The Nautilus

THE NAUTILUS

Volume 115, Number 1
March 31, 2001
ISSN 0028-1344

A quarterly devoted
to malacology.

EDITOR-IN-CHIEF

Dr. Jose H. Leal

The Bailey-Mattliews Shell Museum
3075 Sanibel-Captiva Road
Sanibel, FL 33957

MANAGING EDITOR
Christina Petrikas

The Bailey-Matthews Shell Museum
3075 Sanibel-Captiva Road
Sanibel, FL 33957

EDITOR EMERITUS
Dr. M. G. Harasewych
Department of Invertebrate Zoology
National Museum of
Natural History
Smithsonian Institution
Washington, DC 20560

CONSULTING EDITORS
Dr. Riidiger Bieler
Department of Invertebrates
Field Museum of
Natural History
Chicago, I L 60605

Dr. Arthur E. Bogan

North Carolina State Museum of

Natural Sciences

Raleigh, NC 27626

Dr Phihppe Bouchet
Laboratoire de Biologie des
Invcrtebres Marins et Malacologie
Museum National d'Histoire Naturelle
55, rue Buffon
Paris, 75005 France

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

Dr. Eileen H. Jokinen
8234 E. North Shore Road
Sault Ste. Marie, MI 49783

Dr Douglas S. Jones
Florida Museum of Natural History
Universit)' of Florida
Gaine.sville, FL 32611-2035

Dr. Harry G. Lee

1801 Barrs Street, Suite 500

Jacksonville, FL 32204

Dr. Charles Lydeard
Biodiversity and Systematics
Department of Biological Sciences
University of Alabama
Tuscaloosa, AL 35487

Dr. James H. McLean
Department of Malacology
Los Angeles County Museum of
Natural History
900 Exposition Boulevard
Los Angeles, C A 90007

Dr. Paula M.Mikkelsen
Department of Living Invertebrates
The American Museum of Natural
History-
New York, NY 10024

Dr Diarmaid 6 Foighil

Museum of Zoology and Department

of Biolog)'
University of Michigan
Ann Arbor, MI 48109-1079

Dr Gustav Paulay

Florida Museum of Natural History
Universit)' of Florida
Gainesville, FL 32611-2035

Mr Richard E. Petit

PO. Box 30

North Myrtle Beach. SC 295S2

Dr. Gary Rosenberg
Department of Mollusks
The Academy of Natural Sciences
1900 Benjamin Franklin Parkway
Philadelphia, PA 19103

Dr. Geerat J. Vermeij
Department of Geolog)'
University- of California at Davis
Davis, CA 95616

Dr. G. Thomas Watters
Aquatic Ecology Laboratory
1314 KinnearRoad
Columbus, OH 43212-1194

Dr John B. Wise

Houston Museum of Natural Science

Houston, TX 77030-1799

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TH E€9NAUTI LUS

CONTENTS

vVooos Hole Or.6a''ioar3pr.ic irisutotion

APR I ^ 2Q01

Volunw 115, Number 1
March 31 2001
ISSN 0028-1344

Paulino J. S. de-Souza Rexision o{' tlie Recent Bullata Jousseaume, 1S75

Gary A. Coovert (Gastropoda: Marginellidae) witli tlie description of hvo

new species 1

Maximiliano Cledon Reproduction and brooding of Crepkhda iir^cntiiui.

Pablo E. Penchaszadeh Sinione, Pastorino and Penchaszadeli, 2()()() (Gastropoda:

Calvptraeidae) 15

G. W. Schmelz Revision of the Architectonicidae (Gastropoda:

Allogastropoda) from tlie Miocene Cliipola Formation, Oak-
Grove Sand, and Shoal River Formation of northern
Florida, with descriptions of four new species 22

Angel Valdes On tlie pulilication date, authorship, and t\pe species of

Unihniculuiii and Ti/lixlino (Gastropoda: Opisthobranchia:
Tvkxlinoitlea) 29

Notes

Richard E. Petit A note on Liicina luiilfiliiuatti "Tuoniev and Holmes"

( RivaKia: Lucinidae) 35

Bruce A. Marshall A replacement name for a New Caledonian Callidstditw

species (Gastropoda: Trochidae) 36

Erratum 36

Book Re\iew 37

Notices 38

THE NAUTILUS 115(1):1-14, 2001

Page 1

Revision of the Recent Bullata Jousseaume, 1875 (Gastropoda:
Marginellidae) with the description of two new species

Paulino J. S. de-Souza'

Depto. Zoologia, Inst, de Biolo2;ia,

CCS
Universidade Federal do Rio de

Janeiro
21941-.570 Rio de Janeno
BRAZIL

Can A. Coovert-

Da\ton Museum of Natural Histon
2600 DeVVeese Parkwav
Davton, OH 45414 USA

ABSTRACT

Tlie genus BuUat/i Jousseaume, 1875 is re\ised based on con-
chologicaj characters. All known li\ing species are endemic to
Brazil. Two new species are herein described, B ouerrinii and
B. analuciae. Bullata guerrinii is most similar to B. largillieri
(Kiener, 1841) as both have an enlarged second columellar pli-
cation which overrides and fuses with the first, but differs in
having a darker coloration, wider aperture, and spire onlv
slightly apparent. Bullata analuciae has similar color pattern to
B. largillieri. but differs in having clearly separate first and
second columellar plications, a generallv larger, thinner shell,
a broad aperture and non-denticulated lip. The other 4 known
species are described and discussed and a ke\' for identification
is presented. A cladistic analysis of Bullata was made using 22
conchoiogical characters (53 states). The single most parsimo-
nious tree obtained (lengdi 37, CI = SI, RI = 69) is as follows:
(B. bullata (B. analuciae {(B, largillieri. B. guerrinii) (B. lila-
cina. B mattheusi)))). The monophviv of the genus is sup-
ported b\- S SMiapomorphies.

Additional kei/ words:
cies, Brazil, ph\logen\"

Volutoidea, Neogastropoda. new spe-

INTRODUCTION

The genus Bullata Jousseaume, 1875 includes species
that live in tropical waters from the intertidal down to
70 m. Recent species of the genus, as it is currently
defined (Coovert and Coovert, 1995), are endemic to the
continental shelf off Br;izil. Yet, its fossil record can be
traced to the Miocene of Trinidad (Maun-, 1925) and
the Mio— Pliocene of the southeastern United States
(Redfield, 1870; Olsson, 1916).

As with most Marginellidae, throughout its taxonomic
historw species of Bullata were earlier referred to the
genus Mariiiuclla Lamarck, 1799. The genus was first

' Current address: Museu de Zoologia da Universidade de Sao
Paulo, RO. Box 42694, Sao Paulo, 04299-970, Brazil.
-Current address: 36 Prospect Avenue, Davton, OH 45415-
2614 USA.

differentiated as a group hv Svvainson (1833) who erect-
ed the genus Vohitclla, a name preoccupied, however,
by Vohitclla Pern', 1810 ( = Vasuin Roding, 1798), This
designation was, nonetheless, widelv used. In 1875, Jous-
seaume established Bullata as a replacement name for
Vohitclla hut put too much emphasis on the involute
spire as a diagnostic character for included species.
Thus, in Jousseaume's monograph, the genus was used
to encompass species that later were assigned to other
genera and families, such as Closia Grav, 1857, Pniuum
Herrmannsen, 1851, Cn/ptospira Hinds, 1844, Grauu-
lina Jousseaume, 1888; Gihhcnila Swiiinson, 1840; Pcr-
sicula Schumacher, 1817 (the last 3 genera belong in the
Cvstiscidae). Coovert and Coovert (1995), in their recent
taxonomic revision of the Marginellidae and Cystiscidae.
provided a more refined diagnosis ot the genus,

Similarlv to what probabK occurs widi other margi-
nellids, the two new species described herein have elud-
ed Brazilian malacologists for a long time due to their
limited ranges and subtlv different conchologiciil char-
acters. Both species could easily be mistaken for Bullata
largillieri (Kiener, 1841) due to somewhat similar color
patterns.

This work is part of an ongoing revision of the Bra-
zihan Marginellidae b\' the senior author. Institutional
abbreviations used are: ANSP: Academv of Natural Sci-
ences of Philadelphia, Philadelphia; BMiNHh The Nat-
ural Histon, London; BMSM: The Bailey-Matthews
Shell Museum, Sanibel, Florida: GAC; collection of Gar\'
A. Coovert; IBUFRJ: Instituto de Biologia AJniversidade
Federal do Rio de Janeiro, Rio de Janeiro; MNHN; Mu-
seum National d'Histoire Naturelle. Paris; MNRJ: Mu-
seu Nacion;il/Universidade Federal do Rio de Janeiro;
MORG; Museu Oceanognifico "Prof. Eliezer de Canal-
ho Rios", Rio Grande; MZUSP; Museu de Zoologiii/
Universidade de Sao Paulo, Sao Paulo; PMC; collection
of Paulo Marcio S. Costa. Rio de Janeiro; USNM; Na-
tional Museum of Natunil Histon', Smithsonian Institu-
tion, Washington,

Paee 2

THE NAUTILUS, Vol. 115. No. 1

SYSTEMATICS

FamiK Marginellidae Fleming, 1828
Subfamily Marginellinae Fleming, 1828
Tribe Prunini Coovert and Coovert, 1995
Genus Bullata Jou.sseaume, 1875

Bullntn Joii.sseaume. 1S75: 167, 250, nomen novum for Vohi-
tella Swain.son, 1833 {non Volutella Pern-, 1810)

Marginella (Volutella) Swainson, 1833: (2)1, Marginella pi. 1
[t\pe species: Marginella bullata Lamarck, 1822 = Valuta
bullata Bom, 1778: original designation] (nan Volutella
Pern-, 1810)

Gibbenilina Monterosato, 1884:139 (see Coovert, 1987: 27, for
further details).

Type species: Voluta bullata Bom, 1778; hv tautono-
m\-.

Diagnosis: Shell moderateh' large to ven- large for
famiK, colored with bands or other patterns, narrowK- to
broadlv obo\ate; spire immersed or nearly so; lip thick-
ened, denticulation usualK' present, absent in some: pos-
terior end of lip arched above ape.x; external varix pre-
sent; varix groove adjacent to body whorl, distinctly col-
ored; siphonal notch absent; parietal callus present to
nearly obsolete in type species; columella with 4 strong
phcations occupying less than half the aperture but not
crowded in anterior direction.

Description: Shell 14-98 mm in length, narrowK' to
broadl)' obovate (length: width ratio 1.53-1.8.3); surface
glossy, unsculptured, with 4-5 whorls. Color pattern
composed of indistinct darker spiral bands, with or with-
out rows of irregular white spots. Spire immersed to vers'
low and apparent, often covered bN' callus. Protoconch
apparent to completely concealed. Outer lip moderateh'
to heavily thickened, internally denticulated or smooth,
with a distinct, colored extemiil varix, particularK^ strong-
ly colored in varix groove adjacent to the bod\' whorl.
Lip with or without a distinct, flattened, beveled area in
anterior direction. Aperture narrow to moderately
broad, wider in anterior direction, some .species ■with an
obscure to distinct trough or siphonal "gutter" (best seen
in apical \iew'). Parietal callus nearl\' obsolete to strongh'
developed: posterior callus nearly obsolete to strongly
developed; left antero-ventral callus extending from the
anterior end of varix to level of third plication, often
resembling a fasciole. (Columella straight to concave.
witli 4 strong plications; phcations subetjual, or the first
and/or second strongly developed, sometimes fused;
third and fourth plications confined to aperture or slight-
ly emerging, distal ends truncate or gradualK diminish-
ing. Space behveen plications increases in posterior di-
rection, fourth plication often remote. Plication angle in-
creasing in posterior direction relative to longituilinal
axis.

