 mm. Protoconch is 
the other: the small species has a projecting protoconch 
while Rissoa precipitata has a depressed protoconch. One 
of us (D.R.M) had obtained specimens of the small species 
during the MAFLA (Mississippi, Alabama, and Florida) 
study (1974 to 1975) in the eastern Gulf of Mexico. This 
small species was first reported by one of us (N.S.R.) as 
Alvania sp. 1 (see literature cited; Rodriguez, 1983). 



ABBREVIATIONS 

USNM = National Museum of Natural History, Smith- 
sonian Institution. 

FSBS I = Florida Department of Natural Resources, 
Marine Research Laboratory, St Petersburg. 

MCZ = Museum of Comparative Zoology, Harvard Uni- 
versity. 

UMML = University of Miami Marine Laboratory. 

CNMS = Colombian Natural Museum of Science, Na- 
tional University, Bogota 



MATERIALS AND METHODS 

All the specimens to be photographed in the SEM were 
cleaned in 100% sodium hypochlorite for 2 minutes, then 
rinsed in distilled water twice. If the specimens still 
showed residual material (e.g., sand grains), they were 
sonicated for 60 seconds. Finally, the specimens were air 
dried and put on a SEM stub with double-sided Scotch 
tape. 



Alvania (Alvania) colombiana new species 

(figures 1, 2) 

Description: Shell 1.0 to 1.3 mm in length. Protoconch 
multispiral, glossy, light brown, with 3'/2 whorls that are 
sculptured with 1 spiral zig-zag line of tiny nodules just 
above the suture and 1 undulating line of the same nod- 
ules in the middle of the whorl. Nodules irregularly ar- 
ranged over entire protoconch. Teleoconch of approxi- 
mately 2 whorls, with numerous axial ribs. Adult shell 
translucent, very light brown in color. 

Axial ribs slightly curved, 26 ribs on the body whorl 
fading out on the base of the shell. Body whorl with a 
narrow spiral groove just below the suture and another 
7 to 9 grooves between the periphery of the final whorl 
and the base of the shell. Varix terminal with a narrow 
extension forming outer lip of semilunate aperture. Some 
specimens weakly umbilicate. 



Page 132 



THE NAUTILUS, Vol. 102, No. 4 




Figures I, 2. Alvania colomhiana new species. I. Hololype, USNM miriiber 8.59339, 1.2S mm in length. 2. Protoconch, 280 x. 
Figures 3, 4. Rissoa xanthias (Watson, 1886), from off Miami, in the junior author's collection, 2.26 mm in length. 4. Protoconch, 
220 X. 



Type locality: Off the west coast of Florida at 29°35'N, 
87°20'06"W, depth 107.3 m. 

Holotype: USNM number 859339. Length, 1.28 mm. 
W'idtii, U.78 mm. 

Paratypes: One from off the west coast of Florida at 
29°35'N, 87°20'06"W, depth 107.3 m, USNM number 



859340; 2 from off the west coast of Florida at 28°24'N, 
85°15'06"W, depth 164.6 m, FSBC I number 33113; and 
off the west coast of Florida at 29°49'30"N, 86°25'30"W, 
depth 82.3 m, FSBC 1 number 33114; 2 from off Puerto 
Rico at 17°53'24"N, 66°35'10"W, depth 221 m, MCZ 
number 297220; and off Miami at 25°47'N, 80°01'30"W, 
depth 137 m, MCZ number 297219; 2 from off Miami 



N. S. Romer and D. R. Moore 1988 



Page 133 




Figure 5. Drawing of Alvania precipitata (Dall, 1889), ho- 
lotype, MCZ 7470, 3.92 mm in length. 



at 25°47'N, 80°01'30"W, depth 137 m, UMML number 
8349; and off Miami at 25°46'30"N, 80°00'08"W, depth 
76.81 m, UMML number 8350; 2 from off the Caribbean 
coast of Colombia at 09°52'35"N, 75°47'25"W, depth 72 
m, CNMS number ICN-MHN(MO)522; and off Miami 
at 25°46'30"N, 80°00'08"W, depth 76.81 m, CNMS num- 
ber ICN-MHN(MO)523. 

Distribution: This species was found in 31 samples from 
off the Caribbean coast of Colombia collected between 
1979 and 1981. The specimens were found at depths 
ranging from 45 m to 261 m, but were most abundant 
at depths between 65 m and 160 m. Other specimens of 
this species were found from off the west coast of Florida, 
off southern Puerto Rico, and the Straits of Florida east 
of Miami. The bottom in almost all cases consisted of 
sandy mud. The species is probabK' found in these depths 
throughout the Caribbean, the southern and eastern Gulf 
of Mexico, and the Bahamas. All the specimens were 
found dead. 

Discussion: Alvania colombiana appears to be a com- 
mon widespread species throughout the Caribbean and 
adjacent areas. This is a very small compact species that 
should not be confused with any other in its range. Some 
features, however, are lacking or lost on the protoconch 
of some specimens. These include the fine nodules dis- 
tributed all over the protoconch, and the undulating line 
of nodules on the periphery. These features are seen only 
under high magnification, so are difficult to observe for 
routine identification. Details of the teleoconch sculp- 
ture, however, should be easy to make out under low 
power, even in a somewhat worn specimen. 



Alvania colombiana has been confused with Rissoa 
precipitata (Dall, 1889) (figure 5). This was due to the 
fact that specimens of Rissoa precipitata were not avail- 
able for comparison. The holotypc and only known spec- 
imen, MCZ 7470, is a thin shell v\itli no terminal varix. 
In fact, the sculpture fades away almost completely on 
the last half whorl (except for the sub-sutural tubercles). 
The protoconch is of the form seen in gastropods without 
a planktonic larval stage: large, rounded, unsculptured, 
and with a smooth transition from protoconch to teleo- 
conch. The type locality is Yucatan Strait at a depth of 
640 fathoms (not 670 as is given by Dall, 1889:280). 

Another similar species is Rissoa xanthias (Watson, 
1886) (figures 3 and 4). It, however, is about twice the 
size, has just over half as many axial ribs, and has a 
different and very distinctive protoconch. In this species 
the protoconch has 3'/2 whorls, in which the sutures are 
deep and ornamented with a line of spirally arranged 
vertical pustules. Just above the suture there is a wide 
canal whose edges are formed by an undulating line with 
vertical pustules equally spaced and directed downward. 
Each canal also has in the middle dots and small nodules 
randomly distributed. Watson reported it from Chal- 
lenger Sta. 24 off Culebra Island, 18°38'30"N, 65°05'30"W, 
depth 715 m, and from Sta. 122, 9°10'S, 34°49'W to 
34''53'W, depth 640 m. 

Finally, Rissoa pyrrhias (Watson, 1886) is another sim- 
ilar species. It is slightly larger than Rissoa xanthias, and 
its spiral grooves are not confined to the base of the shell. 
According to Watson (1886), the protoconch has 2'/2 whorls 
which are scored with coarse but feeble spiral threads. 

ACKNOWLEDGEMENTS 

We are very thankful to Dr. Patricia Blackwelder and 
her assistant team at the SEM lab for their collaboration 
in the use of and picture taking on the SEM. We also 
thank Dr. Kenneth J. Boss for the loan of the holotype 
of Rissoa precipitata. 

LITERATURE CITED 

Dall, W. H. 1889. Reports on the results of dredging ... in 
the Gulf of Mexico (1877-78) and in the Caribbean Sea 
(1879-80) by the U.S. Coast Survey Steamer "Blake" . . . 
29. Report on the Moilusca, part II. Gastropoda and Sca- 
phopoda Bulletin of the Museum of Comparative Zoology 
18:1-492. pis. 10-40. 

Dall, W. H. 1927 Small shells from dredgings off the southeast 
coast of the United States b\ the United States Fisheries 
Steamer "Albatross" in 1885 and 1886. Proceedings of the 
L'nited States National Museum 70(2667);1-134. 

Rodriguez, (Romer) N. S. 1983. Micromoluscos recientes de 
la plataforma Caribe Colombiana entre Bocas de Ceniza 
e Isla Fuerte. Thesis, Universidad Jorge Tadeo Lozano, 
247 p. 

Watson, R. B. 1886. Report on the Scaphopoda and Gaster- 
opoda collected by H. M. S. Challenger during the years 
1873-1876. Voyage of H. M. S. Challenger, Zoology 15: 
1-675; appendix A, 677-680; geographical distribution, 
691-722; index, 723-756; pis. 1-50. 



THE NAUTILUS 102(4):134-148, 1988 



Page 134 



Micromorphology of Neogastropod Egg Capsules 



Charles N. D'Asaro 

Department of Biology 
L iii\ersit\ of West Florida 
Pensacola, FL 32514, USA 



ABSTRACT 

Egg capsule micromorphology of eight species of neogastropods 
{Chicoreus florifer dileclu.s. PhijHonotus pomum. Cantharus 
inuUangiilus, C. cancellariti.s. Conus floridanus floridensis, C. 
ju.spidcus stearnsi. Cranulina ovuliforniis. and Marginella au- 
reocincta) was studied with light microscopy hy examining 
very thick, toluidine blue stained sections Laminae exposed 
hy fracturing the sections, and their reactions to the stain, 
provided the characters used to describe micromorphology. 

The results showed that muricaceans and buccinaceans have 
complex but microstructurally similar egg capsules, while co- 
nids and marginellids have egg capsules with taxonomically 
distinct rnicrostructural characteristics. In the Muricacea and 
Buccinacea, four structural laminae of similar origin and func- 
tion are usually present in the capsule wall, the second from 
the outermost having the most complex pattern of fibers and 
the greatest thickness. The third outermost lamina is continuous 
with one or more components sealing the escape aperture. In 
some buccinaceans, only a trace of the third lamina exists in 
the wall. Four layers, including one or two mucoid plugs, close 
the escape aperture. Conid egg capsules differ in that they 
include only three structural laminae in the capsule wall and 
three layers, including a riuicoid plug, sealing the escape ap- 
erture. Microstructurc and fiber pattern in the middle lamina 
are probably unique to this family. Egg capsules of marginellids 
have a distinctive thin, dense wall not separated into laminae 
and lack an escape aperture closed by a mucoid plug. There 
is a preformed suture that fractures at hatching in the wall of 
most marginellid egg capsules. 

Ke\i words: Reproduction; egg capsules; Neogastropoda; Chi- 
coreus. Phyllonotiis. Cantharus, Conns, Cranulina: Margi- 
ncllti 



INTRODUCTION 

Encapsulation of early ontogenetic stages is typical of 
higher gastropods, especially neogastropods. The highly 
refractory, layered envelopes of carbohydrate and pro- 
tein are structurally and chemicalK complex (Hunt, 1971; 
Flower, 1973; Goldsmith et ai, 1978; Gruber, 1982; Sul- 
livan and Maugel, 1984). Encapsulation and the for- 
mation of egg masses provides protection and has con- 
siderable survival value (Tamarin and C^arriker, 1967; 
Pechenik, 1979, 1983; Abe, 1983; Lord, 1986). Although 
there are thousands of neogastropod species, it still has 
not been clearly established how the egg capsules are 



formed in the oviduct and ventral pedal gland. Nor is it 
known if a common capsular microstructurc exists in 
various neogastropod taxa. 

Macromorphology of the often species-specific egg 
capsules and egg masses produced by neogastropods has 
been studied more frequently than any other aspect of 
neogastropod egg encapsulation. This was illustrated in 
the review of prosobranch reproduction by Webber 
(1977), in a more recent review of egg encapsulation for 
all mollusks by Pechenik (1986), and in descriptive re- 
ports on the external structure of neogastropod egg cap- 
sules by Bandel (1975. 1976a,b,c, 1982) and D'Asaro 
(1986a). 

In early reports on neogastropod capsule-wall micro- 
morphology (e.g., .\nkel, 1937; Hancock, 1956), three or 
four laminae were described, including specific patterns 
of fibers. However, it was not established if these are 
structural relationships common to the species studied 
or to higher taxa. Fretter (1941) provided for Nttcella 
lapillus (Linne, 1758), Ocenebra erinacea (Linne, 1758), 
Nassarius reticulatus (Linne, 1758), and Buccinum un- 
datiim Linne, 1758 the most frequently cited descrip- 
tions of how these neogastropods construct, in the oviduct 
and ventral pedal gland, three or four lay ered capsules. 

To characterize the laminated egg capsule of Uro- 
salpinx cinerea follyensis B. Baker, 1951, Tamarin and 
Carriker (1967) published the first comprehensive study 
employing light microscopy — including polarized light — 
and electron microscopy. They described a capsule with 
four laminae composed of asymmetrical protein-like 
molecules bound into collagen-like matrices. They also 
noted that the great structural complexity of the capsular 
wall could only be partialh explained b\ Fretter's (1941) 
description of the formative process. Bayne (1968) con- 
ducted a histochemical study of several gastropod egg 
capsules, including those of Nucella lapillus, and found 
mucopolysaccharides in at least three capsular laminae 
of that species. He was one of the first to use toluidine 
blue for this purpose. Flower et al. (1969) and Flower 
(1973) described the origin and the ultrastructure of neo- 
gastropod capsular proteins. In another histochemical 
study, (Jruber (1982) described six structural layers in 
the capsule wall of Eupleura caudata etterae B. B Baker, 
1951, and three additional layers associated with tlie 
escape aperture and la\ ers of albumen. 



C. N. D'Asaro 1988 



Page 135 



Table 1. Comparative micromorphology of muricacean egg capsules stained in most cases with toluidine blue (meta = meta- 
chromatic, mp = mucoid plug, nr = not reported, ortho = orthochromatic, pre = present, tt = two types of albumen cells in the 
oviduct, z = zone). 















Lasers 








Width 










closing 


Lavers ot 




Species 


of wall 


LI 


L2 


L3 


L4 


aperture 


albumen 


Author 


Chicoreus florifer 


64 ^m 


meta. 


meta. 


meta. 


meta. 


LI, mp, L3 


two meta 


this report 


dilectus 




4 ^m 


54 ixm 


2 Mm 


4 Mm 


(mp), L4, 
315 Mm 






Phylloiiotus po- 


51 nm 


meta. 


ortho in 


meta. 


meta. 


LI, L3 (mp). 


two? meta 


this report 


imini 




1-2 MITl 


part. 

47 /um 


1-2 Mm 


2 Mm 


mp, L4, 

429 Mm 






S'uccllu lapillus 


nr 


meta 


pre 


meta 


nr 


Li'' (mp) 


meta matrix 


Ankel, 1937; 
Fretter, 
1941; 
Bayne, 1968 


Urosalpinx cine- 


109 Mm 


ortho. 


ortho. 


pre, 2 Mm 


ortho. 


outer z, dense 


pre 


Hancock, 


rea 




30 Mm 


75 Mm 




2 um 


z, L3 or dif- 
fuse z, L4, 
500 Mm 




19.56; Tam- 
arin and 
Carriker, 
1967 


Ocenebra erinacea 


nr 


nr 


pre, two fi- 
brous 
lavers 


pre, mu- 
coid lay- 
er 


nr 


mp 


tt 


Fretter, 1941 


Eupleura caudata 


nr 


meta, (1)* 


(2-5) 


(6?-7) 


meta (8) 


(1, 6?, 7 = 


two meta 


Gruber, 1982 


ctterac 












mp, 8) 







* Ten lasers were described; possible relationships for eight are indicated in parentheses. 



In the most comprehensive and detailed study to date, 
Sullivan and Maugel (1984) used transmission and scan- 
ning electron microscopy, histochemistry, and poly- 
acrylamide gel electrophoresis to examine the egg cap- 
sule of Ihjanassa obsoleta (Say, 1822). They were able 
to determine the physical and chemical structure of the 
capsule wall and confirm the origin of four structural 
laminae in the oviducal capsule gland. A fifth, nonstruc- 
tural layer was identified as a product of the ventral 
pedal gland. 

Since the previous work did not consider egg capsule 
micromorphology in a taxonomic perspective, the pur- 
pose of this stud) was to describe the complex laminar 
structure of egg capsules from two species in each of 
four diverse neogastropod taxa: Muricacea, Buccinacea, 
Marginellidae, and Conidae. These descriptions, as well 
as those in a few papers with similar data, were used to 
compare neogastropod egg capsule micromorphology in 
order to determine if there is a common wall structure 
in the selected taxa, and indirectly, if a common mech- 
anism to produce neogastropod egg capsules exists. 



MATERIALS AND METHODS 

The eight species from St. Joseph Bay in northwest Flor- 
ida selected for study are listed in tables 1-3. Egg capsule 
macromorphology of each species is known (refer to the 
Results for citations and museum voucher numbers). 
Specimens for sectioning were preserved in 10% buffered 
seawater formalin, a fixative considered acceptable be- 



cause egg-capsule proteins are very stable in a range of 
acids and bases and are only slightly soluble when au- 
toclaved in water (Hunt, 1971). Capsules were taken 
from near the center of well-formed egg masses, or if 
deposited individually, they were selected to represent 
the average size and shape for that species. 

To determine the number of egg capsules per species 
that must be sectioned to account for possible variations 
in staining and micromorphology, preliminary histolog- 
ical procedures were performed on two egg capsules 
from each of five egg masses of Cantharus multangulus 
(Conrad, 1846) deposited by different females. Although 
the sectioned capsules were not microstructuralh iden- 
tical, a recognizable pattern of structural laminae existed 
among all capsules produced by the same and different 
females of this species. Because the pattern for C. mul- 
tangulus could be recognized in each sectioned capsule 
and the literature suggested that it existed for other neo- 
gastropod species, sections for the remaining seven species 
were prepared from only two or three egg capsules pro- 
duced by different females. 

.After fixation, the larger egg capsules were opened 
along one edge in the longest axis with a scalpel to ac- 
celerate diffusion of the embedding fluids. Following 
dehydration in ethyl alcohol, capsules were cleared in 
toluene and embedded in paraffin (melting point = 56 
°C). 

Capsules were sectioned at 20-22 um. All sections were 
parallel to the longest axis and included some that passed 
through the sealed escape aperture and the basal plate. 
To separate structural laminae, the capsule wall was frac- 



Page 136 



THE NAUTILUS, Vol. 102, No. 4 







Figure 1. Schematic drawing of a longitudinally sectioned 
egg capsule of Cbicurciis florifer dilectiis. Figure 2. 
Drawing, like figure 1, of a PhijUonoius pomum egg capsule. 
Numbers with arrowheads indicate positions at which photo- 
graphs for figures 3-12 were taken. Empty spaces in figures 1 
and 2 are artifacts produced during dehydration. A = adhesive, 
DA = dense albumen, E = embryo, IMP = inner mucoid plug, 
LI = metachromatic first lamina — solid line, L2 = second 
lamina — light stipple, L3 = metachromatic third lamina — solid 
line, L4 = metachromatic fourth lamina — broken line, OMP 
= outer mucoid plug, T.'\ = thin albumen. L2 is metachromatic 
in figure 1 and partially orthochromatic in figure 2. 



turcd and partially delaminated by varying the speed at 
\\ liicli the microtome blade cut a section. 

To demonstrate mucopolysaccharides, sections were 
stained with toluidine blue for 1.5-2.0 minutes according 
to the method of Gurr (1962:440). They were then rap- 
id!) delnclrated in ethyl alcohol, cleared in toluene, and 
mounted in methyl methacrylate. 

Egg capsule microstructure was examined using light 
microscopy. Observation of the preliminary sections sug- 
gested that toluidine blue metachromasia would produce 
a range in color from pur[)le through heliotrope (reddish 
purple) but rarely red. Also, some layers would be or- 
thochromatic (blue) or colorless. In the intact capsule 
wall, structural laminae were difficult to trace by follow- 
ing metachromatic la\ers alone. However, when a struc- 



tural lamina was first isolated and defined in a fractured 
or delaminated section, then the lamina could be rapidly 
and comparably traced in whole, stained sections. If a 
capsule fractured and delaminated repeatedK' in a con- 
sistent pattern, then the parts were considered distinct 
structural laminae produced by different types of cells 
or produced at different times. Those less than a few 
micrometers wide could be recognized when they sep- 
arated from the sections and folded to one side. Very 
thick sections also allowed microsculpture on the outer 
laminar surface to be examined {e.g., figure 3). Following 
Tamarin and Carriker's (1967) method, structural lam- 
inae were identified from outermost to innermost as Ll- 
L4. Diagrams of median sections were prepared with a 
drawing tube mounted on a compwund microscope, while 
photomicrography was used to record microstructure at 
selected positions on a section 

Measurements of the capsule wall, structural laminae, 
and the sealed escape aperture were taken from the 
middle regions of each (tables 1-3). These dimensions 
should be considered approximations, as Sullivan and 
Mangel (1984) suggested, because of possible shrinkage 
introduced by the method and the wide range in width 
occurring at approximately the same position on different 
capsules of the same species. 

RESULTS 

Chicoreus florifer dilectus (A. Adams, 1855) 
(figures 1, 3-8; table 1; USNM 860426) 

Egg capsules of this muricid are distinctly vasiform in 
section (figure 1). Macromorpholog\ was described by 
D'Asaro (1970:420, fig. 3). 

The capsules have a thick, highK fibrous wall in which 
four structurally distinct components can be identified: 
LI, a thin, fineK fibered, metachromatic outer lamina; 
L2, a metachromatic lamina, with layered, coarse fibers 
forming the bulk of the wall; L3, the thinnest meta- 
chromatic lamina, closely applied to the innermost layer; 
and L4, a metachromatic lamina surrounding the al- 
bumen (figures 3, 4; table 1). LI is so transparent that 
the coarser L2 fibers can be seen through it. Except on 
the stalk and basal plate, LI is folded in a corrugated 
maruier (figures 3, 4, os). L2 may have fibers directed at 
right angles or parallel to the long capsule axis and ar- 
ranged with regional differences in two or three fused 
layers. When three fibrous components are present, the 
middle one has the coarser fibers arranged parallel to the 
long axis (figure 3, L2). Where the wall was folded during 
formation, L2 fibers separate, forming large \acuoles 
(figure 4, v). These vacuoles are not stained and are 
probably filled with a nonmucoid liquid. L2 fibers fuse 
with a more or less homogeneous inner component in 



Figures 3-8. Sections of the Chicoreus florifer dilectus egg capsule. 3. Wall showing laminae 4. Wall just below the apical plate. 
5. .Albumen fibers. 6. Mucoid plugs in the escape aperture at the junction with 1.2 7. Wall in the stalk 8. Basal plate and adhesive. 
Positions at which photographs were taken are indicated on figure 1. a = adhesive, da = dense albinnen, f = fracture, fl,3 = 



C. N. D'Asaro 1988 



Page 137 




w. .. - V *;^ * A*" I 'fly J -^ V . 



•ts 






%T:y ta 



m 








50 u m 
• ' — I 



fragment of L3, fiL2 licmmm luuus component of L2, imp = inner mucoid plug, isz = intensely stained zone, LI = metachromatic 
first lamina, L2 = metachromatic second lamina, L3 = metachromatic third lamina, L4 = metachromatic fourth lamina, os = 
outer surface of LI, omp = outer mucoid plug, sL2 = outer portion of L2 in the stalk, ta = thin albumen. 



Page 138 



THE NAUTILUS, Vol. 102, No. 4 







Figures 9-12. Sections of the Phyllonotuti pnmum egg capsule. 9. Wall showing all laminae. 10. Escape aperture toward the 
base. 11. Escape aperture toward the ape.x 12. Stalk and basal plate. Positions at which photographs were taken are indicated on 
figure 2. a = albumen, cf = circular fibers, e = embryo, hL2 = homogeneous component of L2, imp = inner mucoid plug, Li = 
metachromatic first lamina, L2 = partially orthochromatic second lamina, L3 = metachromatic third lamina, L4 = metachromatic 
fourth lamina, omp = outer mucoid plug, os = outer surface of LI. sL2 = outer portion of L2 in the stalk, r = ridge. 



contact with L3 (figures 3, 4, hL2). L3 stains less inten- 
sively than L4. In the apical plate, L3 appears to fuse 
with the inner mucoid plug and may have similar com- 
position, .^s described by Ankel (1929: fig. 1) for Nucella 
lapilhis and Franc (1940: fig. 2) for Ocinelmna aciculata 
(Lamarck, 1822), L4 completely surrounds the albumen 
and probably contains it during assembly of the capsule 
while other components are more fluid. Hereafter, L4 
will be called the albinnen retaining layer. 

Two structurally distinct types of albumen occur: an 
outer layer including metachromatic fibers with random- 
ly attached granules in which all embryos arc situated 
(figures I, TA; 5, ta) and, an inner core including denser 
strongly metachromatic fibers (figures I, DA; 5, da). Fi- 
bers of dense albumen ma\ fuse in a continuous layer 
where the albuminous components meet; however, the 
fused layer did not delaminate during sectioning. The 
thin outer albumen does not envelop the denser core 
basally (figure 1). 



The apical escape aperture is closed by a structuralh 
complex barrier composed of four la\ers (figures 1. 6). 
LI iscontiguouswith the outer layer. The metachromatic 
(purple) second layer or outer mucoid plug is larger than 
the aperture and interdigitates with but is clearly sepa- 
rated from the homogeneous iimer component of L2 
(figure 6, omp, hL2). Fractures across the w idth of the 
outer mucoid plug apparently caused during sectioning 
occurred (figure 6, f). The third layer is a lenticular, 
heliotrope mucoid plug, larger than the aperture, w hich 
sometimes fractures during sectioning. L3 is fu.sed to the 
lenticular or inner plug in an intensely stained zone (fig- 
ure 6, isz). L4 is the innermost layer closing the escape 
aperture and is fused to L3 in the same intenseK stained 
zone. 

The stalk and basal plate are formed from the capsule 
wall and an adhesive material (figures 1, A; 7, 8, a). The 
outer portion of both includes LI and up to the longi- 
tudinal, fibrous component of L2 (figure 7, sL2), while 



C. N. D'Asaro 1988 



Page 139 



the base of the capsular lumen is formed from the re- 
maining parts of L2 as well as L3 and L4. Mucoid ad- 
hesi\e, which has orthochromatic and metachromatic 
zones and occasional L2 fibers, fills the center of the stalk 
(figures 7, 8, a). 

Phyllonotus pomtim (Gmelin, 1791) 
(figures 2, 9-12; table 1; USNM 860425) 

The somewhat tongue-shaped capsules and the rather 
massive, communal egg masses were described by Perr\ 
and Schwengel (1955: fig. 338), D'Asaro (1970:422, fig. 
3), Radwin and Chamberlin (1973:107, fig. 1), Bandel 
(1976a:10, fig. 4), and Moore and Sander (1978:253, fig. 
2). Unlike Chicoreus florifer egg capsules, those of Phyl- 
lonotus pomum may have two points of attachment w ith 
a supporting substratum that nearly always is a conspe- 
cific capsule. Sections of such capsules are very variable 
in outline. 

Phyllonotus pomum capsules have a distinctly fibrous 
wall that is only partially metachromatic (table 1). Four 
structural parts e.\ist: LI, a thin, metachromatic outer 
lamina; L2, a thick, coarsely fibered and partially or- 
thochromatic lamina; L3, a strongly metachromatic com- 
ponent; and L4, a metachromatic albumen retaining lay- 
er (figures, 2, 9). LI is as transparent as the homologous 
lamina in Chicoreus florifer dilectus and is corrugated 
on or near folds in the wall (figures 10, 11, os). L2 remains 
almost unstained. It has a core of longitudinal fibers bor- 
dered in most areas by a few circular fibers (figure 9, 
L2, cf). Each side of the fibrous zone may be bordered 
by a narrow, faintly orthochromatic, homogeneous zone 
(figure 9, hL2). L3 has no obvious fibers and appears 
fused to the outer mucoid plug of the escape aperture, 
as noted later. L4 has distinct circular fibers arranged in 
a single layer. 

Albumen, including fibrous and amorphous material, 
is present in sections only near the retaining layer (figure 
10, a). This distribution is an artifact, as noted in Can- 
tharus cancellarius, caused by loss of the more fluid 
contents during dehydration. Embryos are distributed in 
both components of the albumen. 

Positioned on one side of the capsule, the sealed escape 
aperture includes four layers (figures 10; 11, LI, omp, 
imp, L4). The outer component is formed from LI. The 
second outermost layer, including intensely stained 
metachromatic regions, is continuous with but distinct 
from the homogeneous proximal part of L2, and appears 
to be fused or continuous with L3. This pattern is dif- 
ferent from that of Chicoreus florifer dilectus. A third 
layer, which is metachromatic, mucoid with small ves- 
icles, and exceedingly fragile, bulges into the capsular 
lumen, where it is bordered by L4, the albumen retaining 
layer. The more apical and lateral borders of the escape 
aperture, formed from L2, have an inward projecting 
ridge (figure 11, r). 

The basal plate is formed from LI and the outer half 
of L2. A separation occurs in a dense reticulum or vac- 
uolated zone near the midline of the lamina (figure 12, 




Figure 13. Schematic drawing of a longitudinally sectioned 
egg capsule of Canlharus multanguliis. Figure 14. Drawing, 
like figure 13, of a Cantharus cancellarius egg capsule. Empty 
spaces in figures 13 and 14 are artifacts produced during de- 
hydration. Numbers with arrowheads indicate positions at w hich 
photographs for figures 15-23 were taken. A = adhesive, .\D 
= apical depression, BM = basal mucoid material, D.A = dense 
albumen, E = embryo, LI = orthochromatic first lamina — 
solid line, L2 = second lamina — light stipple, L4 = metachro- 
matic fourth lamina — broken Une, MP = mucoid plug, TA = 
thin albumen 



sL2). Where the outer half of L2 was in contact with the 
substratum, a homogeneous zone having some fibers ex- 
tending through it toward the substratum is present. The 
inner half of L2, L3, and L4 form the floor of the capsular 
lumen. No adhesive layer was observed on capsules that 
had been attached to those of conspecifics. 

Cantharus multangulus (Philippi, 1848) 
(figures 13, 15-19; table 2; USNM 947143) 

Macromorphology of the egg capsule was described by 
Perrv and Schwengel (1955: fig. 340), Radwin and Cham- 
berlin (1973:110, fig. 6), and D'Asaro (1986a:85, fig. 3). 
Median sections through the ribbed and spined capsule 
are roughK vasiform in outline and have a rounded, 
apical depression partly covered by a transverse ridge 



Page 140 



THE NAUTILUS, Vol. 102, No. 4 




Figures 15-19. Sections of the Canlhartis mullai^gulus egg capsule 15. Wall showing laminae 16. Escape aperture adjacent 
to apical depression 17. Escape aperture on side opposite apical depression. 18. .Alliunien layers 19. Stalk and adhesive. Positions 
at which photographs were taken are indicated on figure 13. a = adhesive, ad = apical depression, bm = basal mucoid material, 
da = dense albumen, LI = orthochromatic first lamina, L2 = second lamina, L2m = metachromatic component, L2o = ortho- 
chromatic component, mp = mucoid plug, os = outer surface of LL ta = thin albumen. 