Remarks: Several authors (see species svnonymv lists,
q.v.} have variously combined Bullata and C.losia Cray.
1857, apparently based on the superficial resemblance
of shell shape, especially that of B lihirina (Sowerbv.

1846). Closia differs in a number of conchological char-
acters, chiefly the very thin, very sharply formed plica-
tions that are strongly crowded in anterior direction.
Bullata. on the other hand, has much thicker, rounded
plications that are not nearly as crowded in anterior di-
rection. The labial denticulation of Closia is sharply, reg-
ularly formed and widely spaced (i.e., separated by a
distance greater than the thick-ness of a denticle). The
labial denticulation in Bullata is irregularly formed and
crowded or nearly absent. The columella in Closia is
concave, resulting in a sinuous outhne, while in Bullata
the columella is more straight. Closia has a much more
completely immersed spire and a tendency toward a
much lighter shell. Closia is probably closely allied to
Ovaainclla (Coovert and Coovert, 1995: 87) and thus in
the Austroginellini. Bullata has an oesophageal caecum
just posterior to the nerve ring, tspe 6 radula (Coovert
and Coovert, 1995: 56-57), and lacks the Valve of Lei-
blen, character states that allocate the genus placing it
in the tribe Pnmini. .\lso, these genera are restricted in
their biogeographv to specific oceanographic basins: Bul-
lata to Southern West Atlantic and Closia to the Indian
Ocean (Coovert and Coovert, 1995: 87). Thus, we con-
sider Closia to be a separate and unrelated genus, which
includes C. sarda (Kiener, 1834) as the t\pe species, as
well as C. majuscula (Martens, 1880) and C. ptinceps
(Sowerbv, 1901).

Cn/ptospira angustata (Sowerbv, 1846) was included
in Bullata by Jousseaume (1875: 251), due to the im-
mersed spire. But most species of Cn/ptospira can be
differentiated h\ their much stronger tasciole-like callus,
much stronger and more numerous columellar plications
that occupy more than half the aperture, and extensive
callusing posterior to varix. Thus we consider Cn/pto-
spira to be a separate genus with a separate origin but
within the Pnmini, with distribution restricted to the
Indo-Pacific Region (Coovert and Coovert, 1995: 93).

Although described as Bullata lipci Clover. 1990. we
agree with Lipe and Sunderland (1991:15), and consider
this a .species of Pntnum, probably a close relative of
Pninum rostratum (Redfield, 1870). The narrow shape
of the shells, the ventral area of the body whorl just
posterior to the plications, which is built up b\ callus
and forms a groove just posterior to the first plication,
and the very reduced, remote fourth plication, are very
similar in these tvvo species. Also both species are from
the Yucatan region of Me.xico. The involute shell of
Pnaunii lipci (Clover. 1990) with the posterior end of
the lip arching above the spire, represents a combination
of shell traits that appears in other marginellid genera
and, although present in Bullata. is not a diagnostic char-
acter this geiuis.

Bullata gucrhnii new species
(Figures 1, 7, 10)

Diagnosis: Color pattern of irregular, squarish cream

spots; spire slighlK but distinctly app;u'('nt. extremely

p. J. S. de-Souza and G. A. Coo\ert, 2001

Page 3

Figures 1-6. BuUata species in ventral and dorsal views. 1. Holohpe of Bullata <:^ucnimi new species, MNRJ 78 IS. length 27.0
mm, width 16.5 mm, 2. Holotvpe of BuUata analudae new species. MNRJ 7186, lengtli 27.0 mm, v\idth 18.1 mm. 3. Bullata
bullata {Bom, 177S), IBUFRJ 8471, length 82.1 mm, width 45.2 mm. 4. Bullata lar^illieri (Kiener, 1841), PMC 939, length 21.3
mm, width 13.4 mm, 5. Bullata lilacina (Sowerby. 1846). PMC 476, length 24.5 mm, width 14,9 mm

Mol and Turscli, l'-)(-)7v PMC 937. length .39.0 mm, width 22,9 mm.

3
6. Bullata matthewsi (van

Page 4

THE NAUTILUS, Vol. 115, No. 1

Figures 7-12. Biilliiln spi-cit-s: .spire, postcnor labial insertion, and colunicllar structures. 7, 10. Holot\pe ot Bullata gueninii
new .species. 8, 11. Holotspe of B. analuciac new species. 9, 12. B hir<iilliiri (Kiener, 1841), PMC 939.

low, dome-like, partialK covered bv po.sterior Iahi;il in-
sertion and posterior parietal callns; aperture brown in-
ternally, becoming paler toward the lip; first and second
plications distalK' fused, forniins; 1 strong plication ridge.

Description: Shell of medium size (25-29 mm in
length), moderately heavy, obovate (length: v\idtli ratio
1.61-1.67, X = 1.6.39, s = 0.0199, n = 6), v\ith approx-
imately 4.5 whorls. Color pattern of irregular s(juarish
cream spots, arranged in irregular axial rows, on pur-
plish-browii backgroimd, crossed by 4 indistinct darker
spiral bands (1 near suture, 2 narrower ones at mid-
section of bodv whorl, 1 near anterior end). Spire ex-
tremely low, dome-like, slight 1\ but distincth' apparent,
partially covered by posterior labial in.sertion and pos-
terior parietal callus. Protoconch visible or partialK so.
dark-brown. Outer lip thickened intemallv, with distinct,
heavy external varix, thinning abruptly in posti'rior di-
rection, with indistinct, flattened, beveled area thinning
evenly in anterior direction, labial denticulation very
fine, obsolete to indistinct, varix groove excavated. Outer

lip brov\iiish-white, darkening toward \arix groove. Ap-
erture narrow, slightK wider than lip thickness when
measured ventrally wider in anterior direction and
somewhat angled at posterior end. Aperture brown in-
ternally becoming p;iler toward lip. Parietal wall convex
with continuous ventral callus, thickening toward aper-
ture, thinning abniptK along collabral line just outside
aperture. Ventral callus extending in anterior direction,
thinning but smootliK joining left aiitero-ventral callus.
Posterior callus forming rounded nioimd separated from
posterior lip insertion by notch. Left antero-ventral cal-
lns extending from anterior end of varix to level of fourth
plication, with distinct furrow between second and third
jilieations. and continuing as shaqi. distinct, straight con-
tinuation of varix. Ventral ciillus whitish brown. Colu-
mella slightly concave, plications somewhat strong. First
plication much w(>aker than second, both distalK fused,
forming 1 \en strong plication ridge, separated from the
parietal callus by distinct groove. Third plication stron-
ger than fourth, weaker than plication ridg(\ (extending

p. J. S. de-Souza and G. A. Coovert, 2001

Paue 5

Bullata lilacina
Bullala largillieri
Bullatu giierriiiii

Bullulu mattheHsi
•S^ Bullata biilluta
•■•'.'...".'•• Bullata unaluciae

-0°

lOOU km

-20°

Figure 13. Geographic ranges ol the li'
lata

spe

cies ol Bul-

slightK outside aperture; fourth plication confined to ap-
erture. Space between plications increasing in posterior
direction, with fourth plication somewhat remote (i.e.,
distance hehveen third and fourth plications distinctK'
greater than distance between second and third). Third
and fourth plications gradually narrowing distalK'.

Type material: Holot^pe, MNRJ 78LS (length 27.0
mm, width 16.5 mm): ' Paratypes:' BM(NH) 1996417
(length 2.3.;3 mm, width 14.2 mm); IBUFRJ 10000
(length 25.7 mm, width 15.4 mm); MNHN (length 29.8
mm', v\idth 18.3 mm); MZUSP 28849 (length 26.1 mm,
width 15.7 mm); USNM 890896 (length 24.4 mm, width
15.1 mm), all from 40 km NE off Alcoba^a, Bahia State,
Brazil, 10-35 m.

Type locality: Pedra da LLxa (17°41.5- S, 038°59- W),
Parcel das Paredes, 27 km E off Caravelas, Baliia State,
Brazil, 5-15 m.

Geographic range: Known from tvpe locality and,
accoreling to local fishermen, from reefs around Abrol-
hos Archipelago (figure 13).

Remarks: Bullata t^iicniiiii is most similar to Biilhita
largillieri (Kiener, 1841) in their color patterns and fused
first and .second pUcations (figures 10, 12). But these
species differ in their shell shape, general color, spire

(figures 7, 9), \cntral callusing, aperture width, shape of
e.xtemal lip, and shape of the third and fourth plications
(figures 10. 12).

Bullata ouerriiiii is easily distinguishable from B hiil-
luta. B lilacina (Sowerbv, 1846) and B luatfhcwsi (\an
Mol and Tursch. 1967). I)\ its spotted color pattern,
while the others ha\f handed color patterns (figures 1-
6).

For a summar\ ol these differences, refer to dichot-
omous ke\ included later in this paper.

Etymology: iiin-rrinii. spec ies detlicated to Mr. I-licar-
do Cueriini.

Bullata aiialuciac new species
(Figures 2, 8, ID

M/ir^incllii [Chi.si/i) Uuvillicri — lusenhern, 19S1: 12(i. fit;. S

(non ICiener. 1841).
Mnrgindla Inroillicri. — Abbott and Daiici-, 1986: 2.34. nnninii-

bered fig. (non Kiener, 1841).
Bullata aff. bullata— Leal, 1990: 244.
Bullata cf. /flrg/Z/icri.— Lipe, 1991: 2-3. pi. 1, fig. 2.
Bullata .sp.l.— Leal and Boiichet, 1991: 23.
Clo.sia cf. largillii'ii — Bozzetti, 1996: .54, nnniinibered fig.

Diagnosis: (."olor pattern of small irregular cream
spots arranged in ;L\ial rows. e\enl\ superimposed on
caramel-brown background, crossed b\ 4 indistinct dark-
er spiral bands; labial denticulation absent or reduced to
veA' obscure imdulations; aperture wide, broader than
lip thickness when measured \entralK'; first and second
columellar plications completeK separate, sube(ju;d in
size. Thirtl plication is subeipud in size to first 2, fourth
plication weaker and confined to aperture.

Description: Shell small to medium (15-38 mm in
length), obovate (length;widtli ratio 1.49-1.70, x =
1.618, s = 0.0669, n = 18), with approximately 4 whorls.
Surface glossw without sculpture; color pattern ol small.

irregular cream spots arranged in ;L\ial rows, e\'enl\' su-
perimposed on caramel brown background, crossed by
4 indistinct darker spiral bands (1 near suture, 2 narrow^-
er ones at mid-section of bod\' whorl and 1 near anterior
end). Spire almost completeK' inunersed, flattened to
slightK' depressed, exposed but covered b\- glassy trans-
parent callus, partially covered by posterior lip insertion
and posterior parietal callus. Protoconch partialK \isible.
large, dark-brown. Outer lip thickened intenialK'. with
distinct external \arix, thicker medially and thinner to-
ward extremities, lacking distinct, flattened, beveled area
in anterior direction; labial denticulation absent or re-
duced to verx' obscure undulations; varix groove distinct-
Iv excavated. Outer lip white, exteniallv darkening to
bright brownish-orange in varix groove adjacent to body
whorl. Aperture wide, broader than lip thickness when
measured ventrallv, curving in posterior direction,
broadest in anterior direction. Aperture inteinally me-
dium-brown with lilac stain at anterior end. Parietal wall
convex with thin ventral callus wash that is thicker at
anterior and posterior ends. Posterior callus in fully adult

Page 6

THE NAUTILUS, Vol. 115, No. 1

shells forms peaked niouiul separated from posterior lip
insertion In' distinct i^ap. Lett untero-ventral callus ex-
tends from anterior end ot \arix to level of third plica-
tion, producing sharp, distinct, and straight continuation
of varix. Columella slightK- concave, plications slender
First and second columellar plications completelv sepa-
rate, suhequal in size. Third plication is subetjual in size
to first and second, fourth plication weaker and confined
to aperture. Space bet^veen the plications increases in
posterior direction, with fourth plication remote (i.e. dis-
tance between third antl fourth plication cbstinctlv great-
er than distance between second and third). Third and
fourth plications gradualK' narrowing distalK, ending in
sharji points.

Type material: Holot\pe, MNRJ 7186 (length .34.7
mm, width 20.4 mm); Parat\pes: ANSP .390:361 (length
32.8 mm, width 19.3 mm): ANSP .399366 (length 27.7
mm, widdi 18.0 mm); ANSP 399995 (length 34.5 mm,
width 20.4 mm); BM(NH) 1996071 (length 34.2 mm,
width 20.9 mm); BMSM, 1006 (length 31.2 mm, width
19.S mm); IBUFRJ 8463Parat\pe 3 (length 37.2 mm,
width 23.3 mm); IBUFRJ 8464 (lengdi 32.0 mm, width
20.1 mm); MNHN. (length 30.7 mm, width 19.0 mm);
MNRJ 7187 (length 28.rmm, width 16.8 mm); MORG
33311 (length 3.3.4 mm, width 21.0 mm); MORG 3.3312
(length 27.0 mm, width 18.1 mm): MZUSP 28243
(length 31.6 mm, width 20.0 mm); USNM 880120
(length 25.2 mm, width 16.9 mm), all from off Vitoria
(20''20' S, 04OW W), Espirito Santo State, Bnizil, 60-
70 m, trawled b\' shrimp fishing boats on mudd\' sand
bottom; GAG M2861 (length 3.3.6 mm, width 19.9 mm),
off Vitoria, Espirito Santo State, Brazil, .50-70 m, trawled
b\- shrimp fishing boats on silt\- sand; AMNH 213931
(length 29.9 mm, width 18.0 mm), off Espirito Santo
State, Bnizil, 50 m; AMNH 2139.30 (length 30.5 mm,
width 18.2 mm); GAG M1426 (length 30^3 mm, width
18.7 mm), both from off nortliern coast of Rio de Janeiro
State, Brazil, 50 m. mudcK bottom.

Type locality: Off Vitoria (20 20' S, 040 OO' W), Es-
pirito Santo State, Brazil, 60-70 m, trawled b\ shrimp
fishing boats on muddv sand bottom.

Geographic range: From tiie northern coast of Rio
de Janeiro State to the central coast of Espirito Santo
.State, Brazil (figure 13).

Remarks: In color pattern B iinalurkic is most similar
to B lar^illicri and B. ^ucrriuii. but the\- differ in shell
shape, spire form (figures 6, 7, 8), ventral callusing, ap-
erture width, shape and thickness of e.xtenial lip. and
columellar structure (figures 10, 11, 12).

Bitllafa aiwhiciac is most similar, in shell shape and
columellar structure, to B. buUata. but tlie\ differ in the
color pattern, .spire form, ventral callusing, and aperture
shap(> and v\idth.

Bullata lilacina and B. matthewsi differ in color pat-
tern and geographical range.

Etymology-: This species is dedicated to Ana Lucia

Rodrigues PeLxoto who first collected most specimens
aboard shrimp fishing boats.

Btdlata bullata (Bom, 1778)
(Figure 3)

Valuta bullata Bom, 1778: 205-206. Type information could
not be directly obtained, holot\pe fomierlv in the Musei
Caesarei V'indoboneiisis, Vienna, but declared missing
(Tomlin, 1917: 22.5): t\pe lotalih-: Indian Ocean [in error,
endemic to Bahia State, Brazil]: Bom, 1780: 218; Tomlin,
1917: 2.55.

MargincUa bullata. — tl'Orbignv, 1841: 415; Sowerbv. 1846:
401, pi. lw\ii, figs. 158, 159; Petit, 1851: .55: Reeve, 1860:
52: Reeve, 1864, pi. 1; Redfield, 1870: 225; Weinkauff,
1879: 24, pi. 4, figs. 3, 4; Paetel, 1888: 191; Smidi, 1945:
71, fig. 950; Rogers, 1951: 89, pi. 24, fig. 5: Dance, 1976:
193, unnumbered figure.

Valuta aiuin Gmeiin, 1791: 3448.

Mar^iulla magna Swainson, 1822: appendix: 12.

Marginella (Volutclla) bullata. — Swainson. 1833: .^dams and
Adams, 1853: 192; Trxon, 1882: 35, pi. 10, figs. 3. 4: Fi-
scher, 1883: 602: Tryon, 1883: 173, pi. 55. fig. 61.

Marginella bellangeri Kiener, 1834: 27-28, pi. 9, fig. 43; Catlow
and Reeve, 1845: 291; Reeve, 1860: 52.

Marginella cuvieri Deshaves, 1853: 75 (Explicatian cles Planch-
es): 1857-58: pi. 123, fig. 8: Redfield, 1870: 262,

Bullata bullata. — Jousseaume, 1875: 250; Tomlin, 1817: 244;
Coan, 1965: 189: Rios, 1970: 113. pi. 40; Oliveira ef nL
1972: 14; Rios, 1975: 117, pi. .35, fig. 506; Wagner and
Abbott, 1978: 22003; Rios, 1985:121, pi. 42, fig. 537:
Coovert, 1986a: 2: Coo\ert, 1986b: 3; Coovert and Lee.
1989: 4: Coovert and Coovert. 1990: 2, fig. 1; Rios. 1990:
8, fig.; Lipe, 1991: 2. pi. 1 fig. 10: Rios, 1994: 147, pi. 48,
fig. 638: Bozzetti. 1994 p. 54^"; Cooxert and Coo\ert, 1995:
92, fig. 66: Rosenberg, 1996, [gopher://erato.acnat.sci.org:
70/OR 1 8 1575.3- 1 8 1 6546-/.\vasp/.text/w;ispgoph .cxt] .

Bullata cuvieri (Deshaves, 18.53). — Jousseaume. 1875: 251.

Marginella (Volutella) bellangeri. — Kobelt. 1878: 72. pi. 27,
fig. 2.

Marginella (Bullata) bullata.— ThltAe. 1929: .3.55. fig. 429; Ei-
seuberg, 1981: 129, fig. 14.

Closia [Bullata) bullata.— \Xenz, 1943: 1376, fig. 3892.

Marginella (Clo.sia) bullata —Morretes. 1949: 104.

Marginella bullata — Santos. 19.55: 113, luinumbered figure.

Diagnoses: Shell large (34—101 nun), slender; color
pattern of numerous thin to verv thin darker spiral bands
crossed b\- thin ;L\ial lines on ros\-gra\ background; ob-
solete crenulation on outer lip; aperture wide, broader
than lip thickness when measured \entrall\ ; \entral cal-
lus reduced to obsolete wash.

Description: Shell large (34-101 mm), smooth, elon-
gated, narrowlv obovate (length:width ratio 1.51-1.96, x
= 1.83, s = 0.0946, n = 25), with about 5 whorls. Color
pattern composed of numerous thin to very thin darker
spiral bands crossed b\' thin, mostK pale or occasionalK
darker iixial lines on ros\' grav background (usually fading
into pinkish-tan with time, in museum specimens). A.xi;d
lines are apparentk' associated with incremental growth
lines. Spire immersed and covered bv a vitreous callus,
somi'times \isible, rareK partK covered bv lip insertion.

p. J. S. de-Souza and G. A. Coo\ert, 2001

Page 7

which imparts flattened to depressed shape to posterior
end of shell. Protoconch darker than earK- teleoconch
whorls, usualK' completeK' co\ered b\' vitreous callus.
Outer lip diickened internally, with distinct external var-
L\, thinner toward extremities, with an obscure, flat-
tened, beveled area in anterior direction, and often
abruptK' thinned in anterior direction, labial denticula-
don usually reduced to obscure undulations, at most pre-
sent as obsolete denticles. Varix groove generalK filled
forming wide, rounded, or flattened shelf Outer lip
white, e.xtemalK' darkening to orangish-browii, especiallv
on varix groove adjacent to body whorl. Aperture wide,
broader than lip thickness measured ventrallv, curving in
posterior direction, broadest in anterior direction. Ap-
erture intemalK brown, becoming white toward lip. Pa-
rietal wall convex with thin ventral callus wash. Posterior
callus nearly obsolete. Left antero-ventral callus extends
from anterior end of varix to level of third plication in
gradual curve toward columella, resembling a fasciole.
Columella slightly concave, plications slender to some-
what heavy. First and second columellar plications sep-
arate to fused distallv, subequal in size. Third plication
is subequal in size to first and second plications, with
fourth plication subequal to slightly weaker and confined
to aperture. Space beUveen plications increases in pos-
terior direction, with fourtli plication remote (i.e., dis-
tance between diird and fourth plication distinctly great-
er than distance between second and third). Third and
fourth plications gradually narrowing distallv.

Material examined: Tvpes of Maivinclla cinicii De-
shaves, 185.3, MNHN unumbered, Bahia, Brazil; IB-
UFRJ 1.540. off SaKador, Bahia State, Brazil. .5-10 m,
B. Linhares col: IBUFRJ 8471, off Salvador, Baliia
State. Brazil. .5-10 m, B. Unbares col.: MZUSP 547, off
SaKador. Baliia: MZUSP 15686, Mar Grande, Itaparica,
Bahia State, Brazil, G. May col.: MZUSP 27267, off Sal-
vador, Bahia State, Brazil.

Geographic range: Endemic to Bahia State, Brazil
(figure 1.3).

Remarks: BiiUaia hullata is most similar, in size and
color pattern, to B. matthewsi, but they tliffer in back-
groimd color, spire form, callus structure, aperture
width, and labial denticulation. In addition, their geo-
graphical ranges are widely disjunct (figure 13).

See remarks under B. gitcninii and B aualuciac. and
kev for comparison with B. bnllata

BuUata biilhita differs from B. lar^illuri and B lila-
cina in size, shell shape, color pattern, structure of ven-
tral callus, aperture width, and columellar structure.

Apparent spire, yellowish coloration and smaller size
in original illustration of MargineUa cuvieri Deshayes,
1853 seemed to indicate that this species was a senior
synonym of B matlhcusi. But in examining tvpes, it be-
came clear that they were smaller discolored B Jmlliita.
hvo of them with atvpicallv apparent spires (but covered
with callus). The broad medial white band shown in the
orisinal illustration was due to shell deterioration This

"variety" was later discussed bv Sowerbv (1846: 401, sp.
104: pi. 77, figure 159) who did not mention Deshaves's
work.