C. N. D'Asaro 1988 



Page 141 



Table 2. Comparative niicromorplinlogy of buccinacean egg capsules stained in 
c'hronialic, nip = mucoid plug, nr = not reported, ortho = orthoclironiatic, pre = 
oviduct) 



most cases with toluidine blue (mcta = meta- 
present, tt = two types of albumen cells in the 



Species 



Width 

of wall LI 



L2 



L3 



L4 



Layers 

closing Layers of 

aperture albumen 



Author 



Cantharus ntul- 
langulus 



23 iim ortho, ortho and vestige meta, 

1-2 /im meta, 1-2 /tm 

21 ^m 

Cantharus cancel- 22 fim ortho, ortho and vestige meta, 

larius 2 ^m meta, 3— i ^m 

17 urn 
Btisycon carica nr pre pre pre pre 

and B. canalicu- 
latum 
Nassarius reticiila- nr three layers of uncertain relationship mentioned 

tus 
Buccinium undo- nr pre two fibrous mucoid nr 

turn layers layer 

Ilyanassa obsoleta 11-22 meta pre, meta, 1 ^m pre, 60 nm 

(LO pre) Mm 2 nm 9-10 ^m 



LI, L2, mp. 


two 


meta 


this report 


L4, 








210 /im 








LI, L2, mp, 


two 


meta 


this report 


L4, 








117 Mm 








nr 


two 




Harasewych, 

1978 


nr 


tt 




Fretter, 1941 


nr 


tt 




Fretter, 1941 


LI, L2, L3 


nr 




Sullivan and 


(mp), L4, 






Maugle, 


110-150 






1984 


Min 









(figure 13, AD). In the intact capsule, the depression is 
not obvious when viewed apically. Cantharus cancel- 
larius (Conrad, 1846) has similar sculpture on its apical 
plate (figure 14, AD). 

Capsules of C. miiltangulus have a wall in which the 
fibers appear fused in most regions when viewed with 
light microscopy. There are three distinct components 
that appear homologous to three of the four muriccacean 
laminae: LI, a dense, orthochromatic outer lamina; L2, 
a mostly homogeneous central lamina, including vacuoles 
in some regions and fibers in others; and L4, a retaining 
layer surrounding the albumen (figure 15; table 2). LI 
stains intensely and may appear black. L2 may have up 
to four layered but fused components with the outermost 
being metachromatic and the innermost orthochromatic. 
The innermost component, like LI, is intensely stained 
and may appear black (figure 15, L2o). L2 has somewhat 
tubular vacuoles in the metachromatic area, especially 
near the apical depression and where the wall is folded 
into ridges or spines (figures 16, ad, L2m; 17, L2m). L3, 
which in many muricaceans is continuous with a mucoid 
plug in the escape aperture, is not a distinctive feature 
in the capsule wall (see the following comments con- 
cerning the escape aperture). The albumen retaining 
layer is characteristically wrinkled when it separates from 
the wall (figure 15, L4). 

Albumen occurs in two metachromatic components: 
a dense portion, broadest near the apical plate, containing 
short, closely packed fibers with attached, irregular gran- 
ules, and a thinner, more diffuse portion with coarser 
fibers and granules, filling most of the lumen (figures 15, 
da; 18, da, ta). Embryos are positioned in the latter. 

The escape aperture is closed by four layers. LI, the 
outer component, is in continuous contact with the or- 
thochromatic part of L2 (figures 16; 17, L2o). The or- 



thochromatic part of L2 is somewhat folded, intensely 
stained, and may fracture vertically during sectioning. 
The broadest layer closing the aperture is a metachro- 
matic, mucoid plug lying between the orthochromatic 
part of L2 and L4 (figures 13, MP; 16, 17, mp). Apically, 
the mucoid plug lies between L2 and L4 in the homol- 
ogous position for L3 in muricaceans, but it does not 
extend to the base of the lumen as a continuous lamina 
(figure 13). However, above the stalk between L2 and 
L4, there is some metachromatic mucoid material with 
chromotropic characteristics similar to those of the apical 
plug (figures 13, BM; 19, bm). This mucoid material 
could have been formed at the same time as the plug 
and would be homologous to L3. L4 is the innermost 
layer closing the escape aperture. 

The stalk and basal plate are formed from LI and 
most of L2, while the base of the capsular lumen is 
formed from the orthochromatic part of L2, some mu- 
coid material, and L4 (figure 19, L2o, bm, L4). The stalk 
has a core of metachromatic mucoid adhesive (figure 19, 
a). 

Cantharus cancellarius (Conrad, 1846) 
(figures 14, 20-23; table 2; USNM 847140) 

The macromorphology of the egg capsules of this species 
was described by Radwin and Chamberlin (1973:110, 
fig. 8) and D'Asaro (1986a:84, fig. 2). Most median sec- 
tions of the capsules are cylindrical to vasiform in outline 
(figure 14). A rounded depression in the apical plate, 
similar to one observed in C. multangulus, is prominent 
and partially covered by a raised transverse ridge folded 
toward the escape aperture. 

As in C. multangulus, much of the capsule wall ap- 
pears to be composed of fused fibers. Three structural 



Page 142 



THE NAUTILUS, Vol. 102, No. 4 




Figures 20-23. Sections of the Cantharus cancellarius egg capsule. 20. Wall showing laminae 21. Escape aperture opposite 
apical depression. 22. Escape aperture adjacent to apical depression. 23. Basal plate and adhesive. Positions at which photographs 
were taken are indicated on figure 14. a = adhesive, ad = apical depression, bm = basal mucoid material, da = dense albumen, 
LI = orthochromatic first lamina, L2 = second lamina, L2m = metachromatic component, L2o = orthochromatic component, mp 
= mucoid plug, OS = outer surface of LI, sL2 = outer portion of L2 in the stalk, ta = thin albumen. 



laminae, homologous to three of the four muricacean 
laminae, are present. These are: LI, a very thin, ortho- 
chromatic outer lamina; L2, an orthochromatic and 
metachromatic central lamina with tubular vacuoles; and 
L4, a metachromatic albumen retaining layer (figure 20; 
table 1). LI is often denser on the apical plate. Distinct 
tubular vacuoles are present in the central portion of L2, 
especially where ridges e.xist in the wall and apical plate. 
Longitudinally directed vacuoles are positioned nearer 
to the outer surface, while more circularly directed vac- 
uoles are nearer to the ituier surface. Apically, where the 
wall is folded, fibers in the lamina are visible (figure 21). 
As shown for C multangulus, L3 is not present as a 
distinct and continuous lamina throughout the whole 
capsule. The metachromatic albumen retaining layer is 
extremely thin and can be identified only where it is 
pulled away from the albimien or the capsule wall and 
folded to one side (figure 20, L4). Dense metachromatic 



albumen with fine fibers surrounds more coarsely fibered, 
less dense albumen in which embryos are positioned (fig- 
ure 20, da, e, ta). 

The escape aperture is closed by four layers arranged 
in a pattern essentially identical to that of C. multan- 
gulus. Apically, LI forms the outermost layer (figures 
21, 22). The expanded, orthochromatic part of 1,2 is the 
second component. It has \ertical folds or fractures (fig- 
ures 21; 22, L2o). The broadest component is a meta- 
chromatic mucoid plug lying between the orthochro- 
matic part of L2 and L4 (figures 21; 22, mp). This plug 
tapers basally between L2 and L4, but does not extend 
to the base as a continuous lamina equivalent to L3. 
There is a layer of similar mucoid material positioned 
between L2 and L4 at the base of the lumen (figures 14, 
BM; 23, bm). 

Basally, the plate is formed from LI and most of L2 
(figures 14; 23, sL2). Only the orthochromatic, proximal 



C. N. D'Asaro 1988 



Page 143 



part of L2 extends across the base, where it, some mucoid 
material, and L4 form the floor of the lumen. Unlike 
most specimens of C. multangulus. the lumen is sepa- 
rated from the substratum b\ only a narrow layer of 
metachromatic adhesive (figure 23, a). 

Contis jioridanus flohdensis Sowerby, 1870 

(figures 24, 26-29; table 3; USNM 847141) 

Median capsular sections typically have the longitudinal 
axis angling away from the axis of the stalk and show a 
slightly irregular outer surface (figure 24). Macromor- 
phology was described by Perry and Schwengel (1955: 
fig. 360, as C. spurius atlanticits Clench, 1942) and 
D'Asaro (1986a:88, fig. 4). 

The capsule wall has three distinct components: LI, a 
fibrous metachromatic outer lamina; L2, a finely fibered, 
orthochromatic central lamina; and L4, a thin, meta- 
chromatic albumen retaining layer (figure 26; table 3). 
LI is very transparent and has fibers circular in cross 
section. Occasional corrugations mark this lamina, es- 
pecially near the escape aperture and external ridges 
(figure 27, LI). Fibers in L2 tend to form a cross-hatched 
pattern in the upper three quarters of the capsule, but 
some align with the long axis in the stalk and the base 
(figures 26-29). Globular structures buried in the fibrous 
layer occur frequently (figure 27). No homologue of L3 
is present as a distinct layer, nor is there evidence that 
the mucoid plug in the escape aperture is homologous 
to L3. The albumen retaining layer (L4) is not obviously 
fibrous. Although this layer is the third in sequence, it is 
designated as L4, the albumen retaining layer, because 
its position and probable function appear the same as L4 
in Vluricacea and Buccinacea. 

in this species, only one layer of fibrous, metachro- 
matic albumen is visible (figure 26, al). Granules are 
attached to the widely separated fibers. Embryos are 
randomly positioned in the albumen. 

The sealed escape aperture is less complex than similar 
structures formed by muricaceans or buccinaceans. It is 
lined on its outer surface by LI and on its inner surface 
by L4. L2 is replaced by metachromatic, layered mucoid 
plug with which it interdigitates extensively (figure 27, 
mp). The degree of interdigitation suggests that L2 and 
the mucoid plug were formed at the same time. 

The stalk is composed of LI and almost all of L2 from 
both sides, while the basal plate on each side is composed 
of LI and L2 from the same side (figures 28, 29, sL2). 
A fibro-mucoid, metachromatic adhesive, which may ex- 
tend into the stalk, attaches the basal plate to the sub- 
stratum (figure 29, a). Fibers from L2 radiate widely into 
the adhesive, especially where it extends into the stalk. 

Conus jaspideus stearnsi Conrad, 1869 
(figures 25, 30-32; table 3; USNM 847148) 

When sectioned, the lamellate capsules, described by 
D'Asaro (1986a:88, fig. 4), appear pointed at the apex 
with a broad stalk and an escape aperture on one side. 




Figure 24-. Schematic drawing of a longitudinally sectioned 
egg capsule of Conus floridanus floridensis. Figure 
25. Drawing, like figure 24, of a Conus jaspideus stearnsi 
egg capsule. Numbers with arrowheads indicate positions at 
which photographs for figures 26-32 were taken. \ = adhesive, 
.\L = albumen, E = embryo, LI = metachromatic first lamina 
— solid line, L2 = second lamina — light stipple, L4 = meta- 
chromatic albumen retaining layer — broken line, MP = mucoid 
plug. 



In this plane, the basal plate is the widest part (figure 
25). 

Three laminae were identified: LI, a finely fibered, 
metachromatic outer lamina; L2, a complex orthochro- 
matic and metachromatic lamina with fibrous and mu- 
coid components; and L4, a metachromatic albumen re- 
taining layer (figure 30; table 3). LI, with circular fibers, 
is tightly fused to the fibrous portion of L2. Except near 
the escape aperture and on the stalk, this layer is uni- 
formly corrugated (figures 30; 31, os). L2 has an ortho- 
chromatic outer component with coarse, circular fibers 
(figure 30, L2). The inner component of L2 consists of 
at least three metachromatic layers of mucoid material 
containing scattered fibers arranged parallel with the 
long axis (figures 31, 32, mL2). Of the species studied, 
only C. /. floridanus has a similar arrangement of fibers 
in mucoid material, and that occurs only in the stalk. No 
distinct lamina equivalent to L3 is present. L4 may have 
fragments of albumen fused to its inner surface (figure 
32, is). 

Albumen in this species is metachromatic and includes 
scattered transparent spherules and irregular granular 
material that may form layers (figures 30-32). Embryos 
are distributed throughout it. 



Page 144 



THE NAUTILUS, Vol. 102, No. 4 



LI 







n. 




Figures 26-29. Sections ot the Conu:> jioridanm floridensis egg capsule. 26. Wall showing laminae. 27. Escape aperture. 28. 
Stalk 29. Basal plate and adhesive. Positions at which photographs were taken are indicated on figure 24 \ = adhesive, al = 
albumen, is = inner surface of L4, LI = metachromatic first lamina, L2 = orthochromatic second lamina, L4 = metachromatic 
albumen retaining layer, mp = mucoid plug, as = outer surface of LI, sL2 = outer portion of L2 in the stalk. 



Table .'{. Comparative micromorphology of conid and marginellid egg capsules stained in most cases with toluidine blue (meta 
irutai hromatic. mp = mucoid plug, ortho = orlhochromatic. pre = present). 



Species 



Width 
of wall 



LI 



L2 



L.3 



L4 



Lasers 

closing 

aperture 



Layers of 
albumen 



Author 



Conus floridanus 49 ^m meta, 
floridensis .5 nm 

Conus jaspidius 25 niu meta, 
slearnsi 2-3 Mm 



Granulina ovuli- 
formia 



\r> fim 



Marginella aureo- 12 ^m 
cincta 



ortho, not pre 

40 M'li 

ortho and not pre 
meta, 
22 nm 
one lamina present 



one lamina present 



meta, LI, mp, L4, one meta 

3-4 nn\ 330 ^m 

meta, I.I, mp. L4. one meta 

1-2 ^m I N ^m 

suture in wall two meta 



suture? two meta 



this report 
this report 



this report; 

D Asaro, 

1986 
this report; 

D'.Asaro, 

1 9.Sfi 



C. N. D'Asaro 1988 



Page 145 



LI forms the outer covering of the escape aperture, 
w hile the inner lining is L4. A metachromatic, mucoid 
plug replaces L2 (figure 31, mp). The edges of the mucoid 
plug are separated from L2 b> an interdigitating meta- 
chromatic boundary zone, similar but less complex than 
that of C. /. floridensis. None of the scattered fibers in 
the metachromatic mucoid layers of L2 enter the mucoid 
plug. 

The two outer laminae form the stalk and basal plate, 
w hile the base of the capsule is composed of the albumen 
retaining layer and the inner, mucoid part of L2 (figure 
32, mL2). The stalk varies in length; therefore, in some 
specimens the lumen of the capsule may be nearly level 
with the substratum. A dense mucoid adhesive is present 
basalK (figure 32, a). 

Granulina ovuUjormis (Orbigny, 1841) 
(figure 33; table 3; USNM 836973) 

Macro- and micromorphology of the egg capsules, based 
in part on eosinophilic features, were described by D'Asaro 
(1986b: 196, figs. 3-5). Because of structural differences, 
marginellid capsule micromorphology is not describable 
with reference to Tamarin and Carriker's (1967) system 
of enumerating capsule laminae. 

The simple, pustulate capsules are constructed from 
an inner component, enclosing two layers of albumen 
and an embryo, and basement and outer components 
that sandwich and fuse the previously mentioned struc- 
ture between them (D'Asaro, 1986b). Because these com- 
ponents were uniformly unstained and did not delami- 
nate during sectioning, the capsule wall is defined as 
having a single structural lamina of uniform composition, 
possibK formed by the same portion of the oviduct that 
produced the layers of L2 in muricaceans and bucci- 
naceans (figure 33, cw). The outer surface of the capsule 
is coated with metachromatic mucus. No escape aperture 
with a mucoid plug exists. The point at which the capsule 
wall fractures at hatching is marked by a distinct meta- 
chromatic suture (figure 33, s). Dense granular albumen 
lies just below the capsule wall, while a less dense com- 
ponent immediately surrounds the single embryo. Both 
albuminous layers stain metachromatically. The adhesive 
on the basal layer is not stained (figure 33, a, bl). 

Marginella aureocincta Stearns, 1872 
(figure 34; table 3; USNM 836974) 

Macro- and micromorphology of the egg capsule were 
described by D'Asaro (1986b: 195, figs. 3-5). 'With ap- 
plication of toluidine blue, all parts of the capsule are 
metachromatic, except the unstained basement adhesive. 
The capsule wall does not delaminate and stains intensely 
to the point of obscuring its layered structure suggesting 
that, as in Granulina ovuliformis, it should be defined 
as a single structural lamina (figure 34, cw, osw). Dis- 
tinctive granulations, described by D'Asaro (1986b: 195, 
fig. 3), are obvious on the outer surface. No specific 
fibrous layer surrounds the albumen. The metachro- 
matic, finely granular albumen occurs in two compo- 




Figures 30-32. Sections of the Conus jaspideus stearnsi egg 
capsule. 30. Wall showing laminae 3 1 . Escape aperture toward 
apex. 32. Stalk, basal plate, and adhesive. Positions at which 
photographs were taken are indicated on figure 25. a = ad- 
hesive, al = albumen, is = inner surface of L4, LI = meta- 
chromatic first lamina, L2 = orthochromatic and metachro- 
matic second lamina, L4 = metachromatic albumen retaining 
ia\er, mL2 = mucoid component of L2, mp = mucoid plug, 
OS = outer surface of LL 



Page 146 



THE NAUTILUS, Vol. 102, No. 4 





Figure 33. Section of the GranuHna ovitlifurmis egg capsule attached to a Thalassia leaf. Figure 34. Section of the Marginella 
aureocincta egg capsule, a = adhesive, bl = basement layer, cw = capsule wall, da = dense albumen, osw = outer surface of 
capsule wall, s = suture, ta = thin albumen. 



nents, a narrow band of dense material and a more fluid, 
central zone in which the embryo lies (figure 34, da, ta). 

DISCUSSION 

Separation of structural laminae by fracturing and to- 
luidine blue staining allows differentiation and identifi- 
cation of homologous components of neogastropod egg 
capsules. Laminae can be identified in this manner be- 
cause the fibers of which they are composed appear to 
have been secreted and chemically bound together dur- 
ing separate phases of assembly. Separate or combined 
functions related to protection, structural support, release 
of larvae or juveniles, and albumen retention are inferred 
for the structural laminae. 

,\mong the iieogastropods studied, three microstruc- 
tural patterns of laminae were identified, each charac- 
terizing specific higher taxa. Complex, four-layered cap- 
sules with an escape aperture sealed by four layers, 
including two that are mucoid, were found in nearly all 
muricaceans and buccinaceans examined. Three-layered 
capsules having an escape aperture sealed by three layers, 
including one that is mucoid, were found in the Conidae. 
Uncomplicated, single-layered capsules that may only 
have a suture in the wall to facilitate hatching appear 
to be characteristic of the Marginellidae. 

Muricacea and Buccinacea are species-rich taxa hav- 
ing family or species-specific egg capsule morphologies 
with a final shape that results from a molding process in 
the ventral pedal gland (.Ankel, 1929; Gruber, 1982; Sul- 
livan and Maugcl, 1984). Micromorphological similari- 
ties suggest that oviducal mechanisms common to both 
superfamilies are u.sed to construct structural laminae 
that serve the same respective functions in these taxa 
(tables 1, 2). Some species may add additional nonstruc- 
tural layers in the ventral pedal gland (table 2, LO; Sul- 
livan and Maugel, 1984). 

Protection (sealing the fibrous wall) appears to be a 



function of the outer structural lamina, LI. In most 
species, LI is thin (5 ^m or less), usually dense and finely 
fibered, and reflects the final shape (corrugations and 
ribbing) imparted by the ventral pedal gland (Gruber, 
1982; Sullivan and Maugel, 1984). LI seals the surface 
of the w hole capsule including the escape aperture, but 
not the basal plate in contact with the adhesive. Since 
the contents of some prosobranch egg capsules are axenic 
but not bacteriostatic (Lord, 1986), the fine structure of 
this lamina could serve as a ph\sical barrier to invasion 
by microorganisms, especially in species with a vacuo- 
lated and fluid-filled L2. (The basal adhesive may also 
serve as a barrier.) In Urosalpinx cinerea, LI is thick 
with separated fibers, but the interstices are filled with 
dense mucus (Tamarin and Carriker, 1967). LI could 
also serve as a reactive substrate during the molding and 
hardening process in the ventral pedal gland which fixes 
the capsule in its final shape (see Gruber, 1982 and Sul- 
livan and Maugel, 1984). 

L2 comprises the internal skeleton of the capsule wall 
in all neogastropod taxa studied except Marginellidae, 
which has a single structural lamina (tables 1-.3). In some 
taxa, extensive cross-linkages between the protein fibers 
form a homogeneous L2. In others, there are dense ho- 
mogeneous zones on inner and outer surfaces gradually 
.separating into directionally oriented fibers forming most 
of the lamina. In ribs or other sculpture, these fibers are 
loo.sely packed. The homogeneous zones, in addition to 
skeletal support, could pro\ide another ph\sical barrier 
to invasion by microorganisms. 

In Muricacea and Buccinacea, L2 may have fibrous 
components with different axial orientation often de- 
scribed as separate layers (Fretter, 1941; Gruber, 1982). 
Resistance to delamination and cohesiveness suggest that 
these layers were constructed from the same capsular 
protein during a continuous process; thus, the\ should 
be described as parts of a single capsular lamina. If all 
similar fibrovis components anil contiguous homogeneous 



C. N. D'Asaro 1988 



Page 147 



zones from previously described egg capsules of these 
superfamilies are viewed in this manner (as organized 
in tables 1, 2), then more meaningful comparisons be- 
tween taxa can be made. 

L3 is distinct in most muricaceans and buccinaceans 
and appears to be a mucoid extension of the apical plug, 
a part of the hatching mechanism (tables 1, 2). In both 
species of Cantharus, L3 is not present in most of the 
capsule wall; however, the mucoid plug in the escape 
aperture tapers gradually into the wall in a homologous 
position. There is also mucoid material with chromo- 
tropic characteristics similar to the apical plug in a ho- 
mologous basal position suggesting that a thin layer of 
L3 mucopolysaccharide may be present throughout the 
wall but could not be identified with the histological 
technique applied. In the egg capsule of Eupleura cau- 
data etterae, which was described by Gruber (1982: fig. 
14), the singular mucoid plug extending as a layer into 
the capsule wall and surrounding the lumen can be de- 
scribed as L3. Lying between the outer structural layers 
and L3 in E. c. etterae, is another layer (Gruber s sixth 
layer). The position of this layer, in contact with the 
equivalent of L2 and surrounding the outer edge of the 
mucoid plug formed by L3, and its composition suggest 
that it is homologous with the first mucoid plug in Chi- 
coreus florifer dilectus (table 1). 

L4. the albumen retaining layer, is the primary lamina 
in muricaceans and buccinaceans completely surround- 
ing albuminous fluids and embryos (tables 1, 2). Often 
it is not reported in the literature, perhaps because it is 
4 nm or less in thickness and usually bound tightly to 
other components. A function of the layer could be to 
prevent nonrefractory albumen from mixing with re- 
fractory capsular proteins when the outer structural lam- 
inae are assembled by the ciliary mechanism Fretter 
(1941) described. 

Albumen in egg capsules of muricaceans and bucci- 
naceans is stratified to some degree at oviposition (tables 
1, 2). This was rather obvious in capsules of Chicoreus 
florifer dilectus, which had a core of dense albumen, 
free of embryos, that was structurally different from the 
surrounding, less dense material. More typically in other 
species, the core albumen contained the embryos. Dif- 
ferent layers of albumen in newly deposited capsules 
suggest that two or more kinds of albumen producing 
cells exist in the oviducts of the species studied, as Fretter 
(1941) has demonstrated for several neogastropods. 

Muricaceans and buccinaceans have structurally sim- 
ilar barriers closing the escape apertures, which are five 
to nine times as thick as the capsule wall (tables 1, 2). 
The increased thickness of the less refractory mucoid 
plugs possibly serves to prevent premature hatching of 
a capsule. In muricaceans, no fibers from the more re- 
fractory parts of L2 extend into the mucoid plugs. During 
sectioning, mucoid plugs in the escape aperture fractured 
across the width of the capsule wall, while the more 
refractors parts of the capsule delaminated lengthwise. 
Boundary laminae in the wall (LI and L4) appear to 
hold the mucoid plugs in position. 

Conid egg capsules are less complex than those of the 



Muricacea and Buccinacea and contain smaller structural 
fibers, usually embedded in a mucoid component. The 
capsule wall has two laminae (LI and L2) and an al- 
bumen retaining layer (L4). Rather than having ribbon- 
like fibers in the laminae (see Flower et ai, 1969, for a 
discussion of fiber ultrastructure), there are very fine, 
short, and folded or twisted fibers that overlap into a 
cross-hatched pattern. A singular mucoid plug lying be- 
tween the outer laminae, LI, and the albumen retaining 
layer, L4, and interdigitating extensively with the fibrous 
middle lamina seals the escape aperture. Extensive in- 
terdigitation would require simultaneous formation of 
L2 and the mucoid plug. There is no evidence in the 
conids studied of a lamina equivalent to L3 associated 
with the mucoid plug. These differences in micromor- 
phology suggest that important differences in oviducal 
structure and function exist between conids and muri- 
caceans and buccinaceans. 

Pustulate marginellid egg capsules are microstruc- 
turally the least complex of the taxa studied, and differ 
markedly from capsules of other neogastropods. Each is 
constructed of three homogeneous components fused al- 
most indistinguishably into the others to form a capsule 
in which the parts do not delaminate during sectioning. 
Resistance to delamination and a uniform response to 
toluidine blue suggest that only one protein was used to 
form the capsule wall. There is no complex escape ap- 
erture, but in many marginellids it is possible to identify 
a preformed suture at which the capsule will break dur- 
ing hatching. The suture is visible at one end and on the 
sides, which corresponds exactly to the position where 
the capsule wall breaks during hatching. These features 
also suggest that the structure and function of the mar- 
ginellid oviduct is different from that of the major su- 
perfamilies studied. 

Although this study does show that there are micro- 
morphological features common to the egg capsules of 
some prosobranch taxa, it does not provide clear evidence 
that can be used to explain exactly how capsules are 
formed in the oviducal glands. It can be inferred that 
L4, the albumen retaining layer, is deposited around the 
albumen and embryos to hold them in a central position 
\\ hile the more refractory parts of the capsule are formed. 
It can also be inferred that for most species the remaining 
parts of the process involve sequential deposition of struc- 
tural laminae, as Fretter (1941) described for Nassarius 
reticulatits, with L3 and formation of the innermost mu- 
coid plug being the second part of the process. The third 
part of assembly would involve formation of the main 
structural lamina, L2, and the outer mucoid plug. LI 
would be the last structural lamina added. Sculpturing, 
hardening, addition of nonstructural la\ ers, and attach- 
ment with an adhesi\e are functions of the ventral pedal 
gland (Sullivan and Maugel, 1984). 

ACKNOWLEDGEMENTS 

I wish to acknowledge the revisions to the manuscript 
suggested by a colleague. Dr. Paul V. Hamilton. Some 
of the specimens sectioned were provided by Mr. Larry 



Page 148 



THE NAUTILUS, \'ol. 102, No. 4 



Dilmore. This project was financed in part with funds 
from the U.S. Environmental Protection Agency (CR- 
811649) and the University of West Florida. 

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Ankel, W. E. 1929. Lber die Bildung der Eikapsel bei 
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Ankel, W. E. 1937. Der feinere Bau des Kokons der Purpur- 
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das Laichleban. Verliaiidlungen der Deutschen Zoolo- 
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Bandel, K. 1975. Embryonalgehause Karibischen Meso- und 
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Bandel, K. 1976a. Morphologic der Gelege und okologische 
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Bandel, K. 1982. Morphologic und Bildung der friihontoge- 
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Bayne, C. J. 1968 Histochemical studies on the egg capsules 
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D'Asaro, C.N. 1970. Egg capsules of prosobranch mollusks 
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Flower, N. E. 1973. The storage and structure of proteins 
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Flower, N. E., A. J. (;eddes, and K M Kudall 1969 I'ltra- 
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THE NAUTILUS 102(4):149-153, 1988 



Page 149 



Two New Species of Metula (Gastropoda: Buccinidae) with a 
Description of the Radula of the Genus 



Philippe Bouchet 

Museum National d'Histoire Naturelle 
55, rue Buffoii 
75005 Paris, France 



ABSTRACT 

Two species of Metula H. and A. Adams, 1853, lack a radula, 
and it is very small in a third species. The morphology of the 
teeth is significantly different from that in Pisaiiia, and the two 
genera are probably not closeK related Metula crosnicri new 
species, from 400-450 m off SW Madagascar, is a large, broad 
species, with \ery convex whorls and a deep suture. Metula 
africana new species, from the deep continental shelf off West 
Africa, is considered the descendant of the Mediterranean Plio- 
cene M. milraeformis (Brocchi, 1814). This lineage cannot be 
taken as evidence for Mediterranean-Indo-Pacific connections 
in the lower Pliocene as claimed by Grecchi (1978). 



INTRODUCTION 

The chequered taxonomical history of the generic name 
Metula H. and A. Adams, 1853, has recently been sta- 
bilized by Emerson (1986), who clarified the identity of 
its type species, Buccinum clathratitm Adams and Reeve, 
1850. Additional information was provided by Beu and 
Maxwell (1987). 

The familial position of the genus has been the subject 
of a controversy between Ponder (1968, 1973) and Cer- 
nohorsky (1971). Ponder (1968) described the anatomy 
of Ratifitsus Iredaie, 1919, and Iredalula Finlay, 1927; 
he concluded that their peculiar glandular mid-esoph- 
agus as well as other features of the anterior alimentary 
canal justified their inclusion in the family Colubrariidae, 
which he considered to be anatomically distinct from 
the Buccinidae. Cernohorsky (1971) regarded the pres- 
ence of a vestigial radula in Ratifusus and Iredalula to 
indicate placement in the Buccinidae, since the species 
of Colubrariidae have no radula at all, and he suggested 
a placement in the buccinid subfamily Pisaniinae. This 
view has been accepted by most subsequent authors deal- 
ing with Metula (Olsson and Bayer, 1972; Kilburn, 1975; 
Houbrick, 1984; Emerson, 1986), who apparently over- 
looked Ponder s (1973) refutation of Cernohorsky s point 
of view. Ponder (1973) confirmed that Ratifusus, Ire- 
dalula. and Colubraria shared the same anatomical char- 
acters that separate them from the Buccinidae, and com- 
mented on the radular differences: "It thus appears that 
some Colubrariidae have lost the radula and that it is 



relatively small or vestigial in the remainder. It is possible 
that the whole Metula-Ratifusus series discussed by Cer- 
nohorsky (1971) belongs in the Colubrariidae as they all 
have similar shell features" (Ponder, 1973:328). 

The family Colubrariidae is treated as a synonym of 
Buccininae by Ponder and Waren (1988), while Beu and 
Maxwell (1987) recognize a subfamily Pisaniinae, where 
they include Metula, Colubraria, and a number of other 
genera. 

The purpose of the present paper is to provide a name 
for the West African species that has been known in the 
recent literature as Metula clathrata Adams and Reeve, 
and to describe another new Metula from the upper 
continental slope in the Mozambique channel. Several 
additional Indo-Pacific species of Metula, now under 
study, can be distinguished only on the basis of their 
protoconch, which has already been emphasized by Al- 
tena (1949) as a taxonomical character. 



SYSTEMATICS 

The radula of a species of Metula is figured here for the 
first time (figure 1). It is very small for a buccinid (ribbon 
25 ;um wide; central tooth 6.5 ^m wide, lateral teeth 12 
^m wide) and very similar to the radulae of Ratifusus 
and Iredalula figured by Ponder (1968): the central tooth 
has a narrow arched basal plate with 3 long, slender, and 
equal cusps; the lateral teeth also have a narrow basal 
plate and 3 long slender cusps, the outermost one being 
longest. 

A radula has been looked for, but not found in Metula 
amosi Vanatta, 1913, and M. cumingi (Adams, 1853); 
several specimens were examined in each case by A. 
Waren (personal communication). I do not consider pres- 
ence or absence of this very reduced radula to be of 
generic importance. 