BuUata largillieri (Kiener, 1841)
(Figures 4, 9, 12)

MargineUa laraiUicri Kiener. 1841: 4.3-44. pi. 11, fij^. 3. Ho-
lohpe not found, hpes were fornierlv deposited in Rouen
Museum, bnt are now missing (P. Bouchet, personal com-
munication, 1998). Tvpe locality: Baliia. Briizil ("bale de
Baliia" Kiener, 1841: 43, most probably Todos os Santos
Bay)]: Catlow and Reeve, 184.5: 292: Reeve. 1860: 52;
Reeve, 1864, pi. \'I: fig. 22 a. b; Redfiekl, 1870: 240; Tom-
lin, 1917: 275; Dance. 1976: 195 unnumbered figure.

MargineUa rargillieri. — Sowerbv, 1846: 402, pi. kxviii, figs.
178, 179. fSO; Paetel. 188.3: 31.

MargineUa IargiUierti.—?etit. 1851: 52; W'einkauff, 1879: 44,
pi. 8, figs. 2, 3: Rios and Oleiro, 1968: 17: Oliveira et al.
1981: 271.

Mnroinclln iVolutclla) laraiUieri. — .\dams and .\dams, 1853:
192: Kobelt. 1S7S: 72, pi, 27. fig, 9.

MnraincUa oiuiit Rt-eve, 1864: sp. 89. pi. XA4II, fig. 89a. b;
Redfield, 1870: 247: W'einkanff 1879; 65, pi. 12, figs. 6,
7; Paetel. 1888: 195.

MargineUa I'angiUierti. — Paetel, 1869; 38.

Closia largillcri. — Jousseaume. 1875; 255.

Closia pares Jousseaume, 1875: 2.55 (nomen novum for Mar-
gineUa ovum Reeve. 1865); Tomlin, 1917: 287.

MaroincUa (Closia) laroiUieiii. — Tnon, 1882: 47. pi, 12, figs.
'77. 78.

MargineUa l'argUlierti—?deie\. 1888: 194,

MargineUa {Closia) largiUieri. — .Vlorretes. 1949: 104.

Closia largillieri.— Rios, 1970; 114, pi. 40; Rios, 1975: 117, pi.
35, fig. 508: Wagner and Abbott. 1978: 22008.

Closia largillere- — Oliveira rt al., 1972: 14.

Persicula (Closia) largillieri. — Rios. 1985: 122, pi. 42, fig. .542.

Pcrsicnla largillieri.— heal 1990: 171, .399, 417.

Maroinella sp. 65.— Lipe, 1991: 2, pi. 1, fig. 12.

BuUata Inraillieri.—Upt.-. 1991: 2, pi. 1, fig."3: Rosenberg. 1996
[gopher://erato.acnatsci.org:70/()R1816546-181694.5-/
.wasp/, text/waspgoph .cvt ] .

Closia largiUierii—Bozzi'ti, 1992: 10,

Bnllata largiUierti —Rkk. 1994: 147, pi. 4S, fig. 6.39,

Diagnosis: Shell heavv, obovate to broadly obovate:
color pattern of small, irregular cream spots arranged in
axial rows, superimposed on light chocolate-brown back-
ground: spire completeK immersed, covered by poste-
rior labial insertion and jiosterior parietal callus: hibi;il
denticulation sharp and distinct: ventral callus heavA,
continuous; aperture narrow: first and second plication
distallv fused, forming verv strong plication ridge.

Description: Shell of medium size (14-29 mm in
length I, heavy, obovate to broadly obovate (lengtli:width
ratio 1..39-1.67, x = 1..561, s = 0.0.595, n = 25), Color
pattern of small, irregular cream spots arranged in axial
rows, on light chocolate-browii background, crossed bv'
1-3 indistinct darker spiral bands (1 near suture. 1 at
mid-section ol'bodv whorl, and 1 near anterior end, this
latter sometimes absent). Spotting somewhat to distinct-
K reduced on dark spiral bands. Spire completely im-

Page 8

THE NAUTILUS, Vol. 115. No. 1

mersed. partially to coiiipletelv covered h\ po.sterior la-
bial iii.seitioii and posterior parietal callu.s. Protoconeh
not \isil)Ie. Outer lip liea\il\ tliickeiied internalK; with
distinct, heavy external varLx, thicker medially and thin-
ner toward e.xtreniities, especially in po.sterior direction,
with indistinct, flattened, hexeled area in anterior direc-
tion, labiiil denticulation shaip and distinct, composed of
appro.ximateK 24—36 denticles, varix groove excavated.
Outer lip white, externally evenly colored a creamy
brovMiish-orange. only ver^- slightK' darker in varLx
groove adjacent to bod) whorl. Aperture narrow, nar-
rower than lip thickTiess measured ventralH'. Aperture
intemalK- pale-brown. Parietal wall convex with contin-
uous heavT ventral callus, thickening toward aperture.
abniptK' diminishing along collabral line just outside ap-
erture, forming almost planar surface tangentiiil to
whorl. Callus extending to columella, next to distal ends
of third and fourth plications. Posterior callus in fully
adult shells forming rounded mound separated from
posterior lip insertion b\' distinct gap. Left antero-ventnil
callus extending from anterior end of varfx to level of
fourth plication, with distinct furrow between second
and third plications and producing a somewhat rounded
continuation of varix. Columella concave, plications
heavy. First plication much weaker than second, distallv
fused, effectiveK' forming 1 very strong plication ridge,
separated from parietal callus b\' distinct groove. Third
plication is slightly stronger than fourth, lioth slightly
emerging from aperture. Space iiehveen plications in-
creases in posterior direction, fourth plication not re-
mote. Third and fourth plications abruptK truncated and
expanded distalK'. where the\ meet parietal callus.

Material examined: IBUFRJ 8909. Rio do Fogo
Beach. Hio Crande do Norte State. Brazil. L. Couto col.;
MNHJ 5440, off Boipeba. Baliia State. Brazil; MNRJ
HSL 4180, Barra Beach. Salvador. Bahia State. Brazil;
PMC 939, off Salvador Bahia State. Brazil. B. Linhares
col.; GAC M22I5. IBUFRJ 1469. off Itaiwpoana, E.spi'r-
ito Santo State. Brazil.

Gcojjraphic range: Rio Grande do Norte State to
northern coast of Rio de Janeiro State. Brazil (figure 13).

Kemarks: See remarks under B. ^ticninii. B. analu-
cUw, and B. hullata. and ke\' for a compari.son with B
lariiiUicii.

Bullutci largillicri is reailiK distinguished from B. lil-
acina and B. matthewsi by its distinctive color pattern,
different columellar jilications, and its more obo\ate
shape.

A juvenile specimen examined (IBUFRj 1469) shows
second plication strongly angled toward and fused witii
the first, but plications are comparatively thin and sharp.
and not yet heavily thic-kened. This juvenile shell has no
external varix but it appears that it was just i)eginning to
thicken the lip inside the aperture. Lip is thin, gradualK'
beveled, and with a shaqi edge, which is verv narrf)wK
opaque white e.xtenuilK . There is no trace of parietal
callus.

Mar^iiulla otitin Reeve. 1865. renamed Closia jxiros
Jousseaunie. 1S75. seems to represent B lar^illiiii. giv-
en mentions in the original description of its "curiously
swollen" second columellar plication, immersed .spire,
narrow aperture, and denticulated lip.

Biillatii liliiiiiiii (Sowerbv. 1846)
(Figure 5)

Marginclla lilacina Sowerbv. 1846: 402. pi. Ix,y\iii. figs.
176-177. H()lot\pe. BM{NH) 1880.9.18.2. t>pe locality-
unknown; Petit, 1851: 55: Reeve. 1865. pi. .\iv; Redfield,
1870: 240; Weinkauff. 1879: 46. pi. 4. figs. 3. 4; Paetel,
1888: 194; Matthews and Rios. 1967: 72: Kempf and Mat-
thews, 1968: 9.3; Mattliews and Kempf 1970: 5: Matthews
and Matthews, 1979: 71; Abbott and Dance. 1986; 2.34,
unnumbered figure.

Marginella (Voliitella) lilacina. — .-Vdams and .\dams, 1853: 192.

MaroincUa (Closia) lilacina. — Tr\on. 1882: 47. pi. 12. fig. SO.

Maivinclla (Vokarina) lilacina. — Morretes. 1949: 105.

Closia lilacina. — |ousseaume. 1875: 255; Rios. 1970: 114. pi.
40; Rios. 1975: 118. pi. .35. fig. 509; Wagner and Abbott,
1978: 22008; Bozzetti, 1992: 10.

Pcrsicitla lilacina. — Oliveira cf ai, 1981; 270.

Pcrsicula (Closia) lilacina.— Rios, 1985: 121. pi. 42. fig. 538;
Mello and Perrier, 1986: 1.33.

Bullata lilacina.— Upe, 1991: 2, pi. 1, fig. 1; Rios, 1994: 147, pi.
48, fig. 640; Rosenberg, 1996, [gopher;//erato.acnatsci.
org:7()/0R 1 8 1 694.5-1 81 7292-/.wasp/.te.\t/wa,spgoph.c\t].

Diagnosis: Shell broadly obcnate; color pattern of 2
pale-rose bands on ros\-browii background, outer lip
white, externalK pale- to dark-lilac; outer lip with dis-
tinct, flattened, beveled area in anterior direction,
abniptK' thinning in anterior direction forming siphonal
"gutter"; first plication somewhat higher in ventral pro-
file.

Description: Shell of medium size (19-33 mm in
length), broadK' obovate (length:width ratio 1,39-1.59, .x
= 1,.533, s = (').()573. n = 2.5). Color pattern of 2 pale-
rose medial spiral bands, and numerous, thin darker in-
distinct spiral lines, crossed bv thin, mostK' pale or oc-
casionalK darker axial lines on ros\-brown background.
Spiral lines absent on meihal spiral bands, usualK' fading
in collection specimens. A.xial lines apparently associated
with incremental growth lines. Spire immersed, mostly
co\ered b\ callus. partialK coxeretl h\ posterior lip in-
sertion and posterior pariet;i] callus. Protoconeh com-
pleteK' covered to bareK \isible. dark. Outer lip hea\ily
thickened internally, narrowing rather abruptK- in ante-
rior direction and in posterior direction, with distinct.
Hattened. bexeled area in anterior ilirection; abrupt an-
terior thiiming of lip effectiveK formiTig siphonal "gut-
ter" (best seen in apical view). Labial denticulation dis-
tinct, fine to coarse, composed of appro.ximateK- 20 den-
ticles, \'ari\ distinct, heaw. varix groove deepK excaxat-
ed. Outer lip white, extenialK- pale- to dark-lilac,
orangish-brown in varix groove adjacent to bo(K- whorl.
,\]K'rture narrow- to moderately narrow, narrower than
lip thickness when measured ventralK-. broadest at level
ol plie;itions, n;irr<)wing in anterior directioir .\pertnre

p. J. S. de-Souza and G. A. Convert. 2001

Pasre 9

iiitt-niallx lilac witli lilac stain at anterior end. Parietal
wall convex with continiioii.s, moderatel\' hea\-\' ventral
calln.s. thickening toward aperture, thinning along indis-
tinct collabral line just outside aperture, reaching fourth
plication, usualK' indistinctlv trans\erseK' undulated, ex-
tending to colinnella as a wash. Posterior callus in fulK'
adult shells forming ver\- low. rounded mound barely
separated from posterior lip insertion b\' indistinct gap.
Left antero-ventral callus extending from anterior end of
varix to level of tliird plication as heaw wash, somewhat
resembling indistinct tasciole, with lilac coloration, and
slight furrow between second and third plications. Col-
umella slightK- concave, plications heaw. First and sec-
ond columellar plications completely separate, subecjual
in size, and slightK stronger than third and fourth, first
somewhat higher in ventral profile (best seen in lateral
view). Third phcation is subequal in size to foiuth. both
slightK' emerging from aperture. Space bet^\een plica-
tions increasing in posterior direction, with fourth pli-
cation not remote. Third and fourth plications tnmcated
and slightly expanded distalK'.