The radula (figure 2) of Pisania striata (Gmelin, 1791), 
tvpe species of Pisania, is 150 ^m wide; it differs in 
having a central tooth with a large square basal plate 
and 5 short and broad cusps, the outermost 2 being small- 
er; the lateral teeth are more strongly built, with 3 un- 
equal cusps. 

In view of the small variation of radular types in buc- 



Page 150 



THE NAUTILUS, Vol. 102, No. 4 



cinids, this difference is remarkable and probabK indi- 
cates that the two genera are not closeK- related. What- 
ever rank (subfamily or tribe) the Pisania group is given 
in Huccinidae, additional research is needed before the 
Colubraria group is considered a mere s>nonym of it. 

DESCRIPTIONS 

[For a diagnosis of the genus see Altena (1949) as An- 
temetula] 

Metula crosnieri new species 
(figures 3-5, 9) 

Description: Shell solid, fusiform, consisting of 2.5 pro- 
toconch and 6.2 teleoconch whorls. Spire high, body whorl 
comprising 64% of total shell height. Protoconch (figure 
9) with large nucleus of two smooth convex whorls 
abruptK demarcated from teleoconch. Teleoconch whorls 
convex, without sutural ramp, with deeply impressed 
suture. Sculpture of raised spiral cords and curved op- 
isthocline axial ribs; cords and ribs producing beaded 
intersections and pitted intervals. Eight spiral cords on 
spire whorls; 4 minor, intermediate cords on penultimate 
whorl; about 18 cords above periphery of body whorl, 
principal and secondary cords alternating rather regu- 
larly, and 33 cords below periphery, of which about 15 
are set close together in siphonal region. Axial ribs about 
equal in strength to spiral cords on spire whorls; weaker 
on penultimate whorl, with main sculpture being spiral 
on body whorl. In addition to axial ribs, several incre- 
mental scars are obvious, especially on body and pen- 
ultimate whorls. Aperture ovate; inner lip thin, smooth, 
adherent to body whorl, thicker in columellar region; 
outer lip bearing 12 very weak teeth that do not corre- 
spond with position of external spiral cords; two most 
apical teeth slightly stronger. Peristome thickened, form- 
ing broad varix, also covered by spiral cords. Siphonal 
canal long, broad, widely open, and recurved. Fasciole 
indistinct. 

Colour light tan, with a very indistinct darker spiral 
band at periphery of body whorl; incremental scars light- 
er; aperture white. 

Dimensions of the holotype: Height 51 3 mm, width 
19.4 mm; height of the aperture 24.5 mm, width 8 mm; 
height of the body whorl 33 mm. 

The largest paratype is 55.4 mm high. 

Type lf>cality: Mozambique channel, SW Madagascar, 
off Baie de Faiiemotra, 22°15'S, 43°05'E, 470-475 m. 

Type material: Holotype and paratype 1 (MNHN) from 
the type locality, collected by A. Crosnier, Dec. 2, 1973 
aboard R.V. "N'auban"; paratype 2 (MNHN), Mozam- 
bique channel, SW Madagascar, 22°17'S, 43°04'E, 400- 
450 m, collected by R. v. Cosel, Nov. 30, 1986 aboard 
trawler "Mascareignes III". 

Distribution: Known only from llic type material, off 
SW Madagascar. 




Figures 1, 2. Scanning electron niicrographi ol raciulac I. 
Metula africana new species, scale bar lO/iiii. 2. Fisania striata 
((Jnielin. 1791), scale bar .50 nm. 

Remarks: Metula crosnieri has remarkably convex 
whorls and a deep suture when compared with its con- 
geners. The protoconch (figure 9) indicates non-planc- 
totrophic larval development. The combination of these 
two characters distinguishes it from all known Indo-Pa- 
cific Metula. 

In the Atlantic, Bartschia significans Rehder, 1943, 
type species of Bartschia Rehder, 1943, has even more 
convex whorls, and a multispiral protoconch. I cannot 
find characters other than the convexit\- of the whorls 
that sharply distinguish Bartschia from Metula, and con- 
clude that Bartschia should be considered at most a sub- 
genus of Metula. 



p. Bouchet 1988 



Page 151 



I am naming this species after my colleague Alain 
Crosnier, who first collected it during a survey of deep 
water shrimp populations oft Madagascar. 

Metula ajricana new species 
(figures 1, 6-8, 10) 

Metula clathrata Knudsen, 1956;39, plate 1, figure 1; non M. 

clathrata (Adams and Reeve, 1850). 
iKin Metula knudseni Kilburn, 1975:592 (replacement name 

for Buccinum clathratum Adams and Reeve, 1850: see 

Emerson, 1986). 

Description: Shell solid, fusiform, consisting of 7 teleo- 
conch whorls. (Protoconch of holotype partly broken, on 
a paratype consisting of large nucleus and 1.5 smooth 
convex whorls, figure 10.) Teleoconch whorls convex, 
with faint but distinct sutural ramp on early whorls; ramp 
indistinct on penultimate whorl, body whorl evenly con- 
vex. Teleoconch sculpture of raised spiral cords and 
slightly curved opisthocline axial ribs; intersections dis- 
tinctly beaded in sutural ramp area, only slightly nod- 
ulous below ramp. Spiral and axial sculpture of similar 
strength on early whorls; later spiral sculpture gradually 
dominates. Nine primary spiral cords per whorl on spire 
whorls, first (adapical) and third cords stronger, limiting 
sutural ramp. Fine spiral threads present between cords 
on penultimate and bod\ whorl, several eventually de- 
veloping into secondary spiral cords, with one present in 
sutural ramp area of body whorl. About 22 primary cords 
below periphery of body whorl, plus another 10 in si- 
phonal area. Incremental scars distinct; growth lines very 
distinct between axial ribs. Aperture ovate, narrow. Inner 
lip thin, smooth, adherent to body whorl, thicker in col- 
umellar region. Outer lip regularly convex, bearing small 
teeth that correspond at least partly with position of spiral 
cords on peristome; a group of 5 teeth forms a small 
callus in apical portion of outer lip, delimiting small anal 
canal. Peristome forming a thickened varix, over which 
spiral sculpture extends. Siphonal canal short, narrow, 
open, only slightly recurved. 

Ground colour of shell beige cream, with 3 brown 
spiral bands; 3 adapical cords brown, interval between 
them beige cream. Below, uniformly beige band extends 
over next 2 spiral cords, occupies central position on spire 
whorls. Darkest band occupies next 4 spiral cords and 
intervals between them; this band occupying suprasu- 
tural position on spire whorls, and a central position on 
body whorl. Below brown band a second beige band 
extends over 4 spiral cords; next 2 cords brown, with 
brown colour fading towards base of shell. Aperture cream 
colored. 

Dimensions of the shell: Height 54.5 mm, width 17.5 
mm; height of the aperture 27 mm, width 7 mm; height 
of the body whorl 35 mm. 

Type locality: Off Saint Louis, Senegal, in 300-600 m. 

Type material: Holotype (MNHN), paratype 1 (MNHN) 
and paratype 2 (AMNH 198755), all from the type lo- 
cality, collected by M. Pin on the trawler "Louis Sauger". 





Figures 3-8. Back, side, and front views. 3-5. Metula cros- 
nieri new species, holotype, 51.3 mm. 6-8. Metula ajricana 
new species, holotype, 54.5 mm. 



Other material examined: SENEGAL: off Saint Louis, 
"deep water", 4 shells (MNHN, leg. M. Pin), 10 shells 
(coll. M. Pin, Dakar); IVORY COAST: off Abidjan, 1 
shell, coll. Marche-Marchad (MNHN), and 1 shell, P. Le 
Loeuff coll. (MNHN); EQUATORIAL GUINEA: Atlan- 
tide Sta. 120, 02°09'N, 09°27'E, 250-850 m, 1 specimen 



Page 152 



THE NAUTILUS, Vol. 102, No. 4 




Figures 9. 10. Protoconchs. 9. Metula crosnieri. 10. Metula 
ajricana. Scale line 1 mm. 



(Kiiudsen, 1956) (ZMC); CONGO: West of Pointe-Noire, 
100 m, 1 shell, A. Crosnier coil. 1962 (ANSP 333810). 

Distribution: Deep continental shelf and upper slope 
of West Africa, from Senegal to Congo. 

Remarks: Metula africana has been figured three times 
in the literature: by Knudsen (1956: plate 1, figure 1) 
under the name Metula clathrata; by Emerson (1986: 
figures 4, 5) as Metula sp.; finally by Kaicher (1987: card 
4851) as Metula sp. I refer to Emerson (1986) who re- 
viewed the nomenclature of this West African Metula 
and concluded that it represents a new species. 

The protoconch (figure 9) indicates non-planctotro- 
phic larval development. 

There may be clinal variation in adult size along the 
West African coast: The 18 shells examined from Senegal 
have a mean height of 41.7 mm; the 2 shells from Ivory 
Coast, although fully adult, measure 32 and 36 mm; that 
from Equatorial Guinea is 30.6 mm high, and that from 
Congo 25.5 mm. 

The tiny rachiglossate radula (figure 1) was prepared 
from the specimen taken during the .-Ktlantide expedition 
(ZMC). 

The close connection between Mediterranean Pliocene 
and Recent West African marine faunas has been dem- 
onstrated in a number of paleontological papers (for re- 
cent reviews see Ruggieri, 1967; Marasti and Raffi, 1979; 
Sahelli and Taviani, 1984). With a single species of Me- 
tula present in the Pliocene of Italy and a single Recent 
species in West Africa, it is reasonable to assume that 
Metula mitraeformis (Brocchi, 1814) (for figures and 
references see Pelosio, 1966) is the direct ancestor to M. 
africana. I have examined material of the fossil species 
and found it to differ from the Recent one by its much 
weaker axial sculpture, which on the body and penul- 
timate whorls is limited to growth lines. In M mitrae- 
formis, there is a very broad sutural ramp that extends 
over the adapical third or half of early teleoconch whorls, 
and becomes obsolete on the penultimate and body whorls; 
the siphonal canal is also broader. 

Because he was mistaken about the identity, type lo- 



calit\, and distribution of M. clathrata, Grecchi (1978) 
speculated that the presence of its presumed ancestor M. 
mitraeformis in the Mediterranean Pliocene was an in- 
dication of Mediterranean-Indo-Pacihc connections in 
the lower Pliocene after the Messinian salinity crisis. 
With M. clathrata now known to be a West American 
species distinct from M. africana, the history of the M. 
mitraeformis-africana lineage can not be taken as an 
indication of such connections. This lineage probably has 
an Eastern .Atlantic history dating to the Miocene; al- 
though the Neogene West African fossil record is lacking, 
it is far more probable that M. mitraeformis reinvaded 
the Pliocene Mediterranean from West Africa rather than 
from the Indo-Pacifie through unproven maritime con- 
nections. 

ACKNOWLEDGEMENTS 

I thank Dr. A. Waren who prepared the radula of Metula 
africana and commented on the manuscript. A. Crosnier, 
M. Pin, and R. von Cosel collected most of the material 
cited in this paper. Dr. W. Emerson generousK refrained 
from describing M. africana when he learned 1 was 
working on it. Photography is b\ P. Lozouet. 

LITERATURE CITED 

Alteiia, C O van R 1949. The genus Antemetula Rehder 
in the Indo-West Pacific area, with the description of two 
new fossil species. Bijdragen tot de Dierkdunde 28:385- 
393. 

Beu, A. G. and P. A. Maxwell. 1987. A revision of the fossil 
and living gastropods related to Plesiotriton Fischer, 1884 
(Family Cancellariidae, Subfamily Plesiotritoninae n. 
subfam.) with an appendix: genera of Buccinidae Pisani- 
inae related to Colubraria Schumacher. 1817. New Zea- 
land Geological Survey Paleontological Bulletin 54:1-140, 

Cernohorsky, W. 1971. Indo-Pacific Pisaniinac and related 
buccinid genera. Records of the .\uckland Institute and 
Museum 8:137-167. 

Emerson, W. K. 1986. On the type species of Metula H. & 
\. Adams, 1853: Buccinum clathratum A. .\dams and 
Reeve, 1850. The Nautilus 100(1 ):27-:30 

Grecchi, G. 1978. Problems connected with the recorded 
occurrence of some mollusks of Indo-Pacific affinit\ in the 
Pliocene of the Mediterranean area Rivisla Italiana di 
Paleonlologia 84:797-812. 

Houbrick, R. 1984. A new "Metula" species from the Indo- 
West Pacific. Proceedings of the Biological Society of 
Washington 97(2):420-424. 

Kaicher, S. D. 1987. Card catalogue of world-wide shells. 
Pack 48: Buccinidae, part 3 Published by the author, St. 
Petersburg. 

Killnirn, R N 1975. Taxonomic notes on South African ma- 
rine Mollusca (5): including descriptions of new ta\a of 
Rissoidae, Cerithiidae, Tonnidae, Cla.ssididae, Buccinidae, 
p-asciolariidae, Turbinellidae, Turridae, Architectonici- 
dae, Epitoniidae, Limidae, Thraciidae. Annals of the Natal 
Museum 22(2):577-622. 

Knudsen, J. 1956. Marine prosobranchs of tropical West Af- 
rica (Stenoglossa). Atlantide Report 4:7-110. 

Marasti, R and S. Raffi 1979 Observations on the paleocli- 
matic and biogeographic meaning of the Mediterranean 



p. Bouchet 1988 



Page 153 



Pliocene molluscs. State of the problem. 7th Intenuilinnal 
Congress on Mediterranean Neogene, Athens, 8 unnum- 
bered pages. 

Olsson, A. and F. Bayer. 1972. American Metulas (Gastro- 
poda: Buccinidae). Bulletin of Marine Science 22(4):900- 
925. 

Pelosio, G. 1966. La Malacofauna dello stratotipo del Tabi- 
aniano (Pliocene inferiore) di Tabiano Bagni (Parma). Bol- 
lettino della Societa Paleontologica Italiana 5(2):l()l-183. 

Ponder, W. 1968. Anatomical notes on two species of the 
Colubrariidae. Transactions of the Roval Society of New- 
Zealand, Zoology 10(24):217-223. 

Ponder, W. 1973. The origin and evolution ot the Neogas- 
tropoda. Malacologia 12(2):295-338. 



Ponder, W. and A Waren 1988 Classification of the Cae- 
nogastropoda and I leteroslropha — a li.sl of the family-group 
names and higher taxa. Malacological Review, supplement 
4:286-32-1. 

Ruggieri, G. 1967. The Miocene and later evolution of the 
Mediterranean Sea. In: Adams, C. G. and D. V. Ager (eds.). 
Aspects of Tethyan biogeography. The Systematics Asso- 
ciation, London. P. 283-290. 

Sabelli, B. and M. Taviani. 1984. The paleobiogeographic 
distribution of the Mediterranean benthic mollusks and 
the Messinian salinity crisis or where did the mollusks go? 
Annales de Geologic des Pays Helleniques 32:263-269. 



THE NAUTILUS 102(4):154-158, 1988 



Page 154 



A New Species of Intertidal Terebra from Brazil 



Kurl Auffenberg 

Malacolog> Division 
Florida Museum of Natural History 
University of Florida 
Gainesville, FL 32611. USA 



Harry G. Lee 

709 Uomax Street 
Jacksonville, FL 32204, USA 



ABSTRACT 

Terebra imitatrix new species is described from northern Brazil 
and compared to morphologically similar species of Hastnla. 
The three known terebrid feeding types are briefl\ reviewed. 
Dissection of the foregut of this new species revealed characters 
that overlap two of the major feeding types. 

Key words: Gastropoda; Terebridae; Terebra; anatom\ : Bra- 



INTRODUCTION 

While compiling distributional records of the West At- 
lantic Hastula cinerea species group, the junior author 
located an unidentified lot of 23 specimens from Brazil 
in the Academy of Natural Sciences of Philadelphia 
(ANSP 299957). These specimens were compared to the 
known terebrids from the Atlantic and were found to 
belong to a distinct undescribed species. An additional 
lot of the same species was subsequently found in the 
American Museum of Natural History (AMNH 129280). 
Matthews et al. (1975:99, fig. 31) in their treatment of 
Hastula ciiterea describe and illustrate a protoconch con- 
sistent with this new species. The Brazilian specimens of 
Hastula salleana (Deshayes, 1859) figured by Rios (1970: 
123, pi. 47, 1975:127, pi. 38, fig. 560, 1985:131, pi. 45) 
are probably referable to this new species, but the figured 
specimen was unavailable for examination. This paper 
describes this new species and compares it with mor- 
phologically similar species of Hastula. A description of 
the foregut anatomy reveals that it does not conform to 
any of the three known feeding types, which are re- 
viewed herein. 

MATERIALS AND METHODS 

Only shells possessing 10 or more teleoconch whorls and 
with intact protoconchs and apertures were measured 
with Vernier calipers. All dissections were made under 
a Wild M-5 dissecting micro.scope and line drawings 
made with the aid of a camera lucida. Description and 
discussion of the anatomy is limited to the foregut due 
to poor preservation in the upper whorls. Anatomical 
and protoconch measurements were made with an ocular 



micrometer and converted to millimeters. Terminology 
follows that of Miller (1970, 1971). Two preserved but 
completeK retracted adult specimens of the new species 
(ANSP 299957), Hastula maryleeae Burch (UF 113539) 
and Hastula salleana (Deshayes) (UF 48197, 113540), 
were dissected from each lot. Two dried-in specimens of 
Hastula maryleeae Burch (T. Bratcher collection) were 
rehydrated in a weak solution of potassium h\dro.\ide, 
transferred into water, and dissected. The radular sac 
was extracted and dissolved in a weak solution of potas- 
sium hydroxide. Radular teeth were individual!} mount- 
ed on scanning electron microscope specimen stubs cov- 
ered with double-sided tape. Micrographs were made 
with a Hitachi 5-415.^ scanning electron microscope. 

Repositories of examined specimens are indicated by 
the following abbreviations: 

AMNH American Museum of Natural Histor\' 
ANSP Academy of Natural Sciences of Philadelphia 
UF Florida Museum of Natural History 



SYSTEMATICS 

Family Terebridae Morch, 1852 
Genus Terebra Bruguiere, 1789 

Terebra imitatrix new species 
(figures 1-6, 8, 9, table 1) 

Description: Shell (figures 1-3, 6) medium in size, 
broadening anteriorK ; color variable, ranging from 
banded, or cream, to purplish-brown; teleoconch whorls 
10-12; sides flat to slightly convex. Protoconch whorls 
1.5-2.0; glassy, transparent (figure 6). .\xial sculpture of 
close-set, recurved ribs of \ariable strength, generally 
distinct near the suture, becoming obsolete anteriorly; 
major axial ribs 34-57 (x = 41.6) on the penultimate 
whorl. Spiral sculpture of very faint microscopic incised 
lines most distinct in the intercostal spaces on the upper 
whorls, obsolete on later whorls, rarely cro.ssing the axial 
ribs; spiral rows of pits absent. Last whorl with obsolete 
axial ribs and spiral incised lines; color pattern variable, 
typicalK of five diffuse color bands: one white pre-sutural 
band usiialK w ith distinct brown spots that become ob- 
solete toward apertural lip, one broad bluish-black zone 
at shoulder, one pinkish band at periphery, one purplish- 



K. Auffeiiberg and H. G. Lee 1988 



Page 155 




4 




Figures 1-3. Terebra imilatrix new species. 1. Holotype, ANSP 299957 (27.5 mm shell length) 2, 3. Paratypes, ANSP 369293, 
all from Rio Grande do Norte, Brazil, -sand island at mouth of the inlet at Areia Branca. 



brown band below periphery, and one white basal zone. 
Columella brown, centrally concave and slightly re- 
curved; rounded rib present on anterior edge. Parietal 
callus thin, transparent to light brown. Fasciole white to 
bluish-gray with distinct white rib. Anterior siphonal 
notch moderately broad, straight. Aperture light to dark 
brown with white band. 

External anatomy: Animal cream-colored with no ap- 
parent pattern in alcohol-preserved specimens. Oper- 
culum corneous, small, thin. Eyes on short, broad eye- 
stalks. Labial tube large, spoon-shaped (figure 4). Anterior 
end of labial tube terminating in thick muscular lips 
bounding mouth slit; well-developed sphincter lacking. 

Foregut anatomy: Labial cavity large, dominated by 
massive (3.7 mm), extendable, muscular organ (accessory 
feeding apparatus; Miller, 1970, 1971) that tapers an- 
teriorly; muscular organ with two rows of papillae on 
ventral side (figure 4); attached posteriorly to the left 
side of the cephalic hemocoel by connective tissue; re- 
tractor muscle large, originating in foot below and slight- 
ly posterior to anterior siphon, passing through cephalic 
hemocoel. Buccal tube muscular, short (0.9 mm), taper- 
ing anteriorly. Buccal cavity small (0.6 mm), rounded. 
Pre-ganglionic esophagus enters buccal mass posteriorly 
(figure 4). Salivary glands not located. Radular sac blade- 
shaped, small (0.8 mm). Radular caecum (0.4 mm) with 
well-developed groove running posteriorly, two distinct 



bulbs anteriorly (figure 5). Radular organs attached to 
right side of anterior portion of buccal cavity by very 
short duct (0.1 mm). Two rows of radular teeth situated 
obliquely in radular sac and caecum (figure 5). Radular 
teeth about 30; 0.1-0.2 mm in length, slightly curved, 
not barbed (figures 8, 9). Poison gland long (6.2 mm), 
extremely convoluted, entering right side of buccal cav- 
ity slightly behind radular sac (figure 5). Poison bulb 
small (0.9 mm), weak, seemingly vestigial, lying at ven- 
tro-posterior end of cephalic hemocoel. 

Etymology: From the Latin feminine noun meaning 
one who imitates, in reference to the deceptively close 
resemblance of the shell to that of Hastula cinerea (Born, 
1778), with which it occurred in the type lot. 

Table I. Measurements (mm) of shell characters of Terebra 
imitatrix new species and Hasttila rnaryleeae Burch 



Species 


Character 


X 


Range 


SD 


T. imitatrix 


Shell length 


27.3 


24.1-29.7 


1.9 




Shell width 


6.4 


5.7-7.4 


0.5 




Protoconch 


0.50 


0.46-0.52 


0.03 




width 








H. rnaryleeae 


Shell length 


18.2 


15.0-24.1 


2.7 




Shell width 


4.6 


3.7-5.5 


0.6 




Protoconch 


40 


0.38-0.42 


02 




width 









Page 156 



THE NAUTILUS, Vol. 102, No. 4 




•■s be 






Figures 4-6. Terebra irnitatrix new species. 4. Diagrammatic dorsal view nf the organs of the foregut 5. Right side of the buccal 
tulie showing radular organs and insertion of the poison gland. 6. Protoconch. 7. Hastula manjleeae Burch, protoconch (.-WINH 
191815) Scale bars = 1 mm. af, accessory feeding apparatus; be, buccal cavity; bt, buccal tube; It, labial tube; pb, poison bulb; pe, 
pre-ganglionic esophagus; pg, poison gland; re, radular caecum; rm, retractor muscle; rs, radular sac. 



Type locality: Brazil, Rio Grande do Norte, sand island 
at mouth of the inlet at Areia Branca, 04°57'S, 37°08"W, 
G. & M. Kline et al., 14 December 1963. 

Holotype: ANSP 299957, shell length 27.5 mm, width 
6.7 mm. 

Paratypes: Paratypes 1-20, ANSP 369293, from the 
tvpc locality (15 dry, 5 in alcohol). Paratypes 21-22, UF 
115180, from type locality. Paratypes 23-24, AMNH 
129280, Brazil, Ceara, Acaraii, 02°53'S, 40°07'W. 

Distribution: PresentK know n onl\- from northern Bra- 
zil. 

Ecology: Based on the locality label of the type lot, this 
species is found in sand near or at inlets much like some 
populations of Hastula. One specimen of Hastula ci- 
nerea (Born, 1778) was found in the type lot and another 
(AMNH 129269) from the same locality as the AMNH- 



paratypes. Although these two species may occur micro- 
s\ mpatrically, we assume there is no trophic competition 
due to the strikingly divergent feeding organs (see dis- 
cussion). 

Comparative remarks: Terelrra irnitatrix is similar in 
shell morpholog\- to the West Atlantic Hastula cincrea 
group. It can be easily separated from Hastula cinerea 
and Hastula salleana by the lack of spiral rows of pits 
and the generally more numerous, but less prominent, 
axial ribs. .Although similar in size, T. irnitatrix is usually 
broader anteriorly . Hastula cinerea and H. salleana have 
3.5-4.0 protoconch whorls, while T. irnitatrix has 1.5- 
2.0. The typical color pattern of T. irnitatrix is more 
distinctK banded with larger and more distinct brown 
spots al the suture and a broader white subsutural band. 
Terelna irnitatrix and Hastula manjleeae are more dif- 
ficult to separate, particularly southern Caribbean pop- 



K. Auffenberg and H. G. Lee 1988 



Page 157 



Illations of H. niarijleeae described as Terehra toba- 
goensis Nowell-l'slicke, 1969, now placed in the 
s\non\ my of H. manjleeae by Bratcher and Cernohorsky 
(1987). Examination of the lectotype (AMNH 195453 
designated bv Bratcher and Cernohorsky, 1987:194, pi. 
60, fig. 235c)' and paralectotypes (AMNH 191819) of T. 
tobagoensis. as well as several other lots from Tobago, 
typical specimens of H. manjleeae from Te.xas, and spec- 
imens of a weakly ribbed form of H. maryleeae from 
the Dominican Republic (see Bratcher and Cernohorsky, 
1987, for discussion) revealed consistent shell characters 
b\ which the two species may be separated. Anatomical 
features are discussed below. Typical H. maryleeae is 
easily separated from T. imitatrix by the distinctive shell 
shape caused by the enlarged nodes of the axial ribs at 
the suture and by the slight crenulations at the sutures 
of the upper whorls. Hastula manjleeae has a much 
smaller shell than T. imitatrix at the same whorl count 
(table 1). The shell of T. imitatrix broadens more an- 
teriorK- and has a proportionately slightly larger aper- 
ture. The protoconchs of both species have 1.5-2.0 whorls, 
however, the protoconch of T. imitatrix is more bulbous 
(figures 6, 7; table 1). Typical T. imitatrix also resembles 
the West African Hastula aciculina (Lamarck, 1822), 
particularK- in color pattern. However, T. imitatrix lacks 
the supra-sutural groove and callosity found in H. acic- 
ulina (Bouchet, 1982; Bratcher and Cernohorsky, 1987). 



DISCUSSION 

Miller (1970) proposed a division of the Terebridae into 
three major groups based on the anatomy of the foregut 
(feeding type) and later published a series of papers 
(Miller, 1971, 1975, 1979) on this subject supported by 
in-depth life history studies. These data are reviewed 
and the species assigned to each group are listed in 
Bratcher and Cernohorsky (1987). Type I species have 
a long labial tube and a short buccal tube. They do not 
possess a radular apparatus or poison organs. This group 
is further divided into two subgroups, lA and IB, based 
primarily on habitat and pre%' (see Bratcher and Cer- 
nohorsky, 1987). Type II encompasses species exhibiting 
typical toxoglossan feeding characters. The labial tube 
is long and eversible and the buccal cavit\ is relatively 
large. The buccal tube is long and retractile. They have 
a poison gland and bulb as well as a radular sac containing 
two rows of harpoon-like radular teeth. This feeding type 
is further divided into two subgroups, types IIA and IIB, 
based primarily on habitat and behavior. Type IIA in- 
cludes several Indo-Pacific species, as well as the West 
Atlantic Hastula discussed in this paper (see Bratcher 
and Cernohorsky, 1987). Type III species possess an ac- 
cessory feeding organ which grasps prey and pulls it into 
the labial cavity. The\' lack a radular apparatus and some 
have lost the buccal tube and salivary glands. This group 
presentK contains no .\tlantic species, but is represented 
b\' several Indo-Pacific taxa (see Bratcher and Cerno- 
horsky, 1987). 

Terebra imitatrix has very different foregut anatomy 




Figures 8, 9. Radular teeth of paratype (ANSP 369293) of 
Terehra imitatrix. new species. 8. Whole tooth, 60 x . 9. Tip 

of lootli, 200 X. 



from the West Atlantic Hastula. We have dissected H. 
salleana and H. manjleeae for comparison, and both 
possess a Type IIA polyembolic proboscis (Miller, 1970, 
1971). Terebra imitatrix has a large spoon-shaped labial 
tube, presumably ineversible, a labial cavity dominated 
by the accessory feeding apparatus, an extremely short 
buccal tube incapable of extending outside the mouth 
and the buccal cavity, and the associated radular organs 
are minute in comparison to those of Hastula. The ves- 
tigial poison gland and bulb are barely recognizable as 
such and are considered homologous to the massive poi- 
son gland and bulb of Hastula only by the similarity of 
position and the site of its entrance into the buccal cavity. 
The radular teeth are similar in size and shape to those 
of Hastula maryleeae (0.15 mm). The teeth of H. sal- 
leana and H. cinerea are larger (0.5 mm) and barbed. 
The presence of an accessory feeding apparatus and the 
small size of other foregut organs place T. imitatrix near 
the group possessing a Type III polyembolic proboscis. 
However, the presence of radular and poison organs, such 
as occur in T. imitatrix, has not been reported for this 
proboscis type. Also, the shell of T. imitatrix is very 
different from the species in this group, all of which have 
shells with deeply impressed sutures, a sub-sutural groove 
and strong to moderate axial sculpture. 

The feeding behavior of Hastula cinerea and of H. 
inconstans (Hinds, 1844) have been well documented 
by Marcus and Marcus (1960) and Miller (1979), re- 
spectively. The long eversible labial tube forages for prey 
items. A single radular tooth is passed through the labial 
tube, held in the tip, and inserted into the prey to fa- 
cilitate penetration of the venom. The prey is then in- 
gested via the labial tube. Miller (1970) suggests that 
species with a Type III proboscis forage by utilizing the 
accessory feeding apparatus, and that food items are 
passed into the opening of the interior buccal tube. We 
do not know if the radular and poison organs are func- 
tional in Terebra imitatrix. If so, the radular tooth is 
either transferred from the buccal tube to the accessory 
feeding apparatus and inserted into prey outside the 
body, or once it reaches the buccal tube. 



Page 158 



THE NAUTILUS, Vol. 102, No. 4 



The presence in Terebra imitatrix of the accessory 
feeding apparatus recorded in a few Indo-Pacific Tere- 
bra and the poison anil radiilar organs of Hastula make 
this species unique. Terelna imitatrix may be closely 
related to the West African species, Hastula aciculina, 
however, this species must be studied anatomically be- 
fore this assignment can be verified Generic limits within 
the Terebridae are presently p<}ori\ understood with much 
overlap in shell and anatomical characters. Additional 
anatomical studies are needed to clarify the taxonomic 
and e\oiutionar\ significance of the foregut in the Tere- 
bridae. 



ACKNOWLEDGEMENTS 

We wish to thank several people who have assisted us 
during this study. Special thanks go to Robert Robertson, 
George M. Davis, and Mar) \. (iarback of the .\cademy 
of Natural Sciences of Philadelphia for allowing us to 
borrow and dissect the specimens on w hich this descrip- 
tion is based. William K. Emerson, Harold Feinberg, and 
Walter Sage at the .American Museum of Natural Histor\ 
allowed us to visit their department and borrow several 
lots of \ery pertinent material. Twila Bratcher of Hol- 
lywood, California kindly provided comparative mate- 
rial, as did Eliezar C. Rios (Funda^ao Universidade do 
Rio Grande do Sul, Brazil), the late Joseph Rosewater of 
the U.S. National Museum, and Thomas Pulley and Con- 
stance Boone of the Houston Museum of Natural Science. 
Marcos Guianaldo and Armando Garcia, Florida Mu- 
seum of Natural History, prepared figures 4-7. The mi- 
crographs were made by the senior author with the scan- 
ning electron microscope housed in the Department of 
Zoology, University of Florida. In addition, we thank 
Richard S. Houbrick, William K. Emerson, and Fred G. 
Thompson for reviewing earlier drafts of the manuscript. 