Material examined: AMNH 194277. off Fortaleza,
Ceani State, Brazil; MNRJ 4175, P090 Beach, Paraiba
State, Brazil; MNRJ HSL 4176, Camocim, Ceani State.
Bnizil; PMC 476. off Fortaleza, Ceara State, Brazil; IB-
UFRJ 8924, off Rio do Fogo, Rio Grande do Norte
State. Brazil, 5-6 m, 04/1997; MZUSP 16339, off For-
taleza, Ceara State, Bnizil; MZUSP 29716, off Fortaleza,
Ceara State, P. Montouchet col. cx-piscc Aiiipliichthys
cn/ptoccutnis (Valenciennes, 1837).

Geographic range: Amapa State to C-eara State, Atol
das Rocas, Brazil (figure 13).

Remarks: Although B. lilaciiui has been reported
from coast of Bahia (Rios. 1994). all specimens collected
in Bahia tliat we obser\ed, were in fact bleached B hir-
giUieri, a species easily recognizable b\ its columellar
structure.

The color pattern of B Ulacina in most presened
shells seems to consist of onK' 2 wide whitish bands. The
e.xaniination fresh-collected shells (IBUFRJ 8924) re-
vealed thin and numerous spiral and axial lines, which
apparentK' fade with passing of time.

Biillata lilaciiia differs from B nuitthciLsi in the spire
form, and stmcture of parietal callus and columella. For
comparison with other species, see previous remarks and
kev.

Bullata iimtthcwsi (van Mol and Tursch, 1967)

(F'igure 6)

Mar^inellii (Pniniim) mntthcwsi \an Mol and Turscli, 1967:
196-197, fig. 1, holot>pe: Stanford Universit)^ Paleo-T%pc
Collection 9S56, t)pe localitv': off Fortaleza, Ceara State,
Brazil, 36 m, ex-piscc Aiuphichhjs cnjptoccntnis (Valen-
ciennes, 1837).

Marginella matthetvsi^ — Matthews and Rios. 1967: 72; Kcnipf
and Matthews. 196S: 93; Matthews. 196S: 24S; .-Vbhott and

Dance. 1986: 2.'55. unnmiihrrcil fiijiire; Matthews and
Matdiews, 1979: 71.

Pntniiiii inatllwu-si. — Wagner and .\hhott. 1978: 22009.

Bullata mnttlu-w.si.—Rios. 1970: 113. pi. 40; Rios, 1975: 117.
pi. 35, fig. 507; Rios, 1985: 121. pi. 42. fig. 538; Rios, 1990:
9, unniniibered figure; Lipe, 1991: 2, pi. 1 fig. 4; Bozzetti,
1994: 54, fig. 1; Rios, 1994: 147, pi. 48, fig. 638: Rosen-
berg, 1996, |goplier://erato.acnatsei.org:7()/()RlS17292-
1817496-/. wasp/.text/waspgoph.cvtj.

Diagnosis: Shell moderately large: color pattern of
mnnerous thin to veiT thin tawTiv- to brownish-orange
spiral bands crossed b\' mmierous axial lines on pale-
orange background; spire slightK' but chstinctK' apparent:
outer lip in anterior direction with verv distinct, flat-
tened, broadened, beveled area and distinct anterior
trough or siphonal "gutter": parietal wall with continu-
ous, thin but distinct, ventral callus; first columellar pli-
cation distinctK larger and thicker than other 3.

Description: Shell moderateK' large (39-53 nun in
length), obovate (length;width ratio 1.6.5-1.71. .x =
1.681. s = 0.0211. n = 7). with 4.5-5 whorls. Color
pattern of numerous thin to very thin tawin- to brown-
ish-orange spiral bands crossed h\ thin. mostK pale or
occasionally darker axial lines on pale-orange back-
ground. Two medial spiral bands of less dense coloration
present. Axial lines apparentK associated with incremen-
tal grov\tli lines. Spire distinctK' apparent. extremeK' low,
dome-like, partialK' covered b\' posterior lip insertion
and to a limited extent b\' posterior pariet;d callus. Pro-
toconch mamillated, completely \isible. Outer lip thick-
ened internally, with distinct external varix, rather
abniptK' narrowed in posterior direction, with \er\' dis-
tinct, flatteneil. broadened, beveled area in anterior di-
rection; abniptlv thinning in anterior direction, effec-
tively forming a distinct trough or siphonal "gutter" (best
seen in apical view), labial denticulation indistinct to dis-
tinct, fine to coarse, compo.sed of approximately 27-30
denticles; varix groove distinctly excavated. Outer lip
white, externally pale orange-cream, darkening to a p;ile
brownish-orange in varix groove adjacent to body whorl.
Aperture wide, broader than lip thickness measured ven-
trallv, curving in posterior chrection, broadest in anterior
direction. Aperture internallv nearlv white, darker band-
ing showing through ven' faintly. Parietal wall convex,
with thin, but distinct and continuous ventral callus,
forming verv indistinct collabral ridge just inside aper-
ture, extending to columella as a wash. Second indistinct
collabral callus ridge present further inside aperture and
abruptly ending just posterior to fourth plication, leaving
distinct gap behveen them. Profile of collabnd ridge
even with die tops of the plications. Posterior callus in
fully adult shells forming indistinct, low rounded mound
barely separated from posterior lip insertion by indistinct
gap, and continuing in short arc toward aperture. Left
antero-ventral callus extending from anterior end of var-
L\ to level of third plication as heavv' deposit somewhat
re.sembling indistinct fasciole. with somewhat distinct
furrow between second and third plications. Columella
nearK straight, plications strong. First columellar plica-

Page 10

THE NAUTILUS, Vol. 115, No. 1

tion distinctly larger and thicker than other three, dis-
dnctlv rai.sed in ventral profile (best seen in lateral view).
broadened, flattened, and abniptK' merging with ante-
rior edge. First and second coluinellar plications com-
pletely separate. Posterior three plications subequal in
size, all emerging from aperture to progressively greater
e.xtent in anterior direction. Space between plications
subecjual, widi fourth plication not remote. Posterior
three plications gradualK narrowing distally.

Material examined: Paratope, AMNH 134492, off
Mucuripe, Ceara State, Brazil; PMC 9.39, off Fortaleza,
Ceara State, Brazil; IBUFRJ 892.5, off Fortaleza, Ceara
State. Brazil; GAC M1325. off Fortaleza, Ceara State,
Brazil; GAC M2704, off Fortaleza, Ceara State, Brazil;
MZUSP 27308, off Fortaleza, Ceara State.

Eastern
Distribution

Geographic range:

do Norte State
13).

Maranhao State to Rio Grande
and Fernando de Noronha Island (figure

Remarks: BiiUata matthcwsi has a distincth- apparent
spire, similar to that of B ^ucninii but the\' differ in
spire height, color pattern, color patter For comparison
viith congeneric species see specific remarks above and
dichotomic key.

PHYLOGENY AND BIOGEOGRAPHY

Preliminar\' results of a cladistic anaKsis of the genus
Btillata. using conchological characters, are reported;
furdier anaKsis using anatomiciil data is intended and
will be published at the earliest opportunity.

In this analysis, 22 conchf)logical characters with a to-
tal of .53 states were used (Appendi.x). Character polar-
ization was done through outgroup comparison method
(Maddison et al, 1984). The t\pe species of Pniinim
Herrmannsen, 1850 [P. pniinim (Gmelin, 1791)] was se-
lected since this genus was proposed as the sister group
to Bullata by Coovert and C:oovert (1995: 92). The clad-
ogram was calculated using Tree Gardener 2.2 software
(Ramos, 1998) which is basically a graphic interface for
HennigS6 (Farris, 1988).

A single most parsimonious tree (figure 14) was ob-
tained (length = 37, CI = 81, RI = 69). The monophyly
of Bullata (node 1) is sustained by 7 svnapomorjihies.
Node 2, containing all species Bullata except B. huUata,
is sustiiined by 2 sviiaponiorjihies (presence of posterior
callus mound, partially thickened ventral callus, spotted
color pattern). Node .3 is sustained bv 6 svnapomorphies
(very thick external lip, distinct labial denticulation, bev-
eled area on e.xternal lip, narrow aperture, furrow on
anterior callus, fourth plication not remote). Node 3
gives origin to 2 branches, one with B. matthcwsi and
B. lilacina as terminal species, the other with B. lar^l-
lieri and B. guerrinii. Node 4 is defined bv the presence
of siphonal gutter v(>ntr;il callus completely thickened,
larger first plication, and reversion of color pattern to
white spots on a brown background. Node 5 is sustained
bv the labial coloration that extends bevond the varix

..♦*

**

#

Northern
Distribution

.r

# # y y

\^^^

v^*

/

1

' non-hiimiiplusiic

stuapomorphy
- ingroup con\ ergence

/parBlkiism

..^^^^

^
>\\^

Figure 14. Single most parsimonious plniogenetic tree ob-
tained from cladistic anaKsi.s of species of Bullata based on
shell characters. Pniniim pniniim (Marginellidae) is the out-
group. See Appendix 1 ,

groove, small first plication and an enlarged second pli-
cation fused with first one.

Our feeling before the analyses was that B Indlata and
B. matthcwsi would group as sister titxa, due to similar-
ities in size and color pattern. But the several sinapo-
moqihies of notles 2, 3, and 5, not shared bv B. bullata,
grouped B. matthcwsi as sister .species of B. lilacina.

The character optimization used in colour pattern (21)
is only one of two equally parsimonious interj^retations.
Another possibility would be that the spotted pattern ap-
peared as a parallelism between B. analuciac and node 4.

Coovert (I98(ic) proposed that the direct mode of de-
velopment within a benthic egg capsule present in the
Marginellidae would greatly reduce the dispersal abili-
ties of its species, confining the distribution of species
to a single zoogeographic province and manv with even
narrower distributions within these provinces. Scheltema
(1989) found that there is significant relationship be-
tween the lack of planktonic larvae and distribution
range. The distribution of the species of Bullata confirms
these stateTueuts.with the very narrow distribution rang-
es of B. bullata, B. analuciac, and B. gucrrinii and the
relatively wider, but still confined to parts of the Brazil-
ian coast, distributions of B. lan'illicri. B lilacina, and

p. J. S. de-Souza and G. A. Coovert. 2001

Page 11

B. nuitihcwsi. However, some capabilities tor n()n-lan.al
dispersal misiht lie present, most prohahK- b\- rafting of
egg capsules and jmeniles (Scheltema, 1989; Leal and
Bouchet, 1991), since these three last species were able
to colonize, respectiveK' the Vitoria Seamoimt (Leal,
1990), and Atol das Rocas and Fernando de Noronha
Archipelago (Matthews and Kempt, 1970). All of these
localities are separated from the continental shell bv
depths greater than 2000 ni, with Atol das Rocas and
FemancTo de Noronha, separated hv distances greater
than 300 km.

The geographic distribution of the genus along the
Brazilian coast (figure 13) suggests two major patterns
of distribution, with a slight overlap due to the extensive
ranges of B. lar<yiIUcii and B. lilacina: a northern species
group, comprising B. mattlwwsi and B. lilaciiui: and a
eastern species group, comprising B, hullata. B aiialu-
ciac, B. lanTiUicii and B. gucrriiiii. The resulting clado-
gram is congruent, to a certain e.xtent, with these geo-
graphic groups. The northern group corresponds to
notle 5. The eastern group is paraphxietic and consists
of node 4 plus B. analuciac and B. bullata.

These patterns appear to follow two major oceano-
graphic current patterns present along the Brazilian
coast; the northern group, associated with the Gu\ana
cmrent and the eastern group, associated with the Brazil
current. It is possible that the divergent currents along
with intracapsular mode of development niav have con-
trilnited to isolate northern and eastern populations, fa-
cilitating speciation events.

Kev to species of Bullata:

la. Shell with distinct cream spots on caramel- or
dark-browii background and with taint spiral
banding 2

lb. Shell lacking cream spots, color pattern of tew-
to man\' bands 4

2a. First and second columellar plications complete-
Iv separate, subecjual in size; parietal callus re-
stricted to posterior end, not forming collabral
ridge: aperture much broader than lip thick-ness;
fourth plication remote (i.e. distance between
third and fourth plication distinctly greater than
distance between second and third); lip lacking

denticulation, moderately thickened

Bullata aualuciac new species

2b. Second columellar plication greatly enlarged,
fused with first; parietal callus moilerate to
heavy, forming weak to distinct collabral ridge;
fourth plication not remote 3

3a. Spire completeK' immersed, usually covered by
posterior labial insertion; aperture narrower tlian
lip thickiiess, onl\' shghtK- broader in anterior di-
rection; parietal and other ventral callus white or
nearlv so; lip greatlv thickened; third and fourth
plications distincth' tmncate and expanded at
distal ends Bullata laroillicii (Kiener, 1841)

3b. Spire not immersed, slightly apparent, proto-

conch exposed; aperture as broad or broader
than lip thickness medialK', distinc-tixeK broader
ill anterior direction; parietal and other \entnJ
callusing brown-tinged; lip nioderateK' thick-
ened; third and fourth plications gradually nar-
rowing distalK- Bullata jificninii new species

4a. Shell smaller, 19-33 nun in length; color ros\-
browii with 2 distinct light pink, broad bands; lip
strongly denticulate; aperture narrower than lip
thickness; third and fourth plications distinctly
tnmcate at distal ends; lip strongK' denticulate,
greatK thickened; color pattern of 1 or 2 broad

spiral color bands

Bullata lilacina (Sowerby, 1846)

4b. Shell larger, 34-98 mm in length; color gra\ish-
to orange-brown, v\ithout distinct broad spiral
bantls, with numerous distinct thin lines of varx-
ing thickness; lip at most weakK denticulate; ap-
erture nmch broader than lip thickness; third
and fourth plications gradualK' narrowing distal-
K', ending in sharji points; color pattern of nu-
merous, small spiral bands 5

5a. Lip weakly crenulated, with a flattened, broad-
ened area in anterior direction, forming a weak
siphonal "gutter"; spire slightK but distincth' ap-
parent; first plication nnicli stronger than other
plications and distincth' higher in profile; fourth
plication not remote, distance between third and
fourth at most only slightK greater than distance

between second and third

. . . Bullata inattluusi (van Mol and Tursch. 1967)

5I3. Lip at most faintly crenulated, lacking flattened
area in anterior direction; anterior 2 columellar
plications subeijual in size; spire immersed, apex
flattened to depressed, usualK' covered; first pli-
cation as strong as or weaker than other plica-
tions, not higher in profile; fourth plication re-
niote Bullata hullata (Bom. 1778)

ACKNOWLEDGM ENTS

We are verv' grateful to Paulo Marcio S. Costa who first
drew my attention to B. analuciac and provided many
specimens herein stucUed. To Dr Ricardo S. Absalao,
Dr. Eliezer de C. Rios, Luiz R. Simone, Dr. .\lbert D.
Ditchfield, Dr. Ricardo Pinto da Rocha, Dr M. G. Har-
asewYch, and an anonvinous reviewer: we are tli;uik-f\il for
tlieir comments and criticism. We are deeply indebted to
Jose Coltro and Marcus \inicius C'oltro who generously
provided several specimens herein studied and biblio-
graphic;il help. Our most sincere gratitude goes to Dr GiU^'
Rosenberg (ANSP), Dr. Paula Mikkelsen (AMNH), Dr.
M. G, Harasewvch and Ms. Rave Germon (Nation;il Mu-
seum of Natural llistoiv), who received the senior au-
thor in their institutions. The types oi Marginclla cuiicii
and information on the types B. largillicri were kindly
provided bv Dr Pliillipe Bouchet. This work was funded
by Fundavao Universitaria Jose Bonifacio, Universidade
Federal do Rio de Janeiro, the Academy of Naturid Sci-

Page 12

THE NAUTILUS, Vol. 115, No. 1

ences of Pliiladelpliia ("Je.ssup Award"), and American
Museum of Natural Historv. Senior author i.s supported
by doctoral grant FAPESP'97/11429-.3, Brazil.

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Page 14 THE NAUTILUS. Vol. 115. No. 1

.\ppcndi.\ 1. Character list ami tahle.

Character 1: Spire shape, nonaililitive. 4 steps. CI = 75, RI = 0

[0] — .Apparent, conic. \erv low; [1] — Apparent dome-like; [2] — Involute flat; [3| — hnohitc. sunk.

C^haracter 2: Posterior lahial in.sertion. nonaclditive. 3 steps, CI = 66. RI = 0

[()] — on bodv whorl; [1] — posterior to bod\' whorl; [2] — co\ering ape.x.

C;haracter 3:' \ari,\. nonadiliti\e. 1 step. Cl' = 100. RI = 100

[0] — narrow; [I] — wide.

Character 4: Lip tliickness. 1 step, CI = 100, RI = 100

[0]— thick; [1]— ven^ thick;

Character 5: Labial denticulation. nonadditive. 2 steps, CI = 100, RI = 100

[0] — absent; [I] — crenulated; [2] — distinct;

Character 6; Labial coloration, nonadditive, 2 steps, CI = 100, RI = 100

[0] — restrict to sarix groo\'e; [I] — extending ventrallv; [2] — 3 colors;

Character 7: .\nterior beveled area on external lip, 1 step, CI = )00. RI = 100

[0] — absent; [I] — present;

Character 8: Siphonal gutter, I step, CI = 100, RI 100

[0] — absent; [1] — present;

Character 9: Aperture width, I step, CI = 100, RI = 100

[0] — wide;[l] — narrow;

Character 10; \'entral callus, 1 step, Cl = 100, RI = 100

[0] — absent; [1] — present verv sinuous;

Character 11: Callus thickness, additive, 3 steps, CI = 66, RI = 0

[0]— thin; [1]— thick locallv; [2]— thick all over;

Character 12; Posterior cailus mound, 1 step, CI = 100, RI = 100

[0] — absent; [I] — present;

Character 13: Anterior callus, 1 .step, CI = 100, RI = 100

[0] — simple; [1] — fasciole Hke;

Character 14: Furrow on anterior callus between second and third plications, 1 step, CI = 100, RI = 100

[0] — absent; [1] — present;

Character 15: first plication size, nonadditive, 2 steps, CI = 100, RI = 100

[0] — subequal; [1] — small; [2] — large;

Character 16: first plication in ventral profile, 1 step. CI = 100. RI = 100

[0] — normal; [1] — raised;

Character 17: second plication. 1 step. CI = 100, RI = 100

[0] — subequal; [1] — larger fused with first;

Character IS: fourth plication. I step. CI = 100. RI = 100

[0] — remote; [1] — not remote;

Character 19: False fifth plication, 1 step. CI = 100. RI = 100

[O] — present; [1] — absent;

Character 20: Shape of distal end of plications. 2 steps. CI = 50. RI = 0

[0] — thin evenly; [I] — tnmcate;

Character 21: Color pattern, nonadditive. 4 .steps. CI = .50. RI = 0

[0] — few wide bands; [1] — many thin wide hands crossed bv axial lines; [2] — spotted on wiik' bands;

Character 22: Size, 2 steps, CI = 50. RI = 100.

[0] — small to medium; [1] — large;

THE NAUTILUS 115(1): 15-21, 2001

Page 15

Reproduction and brooding of Crepidiila argentina, Simone,
Pastorino and Penchaszadeh, 2000 (Gastropoda: Calyptraeidae)

Maxiniiliano Cledon

Facultad de Ciencias Exactas v

Naturales— UBA
Departamento de Biologia
C142.S EHA Buenos Aires
ARGENTINA

Pablo E. Penchaszadeh

Museo Argentine de Ciencias

Naturales— CONICET— UBA
A\'. Angel Gallardo 470
C1405 DJR Buenos Aires
ARGENTINA

ABSTRACT

Crepidiila araentina is a protandrie hermaphrodite diat under-
goes complete sex change. Males develop a mature gonad
when the shell length is 4 mm. Sex change begins when shell
lengdi is around S mm and continues until II mm, when the
first pre-vitellogenic oocytes are found. The minimum and
maximum shell lengths for a brooding female are respectively
1.5 and .36 mm. There is a well-defined seasonal reproductive
cvcle; females brood between August and April. Females brood
between 10 and 46 (mean 2.5) egg capsules per egg mass. The
intracapsular fluid is translucent and viscous in recently laid
capsules but becomes less dense closer to hatching. Neither
nurse eggs nor cannibalism have been observed. The mean
number of eggs per mass was 5600 ± .3.300. The egg capsules
have a mean width of 2.1 mm (SD = 0.03 mm, N = 52), and
a mean length of 2.4 mm (SD = 0.04 mm. N = 52). Each egg
capsule contains about 320 embn.os. The vmcleaved egg di-
ameter is about 170 (j.m. .\\\ eggs develop suichronouslv in the
same brood. The embryos hatch as planktotrophic \eligers with
a shell length of 190-2.30 fxm and a bilobed velum about 160
(xm in width. The mean size of the egg capsules, number of
embryos per egg capsule, and number of embryos per brood
are positiveK' correlated with female shell length.

Additiuiwl key word.'i: protandry, reproductive cycle, season-
ality, encapsulation patterns, southwest Atlantic Ocean.

INTRODUCTION

Protandry is the nicxst common kind of liermaphroclitism
among gastropods (Coe, 19.38a). Protanch-y could he fa-
vored when female egg production increase.s with female
size, y\'hile male reproductive success is independent of
size (Collin, 1995). There is not an optimal size for sex
change below which all the indiyiduals are males and
above yvhich the individuals are females; instead, there
is a range of sizes in which se.xes overlap (Coe, 19.3Sa;
Heller 1993; Collin, 1995).