LITERATURE CITED 

Bouchet. P. 1982. Les Terebridae (Mollusca, Gastropoda) de 
r.\tlantique Oriental. Bollentino Malacologico, Milano 
18(9-12):18.5-2I6. 

Bratcher, T. and \V. O. Cernohorsky 1987 Living terebras 
of the world .\ monograph of the Recent Terebridae of 
the world .Xmerican Malacologists, Inc.. Melbourne, FL, 
240 p. 

Marcus, E. and E. Marcus 1960. On Hastula cinerea. Bo- 
leiitim Faculdade de Filosofia Ciencias e Letras. Univer- 
sidade de Sao Paulo, no. 260, Zool. no. 23:25-54, pis. 1-5. 

Matthews. H. R., \. C. Dos Santos Coelho, P. De Sa Cardoso, 
and M. Kempf. 1975. Notas sobre la familia Terebridae 
no Brasil (Mollusca, Gastropoda). .-Vrquivos do Museu Na- 
cional. Rio de Janiero 5585-104. 

Miller. B .A 1970 Studies on the biologN of Indo-Pacific 
Terebridae. Ph D. dissertation. I'niversity of New Hamp- 
sliirc. Durham. 213 p. 

Miller, B. .\. 1971. Feeding mechanisms in the family Tere- 
bridae. .Annual Report of the American Malacological 
I'nion for 1970. 1970:72-74. 

Miller. B. .A. 1975. The biolog\ of Tereiwa gouWi Deshayes, 
1859, and a discussion of the life history similarities among 
other terebrids of similar proboscis tvpe Pacific Science 
29:227-241. 

Miller, B. .\. 1979. The biology of Hastula inconstans (Hinds, 
1844) and a discussion of life histor\ similarities among 
other Hastula of similar proboscis type. Pacific Science 
33(3):289-306. 

Nowell-l'sticke, G. W. 1969 .A supplementary listing of new 
shells (illustrated). Privately published. St. Croix, Virgin 
Islands, 31 p., 4 pis. 

Rios, E. C. 1970, Coastal Brazilian shells Funda(,ao Cidade 
do Rio Grande, Museu Oceanografico de Rio Grande, Bra- 
zil, 255 p.. 60 pis. 

Rios, E. C. 1975. Brazilian marine mollusks iconograph\. 
Fundagao Universidade do Rio Grande, Centro de Cien- 
cias do Mar, Museu Oceanografico, Brazil, 331 p., 91 pis. 

Rios, E. C. 1985. Seashells of Brazil Funda^ao Cidade do 
Rio Grande, Fundagao Universidade do Rio Grande, Mu- 
seu Oceanographico, Brazil, 329 p , 1421 pis. 



THE NAUTILUS 102(4):159-163, 1988 



Page 159 



Notes on the Biology and Morphology of Margaritifera hembeli 
(Conrad, 1838) (Unionacea: Margaritiferidae) 



Douglas G. Smith 

Museum of Zoology 
Universit\ of Massachusetts 
Amherst, MA 01003-0027, USA 



ABSTRACT 

The freshwater pearl mussel Margaritifera hembeli is found 
onK in the Red River basin and a few nearby drainages in 
Louisiana. Though of concern to conservationists because of its 
declining numbers, M. hembeli remains virtually unknown with 
respect to its anatomy and biology. The species contains all the 
anatomical characters that typify margaritiferid species. The 
sexes are separate and the gonads show a definite seasonality 
in activity. Gametogenesis is pronounced in specimens collected 
in the fall, followed by degeneration of reproductive tissues in 
the late winter through to late spring. It is concluded, on the 
basis of observed gonadal activity, that oviposition and spawn- 
ing take place between late November and late January Char- 
acters are evident in the morphology of the visceral nervous 
svstem and the stomach of M. hembeli that clearly distinguish 
M. hembeli from M. marrianae and other eastern North Amer- 
ican margaritiferid species. A distinct relationship between M. 
hembeli and M. marrianae, however, is suggested by the mu- 
tual occurrence of lateral hinge teeth and a corrugated surface 
of the posterior portion of the shell. Due to the lack of knowl- 
edge of the anatomy and biology of other margaritiferid species, 
especially those living in Asia, it is premature to suggest rela- 
tionships lietween M. hembeli and other described margari- 
tiferid species, particularly those with lateral hinge teeth. 

Key uords: Margaritifera: anatomv, reproduction. North 
.America 



INTRODUCTION 

The North American freshwater mussel Margaritifera 
hembeli (Conrad, 1838) was once believed to comprise 
two geographically discontinuous populations ta.xonom- 
ically linked b\ vague similarities of the shell. Johnson 
(1983) separated the two populations taxonomically by 
describing the Alabama group as M. marrianae, thus 
restricting the M. hembeli group to Louisiana. His de- 
scription included characters of the shell only, principally 
the degree of sculpturing on the shell surface and the 
shape of the ventral shell margin. Additional concho- 
logical differences between the Louisiana and Alabama 
populations were noted by Smith (1983) who pointed out 
dissimilarities in the mantle-shell attachment scars on the 
inner nacreous surface. 

Margaritifera hembeli probably had a more extensive 



range in the Red River drainage as indicated by museum 
records, particularly a specimen in the American Mu- 
seum of Natural History (AMNH 193786) from the Red 
River in Arkansas. During the present century, however, 
the range has contracted considerably due to deterio- 
rating environmental conditions. The present range is 
limited to the Bayou Teche drainage (Vidrine, 1985) and 
a single stream in the Red River drainage. The drastic 
reduction in range has elicited concern from the federal 
government, which has provided protection for the re- 
maining populations (Stewart, 1988). 

Despite increased concern for M. hembeli, very little 
is known about this species other than the characteristics 
of its shell. Ortmann (1912) provided a brief description 
of the anatomy of M. "hembeli, ' but the specimens upon 
which he based his description came from Alabama and, 
therefore, are appropriately referred to M. marrianae. 
Hence the anatomy of M. hembeli remains undescribed, 
and nothing is known about its biology. The present study 
provides some information on gonadal activity and de- 
tails of the anatomy of the stomach and nervous system. 
Comparisons are made with other North American mar- 
garitiferid species, including M. marrianae, as studied 
by Smith (1979a, 1980, 1986, and unpublished). 

MATERIALS AND METHODS 

A total of 43 partially or completely relaxed, preserved, 
specimens were studied. All were collected on various 
dates from 1973 to 1986 from Brown Creek, Gardner, 
Rapides Parish, Louisiana. Specimens had been fixed in 
10% formalin, washed in water, and stored in 50% iso- 
propyl alcohol. Five specimens lacked information on 
date of collection and were utilized for dissection pur- 
poses only. The remaining lots were used for histological 
investigations of gonadal activity and sexual character- 
istics, as well as for anatomical studies. The collection 
dates and numbers of specimens used in the study of 
gonadal activity were as follows: October 1, 1973 (1 
specimen); October 5, 1974 (4 specimens); February 22, 
1975 (8 specimens); March 28, 1975 (4 specimens); April 
25, 1975 (3 specimens); June 21, 1975 (2 specimens); 
March 30, 1986 (5 of 16 specimens). 



Page 160 



THE NAUTILUS, Vol. 102, No. 4 




Figure 1. Photomicrograph of a transverse section through 
the gill of Margaritifera hcmheli showing the interlamellar 
junction (ILJ), 30 x 



Portions of the viscera of each specimen were infil- 
trated with paraffin, sectioned at seven micrometer thick- 
ness, and stained with Ehrlich's hematoxylin and eosin. 
Some sections were stained in a picro- ponceau connective 
tissue stain following the method described in Humason 
(1979:147). At least five slides were prepared for each 
specimen. The barren gills of two specimens and a por- 
tion of the posterior mantle lobe of one specimen were 
also sectioned in a similar manner and stained with picro- 
ponceau connective tissue stain. 

Dissections were undertaken on the stomach and vis- 
ceral nervous system of eight specimens. Three speci- 
mens were investigated for gro.ss morphology of the gills, 
nervous system, and excretory system. The method of 
dissection and exposition of specific internal organs, and 
the terminologv used to describe various organ compo- 
nents, follows Smith (1980, 1986). 

.\11 material relevant to this study has been cataloged 
in the invertebrate collections of the Museum of Zoology 
(Nos. MO. 1643-1645), University of Massachusetts, Am- 
herst, Massachusetts. 

RESULTS 

Gross Anatomy 

Anterior and posterior adductor muscles subequal, foot 
musculature and associated pedal and retractor muscles 
well developed Cerebral and pleural ganglia fused, kid- 
ney with both glandular and non-glandular chambers, 
renal pore and gonopore closely set but clearly separate. 
Labial palps falcate and large, gills or demibranchs la- 
mellar, inner gill larger than outer gill, both inner and 
outer gills free from mantle posterior to pallial line. Both 



EX 



^.^: 



^d.r't-'^r' 










Figure 2. Photomicrograph of a transverse section through 
the posterior portion of the mantle of Margaritifera hembeli 
showing the partial!) contracted diaphragmatic septum (DS), 
which separates the exhalent (EX) and inhalent (IN) chambers, 
SO X. 



lamellae of each gill held together b> solid, separate 
interlamellar junctions (figure 1, ILJ), lined with squa- 
mosal epithelium and composed of loose connective tis- 
sue and fine fibers that are more muscular appearing 
than collagenous (in M. margaritifera. see Smith, 1979a). 
Interlamellar junctions for the most part arranged in 
oblique rows in typical margaritiferine fashion (Ort- 
mann, 1912; Smith and Wall. 1983: fig. lb), similar to 
gills of A/. niarr!ana<'(Ortmann, 1912:235). Gill junctions 
are somewhat patternless along lower margin and an- 
terior and posterior extremities of each gill plate. 

Mantle lobes free all around, no indication of division 
of exhalent region of mantle margins into .separate anal 
and supra-anal apertures. Inhalent and exhalent cham- 
bers separated posteriorly by diaphragmatic septa (figure 
2, DS), and though not observed in the living animal, 
are presumed to function similar to M. margaritifera 
(Smith. 1980) Inhalent margin of mantle with densely 
pigmented papillae, exhalent region pigmented, with 
crenulate margin. 

Gonadal Activity and Sexuality 

.All animals examined histologicalK were sexually ma- 
ture, including several small specimens ranging in shell 
length from 49 to fj9 mm (believed to be from 6 to 9 



D. G. Smith 1988 



Page 161 















'f 
^ 



^i V 




>*.. s' 



Figures 3-6. Photomicrographs of histological sections through male and female gonads of Margaritifera hembeli. 3. Gonad of 
a male specimen of M. hembeli collected in Februar\, following spawning of gametes, 65 x. 4. Gonad of a female specimen 
collected in February following spawning, 65 x. 5. Gonad of a male specimen of M. hembeli collected in October and showing 
male gametes which have filled the entire gonadal stroma, 100 x . 6. Gonad of a female specimen of M. hembeli collected in 
October and containing fully developed ova, 100 x . 



years old on the basis of shell annuli). This would suggest 
that M. hembeli matures at an earlier age than North 
American M. margaritifera (Smith, 1979b). No evidence 
of hermaphroditism was observed. 

Although no gravid females were among the available 
specimens, a specific reproductive cycle was indicated 
by the gonads of sectioned specimens. Animals collected 
in Februarv', March, and April showed characteristic post- 
spawning features (figures 3, 4) including partial or com- 
plete occlusion of gonadal acini by granules, presumed 
pycnotic cells, and unspawned gametes in various stages 
of development or cytolysis. Animals collected in June 
showed little change from April specimens indicating 
that complete resorption of reproductive tissues, corre- 
sponding to an undifferentiated stage of non-reproduc- 
tive activity, apparently does not occur. By early Octo- 
ber, gonadal activity is resumed and sex cells, including 
the latter stages of spermatogenesis and oogenesis, are 
abundant within all observed acini (figures 5, 6). It is 
therefore hypothesized that the oviposition of eggs into 
marsupial demibranchs and the spawning of male ga- 



metes takes place sometime between late November and 
late December with release of larvae occurring in late 
December or January. Based on examined specimens, 
there is no evidence that production of glochidial larvae 
takes place at any other part of the year. 

Visceral Nerve Anatomy 

In M. hembeli, the first pallial bifurcation (figure 7, BI) 
of the posterior nerve is well anterior of the mantle nerve 
separation, usually arising from the visceral ganglion it- 
self at a point near the origin of the posterior nerve cord 
(figure 7, PNC). Some variation exists in the position of 
this bifurcation relative to the visceral ganglion. The 
overall pattern differs from that observed in other eastern 
North .American margaritiferids including A/, marrian- 
ae. in which the first bifurcation is normally posterior of 
the visceral ganglion, but similar to that observed in 
Cumberlandia monodonta (Smith, 1980, unpublished 
data). Moreover, in both M. hembeli and M. marrianae, 
the accessory nerve, which is typically present in M. 



Page 162 



THE NAUTILUS, Vol. 102, No. 4 



VG 




PNC 



OES 





Figures 7-9. .\natoniy of the visceral nervous system and 
stomacli of Margarilifcra hcmbeli. 7. Visceral ganglion (VCJ) 
of M. hemheli. 20 x. 8. Morphology of the stomach roof of 
M. hembeli, 5 x . 9. Morphology of the stomach floor of M. 
hemheli. 5 x . APR, anterior protractor muscle; ASA, anterior 
sorting area; .ASR, anterior sorting area of roof; BI, first pallial 
bifurcation; BN, branchial nerve; C, commissure to cerebro- 
pleural ganglion; MN, mantle nerve; PNC, posterior nerve cord; 
OES, esophagus; PSf\, posterior sorting area; HD, right duct; 
RSA, right side sorting area; T, major typhlosole. 

margaritifera and C. monodonta (Smith, 1980), is almost 
aKva> s absent, being observed only once in M. marrianae 
(Smith, unpublished data). 

Stomach Anatomy 

In addition to the posterior sorting area, which is typical 
of other eastern North American margaritiferids, the 



stomach roof of M. hembeli contains a well developed 
anterior sorting area and a continuation of the posterior 
sorting area extending along the right side of the stomach 
roof and separated from the anterior sorting area by a 
ridge (figure 8). 

The floor of the stomach interior (figure 9) is char- 
acterized by an anterior sorting area that is somewhat 
similar to M. marrianae (Smith, 1986; fig. 4b); however, 
the right side sorting area of M. hembeli is completely 
unlike ,V/. marrianae in that a distinct platform is absent 
in A/, hembeli and the sorting ridges run primariK lat- 
eralK , rather than in the anterior-posterior pattern seen 
in M. marrianae. A groove sets off the right side sorting 
area from the anterior sorting area in A/, hembeli and 
the morphologN of the stomach Door sorting areas of M. 
hembeli can be considered more similar to M. marga- 
ritifera in overall appearance than to other eastern North 
.\merican species. 

DISCUSSION 

As demonstrated by anatomical investigations of the vis- 
ceral nervous system and the stomach (this studv), and 
conchological differences discussed elsewhere (Johnson, 
1983; Smith, 1983), A/, hembeli and A/, marrianae clear- 
1\ represent distinct lineages within the genus Marga- 
ritifera. Although both species have lateral teeth, these 
teeth are also present in M. auricularia (southern Europe) 
and A/, laosensis (southeast .\sia). Lateral teeth may 
therefore represent structures that have arisen separately 
in different "stocks of margaritiferid species, or ma> be 
symplesiomorphies indicative of an ancestral relation- 
ship. 