In general, caKptraeids have a protandrie reproduc-
tive cycle uith a similar sequence of sexual phases (Kii-
kenthal, 1925). Youngest indiyiduals are males and are
usualK' not larger than few millimeters. The oldest in-

dividuals are larger than a centimeter. The\- become fe-
males after a transition period that depends on the spe-
cies. Cal\ptraeids are characterized h\ the production of
eggs that are brooded in bunches of stalked egg capsules
attached either to the foot or to the substratum under
the female. Each egg capsule has a thin membrane and
a peduncle, which fixes it to the substratum. Hatching
of planktonic veligers takes place in some species and of
crawling juveniles in others (Hoagland, 19S6).

TraditionalK' the taxonom\- of species of the genus
Crepidula was based e.xclusivelv on shell characters (i.e.,
Parodiz, 1939; Hoagland. 1977). The substratum has a
great influence on the shell form of gastropods of sessile
habits, making shell-based taxonom\' difficult. Hence it
becomes important to include other traits, such as ana-
tomical and reproductive characters, in species charac-
terizations.

The type of embr\ onic development is species-specif-
ic (Hoagland and Ponder, 1998). In some species few
eggs develop and the rest of them remains uncleaved,
becoming nurse eggs: this is the case of C. dilatata La-
marck, 1822 (Gallardo, 1979; Chaparro and Paschke,
1990) and C. philippiana Gallardo, 1977 (Gallardo,
1996), all from Chile. In the latter species, onl\- one
crawling juvenile hatches from each egg capsule. Egg
capsules in other species contain only cle\eloping em-
brvos immersed in a fluid (Hoagland, 1986).

Crepidula ar^eiitinti Simone, Pastorino and Penchasz-
adeh, 2000. was described as a nev\^ species from mate-
rial obtained for the present study. In it, we characterize
the seasonalits' of reproduction, the relationship between
,sex change and size, minimum size at brooding, number
of egg capsules per brood, the number of embr\()S per
egg capsule, mean egg capsule size, uncleaved egg di-
ameter, hatching stage, and the presence or absence of
nurse eggs in C. arocntina.

MATERIAL AND METHODS

Characterization of sexual stages according to the
size: .\roimd 25 indi\iduals per mouth were collected.

Page 16

THE NAUTILUS, Vol. 115, No. 1

The.se were found attached to shells of the mussel A/i/-
tiliis ('(lulls platciisis d'(lrl)i<jn\. lS4fi, from material
landed In' commercial fisheries at Mar del Plata harbor
The mussel banks are distributed in a patchy fashion on
tlie continental shelf oif the Province of Buenos Aires,
from Querandi lighthouse to Necochea, between 35 and
50 m depth on mixed bottoms of sand and shell debris
(Penchaszadeh, 1974).

The shell length of each individual was measured
within 0.1 mm precision. The soft parts were preserved
in Bouin's solution.

To studv gonad development, 30 individuals were col-
lected in Januarv 1999 with size intervals of appro.xi-
mately 1 mm, and their gonads sectioned and stained
with haemato-wlin-eosin.

To confirm results from the macroscopic observation
of gonads, the reproductive stage (as the proportion of
sexual cells in different stages of development), or de-
gree of gonad development, was examined through his-
tological sections each month in ten females of more
dian 20 mm shell lengtli.

Characterization of the reproductive .season: The

minimum shell length at which individuals brood was
used as a minimum threshold for female reproductive
capacitv. A count of the number of broods per sample
gave the monthly proportion of total brooding females.
These data were contrasted with published bottom tem-
perature data for the mussel beds area (40-.50 ni; 37°-
3S°30' S, 56°-57°50' W) (Bamirez et al. 1973; Carreto
et al. 1998).

Characterization of broods and larvae: The egg

capsules were fixed in 4% formalin in seawater and pre-
sened in 70% alcohol. A total of 52 broods was studied.
From each egg mass, 5 egg capsules were randomlv se-
lected for length and width measurements under a Zeiss
stereoscopic microscope with a 0. 1 nun precision ocular
micrometer The number of embrvos was counted and
the mean and standard deviation were calculated for
each variable. A Kniskal-Wallis test was carried out be-
tween initial number of eggs per capsule and the num-
ber of pre-hatchlings per capsule in order to test the
absence of nurse eggs or camiibalism.

We measured the diameter of 10 eggs per egg cap-
sule, from 5 egg masses found with uncleaved eggs.

Simple linear regressions tvpe 2 following natural log-
arithmic (In) transformations were used to test the pos-
sible correlations between female shell length and the
number of egg capsules per brood, the number of em-
bryos per egg capsule, the mean egg capsule length and
width.

In addition to the preserved material we collected sev-
en egg masses to describe the hatching stage. These
were kept in a non-circulating aerated artificial marine
water acjuarium with '-W/cr of salinitv' and temperature
between 20 and 22°C;. Hatchlings were measured with
a digital video svstem attaclied to a Zeiss microscope.

The shell length at mctamoiphosis (protocouch) of

voung male intli\iduals 4-6 mm length was measured
using SEM.

RESULTS

Reproductive stages and shell length: .Males are al-
readv fimctional when they attain approximatelv 4 mm
in length. At this stage thev have sexual cells at different
degrees of development in the same spermatic follicle

(figures 1-2).

Spermatogenesis diminishes when the individual
reaches 8 mm in shell length and intrafollicular absorp-
tion begins at this stage (figure 3). The spermatozoids
Tuove to the seminal vesicle and the remaining spermatic
cells sutler cvtolvsis and orange-vellow lipid vesicles appear

The penis is located behind the right tentacle and is
characterized bv a filament on the distal tip; sperm
groove was not observed (figure 4). Generallv the penis
gets reabsorbed simultaneously with the male gonadal
structures. This process is usually completed when the
size of the individual reaches 11 mm shell length. Nev-
ertheless, we observed a case of an 11.8 mm length in-
dividual that retained a well-developed penis (figure 5)
but in which liistological sections revealed a female go-
nad with vitellogenic oocytes (figure 6).

The female genital papilla is generallv visible in indi-
viduals of more than 10.3 mm in shell length.

Vitellogenesis begins at 11 mm shell length but fe-
males are not reproducing until thev reach 14.5 mm.
When they reach this size the animal holds a stock of
mature oocvtes and is able to brood. This was corrobo-
rated through histological sections of females collected
monthlv. Oocytes at different stages of maturation were
found in all cases (figure 7).

After laving an egg mass, a stock of late vitellogenic
oocvtes is retained in the follicles, and re-absorption of
oocvtes was not tletected.

The largest brooding female was 36 mm shell length
and the largest female 38.5 mm; both contained late vi-
tellogenic oocvtes.

Reproductive season, broods, larvae, and .size at
settlement: The studv of 52 broods showed that each
egg mass consists of 10-46 egg capsules (mean = 25,
SD = 9, N = 52).

The' reproductive season nms from .\ugust to April.
Small females (14-20 mm) lav egg capsules onlv be-
tween September and Febniarv, while larger females re-
produce during the entire reproductive .sea.son (figure 8).

The egg capsules are trianguhu' with roimded edges,
transparent, colorless, and bilateralK' compressed (figme9).
In livv egg capsules, the capsule w;ills ;ue extremelv deli-
cate and often break when handled. The walls have a trans-
piirent double uKMiibrane and an attachment pedimde in
their proximal e.xtremitv. The capsules mea.sure 2.4 umi
width (SI) = 0.4 mm, N = 52) and 2.1 unn length (SI)
= 0.3 mm, N = 52), excluding die peduncle.

There is positive correlation between tlu- In-trans-
formcd egg (Mpsiilc sizi' (width and length* and the In-

M. Cledon and P. E. Penchaszadeh. 2001

Page 11

• ■*

' • et

..^

Figures 4-5. Cn-piihdii (ir;j^iiitiiiii 1. DrI.iil of tin- distal cud
of tln' penis (pe) of a 4.7 iiiiii male. Scale bar = 0.5 mm. 5.
View of the penis (pe) of an indi\idua! of 11.8 mm whose
t;onad has developed oocvles. Scale bar = 2 mm.

transloniiet! length of the female (r = 0.55, r- = 0.31,
F = 21.9 and r' = OSS, r = 0..34, F = 25.79, respec-
tively, with p < 0,0001, ni = 0.45 and 0.42 respectivelv
N = 52) (figures 10-11).

There is also a positi\e (.omlation hehveeii In-trans-
formed egg capsules length antl width (r = 0.81, r- =
0.66, p <'0,0()01, m = O.Vl, F = 96.45, N = 52) (figure
12). The mean number of embr\'os per egg capsule is

323 (81-44S, SD = 76, N..,,,,„., = 52, N,,.,,,, ,„,„„,,., = 208).

The lii-transforined mi-an minilicr of (■m])r\()s per egg

Figures 1-3. Light micrographs of Crepidiild (ir^ciitiiia tes-
tis. 1. Spermatic follicle of a 4.7 mm male, containing cells in
different degrees of development, eg: spennatogonial cell, ct:
spermatocites, ed: speniiatide. Scale bar = SO (xm. 2. View of
a spermatic follicle of a 10.4 mm male containing a large pro-

portion of sperniatozoiils (e/). Scale bar = 100 (iml .'J. View
of a 7.7 mm male gonad in re-absoiptioii ])liase (rel, containing
late spermatogenesis cells. Scale bar = 220 (xm.

Page 18

THE NAUTILUS, Vol. 115, No. 1

V^i

.>*i^.

r?^*;

d

Figures 6-7. Crepidula argcntina. 6. Developing female gonad of an II. S inm long indmdual. Scale Ixii- = 200 |ji.in. 7. Oocytes in
different degree of de\elopnient, previtellogenic {p\') and \itellogemc (vt) in a female of 17.6 mm of sheU lengdi. Scale biir = 70 (xm.

capsule is also correlated with female length (r = 0,45.
r= = 0.2, p < 0.01, m = 0.78, F = 12^:87, N = 52)
(figure 13). The relation between In-transformed egg
capsule size and In-transformed mean number of em-
brv'os contained was also studied (with egg capsule
width: r = 0.62, r- = 0.39, p < 0.000 m = 0.13, F =
32.14; with egg capsule length: r = 0.66. r- = 0.44, p <
0.0001, m = 1.57, F = 39.12, N = 52)

The In-transformed mean number of embrvos per
brood is 5600 (SD = 3300, N = 51) and is correlated
with the In-transformed female shell length (r = 0.37,
r- = 0.14, p < 0.005, m = 0.46, F = 1.34). No corre-
lation was found betv\'een the In-transformed female
shell length and the In-transfornied number of capsules
per brood. As the F value intlicates, ;ill slopes of the
regressions were different from zero.

S? 50

?■ 30

S 10

5 6
Month

8 9 10 II

Figure 8. Frequency ol brooding females of Civpiduhi iir-
l^ciitiiui from Januan.' 1999 to November 1999. The bars rep-
resent the percentage of brooding females from the tot.il in-
spected females vvithin each month. Small females (14-20 mm
in shell length) are represented in white, large females (more
than 20 nun) are represented in black.

The imcleaved eggs were whitish and approximately
spherical with a mean diameter of 170 |jLm (SD = 9, N
= 250). No differences where observed among the five
mothers.

Cleavage follows die holoblastic pattern. First and sec-
ond cleavages are similar, giving rise to four macromeres.
All eggs develop. No cannibalism or nurse eggs were ob-
served (Kniskal-Wallis, p = 0.27). The development of all
embnos in each brood occurs SMichronousK'.