The presence of a corrugated surface of the posterior 
slope of the shell (weakK expressed in A/, hembeli) and 
the close geographical proximit\ of these two species are 
the strongest lines of evidence indicating a relationship 
between them. Ne\ ertheless, the pattern of \isceral nerve 
bifurcation and stomach morphologv in A/, hembeli and 
M. marrianae show interspecific variation as great as 
that observed between either of these two species and 
M. margaritifera or C. monodonta. Biochemical data of 
the sort de\eloped for M. margaritifera. C. monodonta, 
and A^. hembeli b> Davis and Fuller (1981) is not avail- 
able for M. marrianae, precluding a comparison of ge- 
netic distances among the four species. However, the 
available biochemical data i Davis and Fuller, 1981), 
combined u ith anatomical information presented in this 
stud) and elsewhere (Smith, 1980, 1986), clearK supports 
the concept that the North .American species of the Mar- 
garitiferidae have been isolated from one another for a 
considerable period of time (Smith, 1976). 



ACKNOWLEDGEMENTS 

I thank Malcolm F. Vidrine, Mark E. Gordon, and James 
E. Williams for gracioiisK- supplying me with the pre- 
served specimens used in this studv. Mark E. Gordon 
kiiulK read an early draft of the paper. 



D. G. Smith 1988 



Page 163 



LITERATURE CITED 

Conrad, T. A. 1838. Monography of the family Unionidae, 
or naiades of Lamarck (fresh water bivalve shells) of North 
America, Numbers 10, 11. Philadelphia, PA, p. 81-102 

Davis, G. M. and S. L. H. Fuller. 1981. Genetic relationships 
among Recent Unionacea (Bivalvia) of North ,\merica. 
Malacologia 20:217-2.53. 

Humason, G. L. 1979. .Animal tissue techniques, 4th ed W . 
H. Freeman & Co., San Francisco, 661 p. 

Johnson, R. I. 1983. Margaritifera marrianae, a new species 
of Unionacea (Bivalvia: Margaritiferidae) from the Mo- 
bile-.Alabama-Coosa and Escambia River systems, Ala- 
bama. Occasional Papers on Mollusks (Harvard Universitv) 
4:299-304. 

Ortmann, .\. E. 1912. Notes upon the families and genera of 
the najades. .Annals of the Carnegie Museum 8:222-.365. 

Smith, D. G. 1976. The distribution of the Margaritiferidae: 
a re\iew and a new synthesis Bulletin of the .American 
Malacological Union 1976:42 (abstract). 

Smith, D. G. 1979a. Marsupial anatomy of the demibranch 
of Margaritifera margaritifera (Lin. ) in northeastern North 
America (Pelecypoda: Unionacea). Journal of Molluscan 
Studies 4.5:39-44. 

Smith, D. G. 1979b Se.xual characteristics of Margaritifera 



margaritifera (Linnaeus) populations in central New En- 
gland. Veliger 21:381-383. 

Smith, 1) (; 1980 Anatomical studies on Margari/i/era mar- 
garitifcra and Cumherlandia monodonia (Mollusca: Pe- 
lec> poda: Margaritileridae). Zoological Journal of the Lin- 
nean Society (i9:257-270. 

Smith, D. G. 1983. On tlie so-called mantle muscle scars on 
shells of the Margaritiferidae (Mollusca: Pelecypoda). with 
observations on mantle-shell attachment in the Unionoida 
and Trigonioida. Zoologica Scripta 12:67-71. 

Smith, D. G. 1986. The stomach anatomy of some eastern 
North .American Margaritiferidae (Unionoida: L'niona- 
cea). .American Malacological Bulletin 4:13-19. 

Smith, D. G. and W. P. Wall. 1983. The Margaritiferidae 
reinstated: a reply to Davis and Fuller (1981), "Genetic 
relationships among Recent Unionacea (Bivalvia) of North 
America." Occasional Papers on Mollusks (Harvard Uni- 
versity) 4:321-330. 

Stewart, J. H. 1988. Endangered and threatened wildlife and 
plants; final endangered status for the Louisiana pearl shell 
"Margaritifera hembeli." Federal Register 53:3567-3570. 

\'idrine. M. F. 1985. Fresh-water mussels (Unionacea) of 
Louisiana; a zoogeographical checklist of post-1890 rec- 
ords. The Louisiana Environmental Professional 2:50-59. 



THE NAUTILUS 102(4): 164-166, 1988 



Page 164 



The Two Printings of J. F. Gmelin's Systema Naturae, 
13th Edition (1788-96) 



Alan R. Rabat 

Deparlmenl of Mollusks 
Museum of Comparative Zoology 
Harvard University 
Cambridge, MA 02138, USA 



Richard E. Petit 

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



One of the more important 18th century reference works 
in systematics is J. F. Gmelin s l.'3th edition of Linnaeus' 
Systema Naturae. This work represented a considerable 
updating of and expansion upon the 12th edition of the 
Stjstema Saturae (Linnaeus, 1767). Gmelin not only pro- 
\ ided additional bibliographic references for the species 
described by Linnaeus, but he also described numerous 
new species, using the same format as had Linnaeus. 
Gmelin's magnum opus was published in three volumes, 
comprising 10 parts altogether: Regntim Animalium (7 
parts); Regnum VegetaMe (2 parts); and Regnum La- 
pideuni (1 part). For comments on the structure and 
content of Gmelin's work, see Dodge (1958:157-158). 

Few systematists, however, have realized that Gme- 
lin's work actually underwent two separate printings. 
The initial German printing was by Georg. Emanuel. 
Beer of Lipsiae [= Leipzig]; the later French printing 
was by J. B. DelamoUiere (subsequently as Bernuset, 
Delamolliere, Falque et Soc. ) of Lugduni [= Lyon]. Both, 
of course, were written in Latin. 

Hopkinson (1907) provided a valuable collation of the 
Lipsiae printing, based on an analysis of various German 
literature abstracts and review publications. He noted 
that only the first part in the two multi-part volumes had 
a date on the title page and that this date "did not apply 
to the whole of the parts in that \olume " (Hopkinson, 
1907:1036), since the later parts were published in fol- 
lowing years. Hopkinson did not refer to the Lugduni 
printing. Agassiz (1852:69) listed this work as "Leipz. 
1788-1793,. . . Lugduni, 1789-1796." Engelmann (1846: 
103) provided the same dates as had Agassiz, but erro- 
neously gave the place of publication as "Lugd. Batav." 
which is actually Leiden [= Lugduni Batavorum]. Sher- 
born (1902:.\.\xv) gave the same dates, made the same 
geographical transposition as had Engelmann, and spec- 
ified that the second printing was a reprint of the first. 
Hulth (1907:13) and Soulsby (1933:16, 50) provided a 
partial collation ol these two printings; however, their 
dates are based solely on those stated on the title page 
for the first part of each of the two multi-part volumes. 
Stafleu and Cowan (1976:956) briefly mentioned the two 



printings, and noted for the Lugduni printing: type 
reset, no copy seen . Frequently , librarx copies will have 
the same date stamped on the binding of each part or 
hand-written on the first page of each part; this is in- 
correct. The important facts are that the Lugduni print- 
ing was published after the Lipsiae printing, and was 
printed from reset type w hich resulted in a number of 
differences due to printer s error. Therefore, the date of 
publication of each part of the Lugduni printing must 
post-date that of the relevant part of the Lipsiae printing. 

L'nfortunately , we have not been able to determine 
the date of publication for the succeeding parts of the 
Lugduni printing (only for the first part in the multi- 
part volumes). However, we present this comparison here 
in order to alert s\ stematists to the existence of these two 
printings. Should the reader know of further bibliograph- 
ical sources bearing on this matter, we would greatly 
appreciate hearing from them. 

Herein we present as complete a collation as possible 
of the two printings of Gmelin's Systema Saturae. based 
on Hopkinson (1907) and on our examination of complete 
sets of this work in the libraries of the Museum of Com- 
parative Zoology , the British Museum (Natural History), 
and the Museum National d Histoire Naturelle, Paris. 

Gmelin, jo. Prid. [Joliann P^iedrich]. 1788-1793. Ca- 
roli a Linne, Systema Naturae per Regna Tria Naturae, 
Secundum Classes, Ordines, Genera, Species, cum cha- 
ractcribus, diftcrentiis, synonymis, locis. Editio Decima 
Tertia, Aucta, Reformata. Georg. Emanuel. Beer, Lip- 
siae; [Second printing, 1789-1796], J. B. Delamolliere, 
Lugduni. 

Tomus 1, REGNUM ANIMALIUM 

1(1) Mammalia to Aves Picae, pp. xii -I- 1-500 

Lipsiae, 1788; Lugduni, 1789. 
1 (2) Aves Anseres — A\es Passeres. pp. 501-1032 

Lipsiae, 1789; Lugduni (post 1789?). 
I (3) Amphibia — Pisces, pp. 1033-1516 

Lipsiae, 1789; Lugduni (post 1789?). 



A. R. Kabat and R. E. Petit 1988 



Page 165 



I (4) Insecta General, Coieoptera — Hemiptera, pp. 1517- 
2224 

Lipsiae, 1790; Lugduni (post 1790?). 

I (5) Insecta Lepidoptera — Aptera, pp. 2225-3020 

Lipsiae, 1790; Lugduni (post 1790?). 
1(6) Vermes, pp. 3021-3910 

Lipsiae, 1791; Lugduni (post 1791?). 
1(7) Inde.x 1-3, pp. 3911-4120 

Lipsiae, 1792; Lugduni (post 1792?), 

Tomus II, REGNUM VEGETABILE 

II (1) Monandria to Polyandria, pp. .\1 + 1-884 

Lipsiae, 1791; Lugduni, 1796. 
II (,2) Didynamia to Cryptogamia, pp. 885-1662 
Lipsiae, 1792; Lugduni (1796 or later?). 

Tomus 111, REGNUM LAPIDEUM 

111, pp. 1-476, plates 1-3 (of crystals) 
Lipsiae, 1793; Lugduni. 1796. 

It has been our experience that molluscan taxonomists 
who have used Gmelin's work have usually referred sole- 
ly to the Lipsiae printing. Among others, these include 
Dodge (1958:157-158); and Kohn (1966) who provided 
biographical information on Gmeiin and a useful critique 
of his work. .\n example of the taxonomic problems that 
can arise with respect to the two printings of Gmelin's 
\\ ork is that of certain species of Cancellariidae (Gastrop- 
oda) reviewed by Petit (1984). For Buccinum piscato- 
rium Gmeiin. 1791 (p. 3496), the Lipsiae printing gave 
two references: "List. Conch, t. 1024 f. 89"' and "Martin. 
Conch. 4. t. 124 f. 1151, 1152". Based on this, Petit chose 
the figures from the latter reference (i.e., Chemnitz, J. 
H., 1780, Neues Systetnatisches ConchyHen-Cabinet , 
\"ol. 4, Nurnberg) to be representative of this species. 
Subsequently, A. Verhecken [in litt.) inquired as to how 
the Chemnitz illustration could be used in this manner 
since it was not cited by Gmeiin. In later correspondence, 
N'erhecken advised that he had found that he was re- 
ferring to a cop)- of the Lugduni printing. In that printing 
only one reference, the aforementioned "List. Conch, t. 
1024 f. 89" (i.e.. Lister, M.. 1692/1770, Historiae sive 
Synopsis Methodicae Conchyliorum, London), was giv- 
en for the species in question; the Chemnitz reference 
being omitted. ObviousK', the omission of the reference 
to Chemnitz was a printer's error, as a comparison of the 
two printings makes it evident that the type was reset 
and errors were made. 

Incidentally, the third volume of this work (the Reg- 
num Lapideum) was placed on the Official Index of 
Rejected and Invalid Works in Zoological Nomenclature 
(along with the related sections of Linnaeus and of Tur- 
ton), since the "generic names of fossil animals used in 
these works actually corresponded to the classes of Recent 
animals (ICZN, 1954, Opinion 296). 

We conclude that (1) since the Lipsiae printing was 
published first, it should serve as the basis for discussion 
of Gmelin's species; (.2) the Lugduni printing should be 



treated as a reprint (or reissue) in which errors occur, 
and not as a "second edition "; (3) systematists who are 
studying species described by Gmeiin should compare 
the two printings in order to uncover any discrepancies 
with respect to species descriptions or references. 

ACKNOWLEDGEMENTS 

Mr. Andre Verhecken, Mortsel, Belgium, first brought 
the discrepancy in the Gmeiin citations to our attention, 
and then followed up with information on the existence 
of the two printings. His interest and assistance is greatly 
appreciated. We thank the librarians of the Museum of 
Comparative Zoology, the British Museum (Natural His- 
tory), and the Museum National d'Histoire Natureile, 
who allowed us to examine the sets of Gmelin's work in 
their collections. A number of other hbraries were checked 
for this work; most had only partial sets of one or the 
other printing which did not yield additional useful in- 
formation. Prof. Kenneth J. Boss provided a helpful re- 
view of this manuscript. 

LITERATURE CITED 

Agassiz, L. 1852. Bibliographia Zoologiae et Geologiae, Vol. 
3 (of 4). Ray Society, London, 657 p. 

Dodge, H. 1958. .\ historical review of the mollusks of Lin- 
naeus. Part 6. The genus Trochus of the class Gastropoda. 
Bulletin of the American Museum of Natural History 
116(2):153-224. 

Engelmann, W. 1846. Bibliotheca Historico-Naturalis. Ver- 
zeichniss der Biicher iiber Naturgeschichte . . . Erster Band. 
Verlag von Wiihelm Engelmann, Leipzig, x -I- 786 p. 

Hopkinson, J. 1907. Dates of publication of the separate parts 
of Gmelin's edition (13th) of the 'Systema Naturae' of 
Linnaeus. Proceedings of the Zoological Society of London 
1907(49):1035-1037. 

Hulth, J. M. 1907. Bibliographia Linnaeana: materiaux pour 
servir a une Bibliographie Linneenne. Kungliga Veten- 
skaps Societeten I Upsala, 170 p. -H 11 pis. 

International Commission on Zoological Nomenclature, 1954. 
Opinion 296. Suppression, under the Plenary Powers, for 
nomenclatorial purposes, of volume 3 (Regnum Lapi- 
deum) of the Twelfth Edition of the Systema Naturae of 
Linnaeus published in 1768 and of the corresponding vol- 
ume in the Houttuyn (1785), Gmeiin (1793) and Turton 
U806) editions of the above work. Opinions and Decla- 
rations rendered by the International Commission on Zoo- 
logical Nomenclature 8(13):167-178. 

Kohn, A. J. 1966. Type specimens and identity of the de- 
scribed species of Conus III. The species described by 
Gmeiin and Blumenbach in 1791. The Journal of the Lin- 
nean Societv of London, Zoology 46(308):73-102, pis. 
1-3. 

Linnaeus, C. 1766-67. Systema Naturae, per Regna Tria Na- 
turae, . . . Editio Duodecima Reformata. Direct. Laur. Sal- 
vii, Holmiae. Regnum Animale, 1328 -t- 36 p. 

Petit, R. E. 1984. Some early names in Cancellariidae. Amer- 
ican Malacological Bulletin 2:57-61. 

Sherborn, C. D. 1902. Index Animalium sive index nominum 
quae ab .\.D. MDCCL\'III generibus et speciebus ani- 
malium imposita sunt . . Sectio Prima. L niversity Press, 
Cambridge, Ix + 1195 p. 



Page 166 THE NAUTILUS, Vol. 102, No. 4 



Soulsby, B. H. 1933. A catalogue of the works of Linnaeus Stafleu, F. A. and R. S. Cowan. 1976 [in 1976-86]. Taxonomic 

(and publications more immediately relating thereto) pre- literature: a selective guide to botanical publications and 

served in the libraries of the British Museum (Bloomsbury) collections with dates, commentaries, and types. Second 

and the British Museum (Natural History) (South Ken- Edition. Regnum X'egetabile; Bohn. Scheltema & Holke- 

sington). British Museum (Natural History), London, 2nd ma, Utrecht, \'ol. 1 (of 6):xl + 1136 p. 
ed., p. xii -I- 1-312; 7 pis. 



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