The intracapsular liquid is initial!) viscous, but be-
comes less dense toward hatching.

The embno hatches as a feeding \eliger larva. The
globular smooth lar\al shell at hatching measured from
live and presened material was 190-230 |jLm in length
and 195 |xm in width (N = 112).

Of the seven egg masses maintained in the aquarium,
onK- one containing late veligers at the time of collection
hatched. The colorless, transparent, bilobed \elum had
a maximum width of 164 |a.m. Each o\al lobe measured
151 (jLm maximum length. The longest cilia in the pre-

Figure 9. Xicw of an egg capsule of Crepidula ar^cntina
conlaiiiMig imcleawd eggs. Scale bar = 2 mm.

M. Cledon and P. E. Pencliaszadeh, 2001

Page 19

11

• • * . •

7.5 7,7

In egg capsule length

7,9

13

.* R

O 3

I"

.•n»'--r/j«*^.'

7.4 7,6 7,8

In egg capsule length

2,9 3,1 3,3 3,5

In female's shell length

3,7

Figures 10-13. Crepidula argentina 10. Relation between In female shell length and In egg capsule length. \ = .52. 11. Relation
between In female shell length and In egg capsule width. N = .52. 12. Relation between In egg capsule length and width. N = 52.
13. Relation between In female shell length and In number of enibnos per egg capsule. \ = 52, Regression anal\'sis \'alues and
their significance are gi\'en in the text.

oral cilian- band were 32 jjim long. On the anterior part
of the velum there is a pair of ciliated tentacles. The
triangular ciliated foot possesses an oval operculum,
which reaches 67 \x.m in length and 90 |xm in width.

The protoconchs of indi\iduals of 4—6 mm length
measured 6.31 |xm mean length (SD = 25, N = 34)
(figures 14-1.5).

DISCUSSION

Crepidula argentina is a protandric hemiaphroditic species
with complete sex change. The males are already mature
at 4 mm shell length. Sex change begins at appro.ximately
9 mm and is completed h\ 11 mm. E\idence of c\toKsis
was obsened during die w hole process of sex change. Fur-
dieniiore, a full\' developed fem;de gonad can be present
before die penis is absorbed and the genital papilla de\el-
oped; such sex change has also been obser\ed in Cnici-
btilum spimmim Sowerb\-, 1824 (Coe, 1938b). At the end
of testiculiir re-absorption die penis disappe;u's, and the
oviduct develops (Grimpe and Wegler, 1940).

Crepidula ar^^entina does not form stacks as does C
fornicata Linnaeus, 1758. In this latter species, individ-
uals are strongly influenced bv each other in their sexual
expression (Orton, 1909).

In C. arscniina a number of mature oocytes is main-

tained in the gonail even after laving. This suggests that
females produce more than one brood per year. This
possibility could depend on the duration of brooding.

As in C. convexa Say, 1822 (Bandel. 1976), positive
correlations were detected in C ar<icntina between (e-
niale shell length and egg capsule size, mean number of
embryos per capsule, and mean capsule length and
width: bigger females brood more eggs because the egg
capsules are bigger and contain more eggs. However, no
correladon was observ-ed betw-een female size and the
number of egg capsules per brood.

The South American species C. convexa and C plii-
lippiana do not show a reproductive resting period ( Ban-
del. 1975; 1976; Gallardo, 1996) while C. argentina rests
from May to July. Resting periods are also observed in
the South .American C dilatata Lamarck, 1822, and C
fecunda C;dlard(), 1979 iCallardo. 1979). This fact would
be related to seasonal water temperature changes. Tem-
perature data in Ramirez et al. (1973) and Carreto et al.
(1998) indicate that i)ottom temperature at 50 m (where
mussel beds with C argentina are located) follows a sea-
sonal cvcle with the maximiun between Februarv' and
April (17.5°C) and the minimum between July and Sep-
tember (9.5°C). Crepidula argentina does not reproduce
in the decreasing temperature periixl, between Mav' and

Page 20

THE NAUTILUS, Vol. 115, No. 1

Table 1. Cluiraeleristies of the

n-prot:

liietion for different Crepidtila species from South America.

Brooding

.Nnnilu

r ol

.Nuiniier (li

.Male shell

teniale shell

egg capsnies

eggs per

Untleaxed egg

Nurse

H;it(liing

length

length

Species

Reference

per egg

mass

egg capsule

diameter (p.mt

eggs

stage

(mm)

(mm)

c

araentiiin

This stndy

10-46
( mean
SD =

= 25,
9)

81-t48
(mean =
.323. SD:

Mean = 170,
SD = 9

No

Xi'liger lanac;
shell length:
1 90-2.30 Vm

.3.95-8.85

14.5-36

c.

proteti

lloagland.

26-18

76); all eggs
dexelop.
33-120

Not consigned

No

\fligrr lanae;

4.(i-8

7.3-20

1983

(nieiui

=

(mean = 61,

(embnos ol

(size not

31.6. SD =
1.1)

SE = 8); all
eggs devel-

150)

consigned)

c

diUitata

Gallardo,
1977' Cha-
parro and
Piischke,
1990-

22-29

op.
308-1016; ap-
pr(ttimatel\
1.5-50 em-
bnos devel-
op.

195-263

Yes

Crawling juve-
nile; shell
length: 900-
1300 iJini'.
1075-1600

6-26

12-.53

c.

fi'cnndn

CJallardo,

1977

Mean = 542;
all eggs de-
velop.

204-238
(mean =
212)

No

jim-
N'eliger larvae;
shell length:
.500 jiin '

13-i9

31-72

c

phihppUiiui

Ciallardo,
1977; 1996

16-34

145-131; 1 em-
brvo devel-
ops.

144-161

Yes

(a'awling jn\e-
nile; shell
length 2800-
31()0 |jLm

(i-11

14 and up

Julv (winter) and present,s its maximum reproductive ac-
tivitv between Febniarv and March (summer).

Most .species of die genus v\idi hirge females la\' eggs
widi small diameters (Hoaghmd and Coe, 1982) and this
is the case (jf C on^cntina. The iincleaved egg diameter is
170 |i,m widi ven, litde viiriation. There are odier species
in the genus with a similar egg diameter, like C. cciifhicola
C. B. Adams, 1852, C.foniicata, C. echinus Broderip, 1834
(Hoagland, 1986) ;ind C. Uuffthta Gould, 1846 (Collin,
2(MK)), Crcpklula ar<icntiiw differs from the two first .spe-
cies respectivel}' in tliat it lacks nurse eggs and produces a
different number of egg capsules and it also differs in fe-
male size from these two species. Crcpklula ai'fs^cntina h;is
a me;in number of emlinos per capsule (323) e(jui\;ilent
to C. echinus, a species in which also all eggs develop
(HoagliUid, 1986). The absence of nurse eggs or cannib;!]-
ism among sibling embryos accounts for tlie lack of mtirked
variation in shell lengdi at hatching. Embrvoiiic cannibal-
ism occurs in some otlier species of the genus but was not
observ'ed in C. argentina.

The intracapsular liquid is translucent and viscous in
recently laid capsules, and becomes less den.se as hatch-
ing approaches. Because there are no nurse eggs, intra-
capsular fluid is the oiiK possible extra embryonic food
supply. In C. protea a stick)' matrix in which embryos
were embedded, making the embrsos difficult to remove
from the capsules, has been reported (Hoagland, 1983);
this would distinguish C. proteti from all other Crcpiduhi
species, including C. argentina.

In C. argentina the mean number of egg capsules per
brood (25) is similar to those of other congeneric spe-

cies, such as C. plana Say 1822 (Bandel, 1976), C. cer-
ithicola and C. echinus (Hoagland, 1986).

Crepidula argentina is different from C. protea
d'Orbignv, 1841, as described by Hoagland (1983) in al-
most all reproductive aspects examined (table 1).

The individual shell length at metamoqihosis in C. ar-
gentina is ;u'ound 6.30 |jLm in length with reniiirkablv little
variation among individtuils (SD = 25), wluch shows a totid
growth of iiiound 400 |jLm during planktonic life.

( Comparing Crcpklula argentina with other southern
Soudi .'\merican species: C. dilatata Lamarck. 1822 (C;il-
lardo 1977a) has direct development witli nurse eggs. Un-
like C. argentina. the reproductive period is froin Mav to
[anuarv with a maximum in Julv. In C dilatata hatching
occurs at the juvenile stage. Males and females have Uu-ger
shell length ranges tlian C. argentina (table 1). Crepidula
fccunda (Gallardo, 1979), produces more embryos per
brood, its eggs are hirger and shell lengtlis of males and
fem;iles iilso exceed adult sizes of C. argentina (table 1).
Crepidula philippiana is easilv distinguishetl from C. ar-
gentina bv presence of nurse eggs; die egg tbameter fluc-
tuates between 140—160 fim and onK* one laige juvenile
hatches from each capsule (Gallarck) 19771)).

ACKNOWLEDGMENTS

We greatlv appreciate the manv valuable suggestions
jirovidi'd bv Hichard Strathmann to an earlier version of
the manuscript and bv three anonvinous reviewers. We
are also indelited to Guido Pastorino for liis help in
inonnting the jihotographs.

M. Cledon and P. E. Penchaszadeh, 2001

Page 21

Figures 14-15. Cnyidula aroentina. Scaiuiiiii; electron micro-
giaphs of the protoconch on 4-6 mm males. Scide bur = SO |j,ni.

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Bandel, K.1976. Observations on spawn, embnonic dewlop-
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Carreto, J. I., R. Akselman, N. G. Montova, R. M. Negri. H.
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Verona and ]. I. Carreto (eds.) Xmita de Galicia and In-
tergovemamental Oceanographic Comision ot UNESCO,
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Chaparro, O. R. and K. A. Paschke. 1990. Nurse egg feeding
and energv- balance in embrv'os oi Crcpidiiln dilntata (Gas-
tropoda; CaKptraeidae) during intracapsular develop-
ment. Marine Ecology' Progress Series. 65:18.3-191.

Coe, W. R. 19.38a. Influence of association on the sexual phases
of gastropods having protandric consecutive sexualitv Bi-
ological Bulletin 75:274-2.85.

Coe. W. R. 193.Sb. Sexual phases in the gastropod Cnicihidiiin
spinosiiin. Journal of Moqshologv 62:34.5-361.

Collin. R. 1995. Sex, size ,ind jKisitjon: a test of models pre-
dicting size at sex change in the protandrous gastropod
Crepidulu fomiattn. The .'\merican NatunJist 146:815-8.31.

Collin, R. 2000. Sex (Change, reproduction, and development
of Crepidula (tctunrii and Crrpidula liii'^uhitd (Gaslrojioila:
CaKptraeidae). The Wligcr 4.3:24-3.3.

Callardo, C. 1977a. Two modes ol development m the mor-
phospecies Crcpiduld diUitatci (C;astro]ioda: CaKptraei-
dae) from Southern Chile. Marine Biologv .39:241-251.

(Jallardo, C. 1977b. Crcpidida philippiatw n. sp., nuevo gas-
tropodo CaKptraeidae de Chile con especial referenda al
patron de desarrollo. Studies on Neotropical Fauna and
Environment 12:177-185.

(Callardo, C. 1979. Especies genielas del genero (.'.lepidida
(Gastropoda, CaKptraeidae) en la costa de (;hile; una re-
descripcion de C dilatiita Lamarck v descripcion de C.
fccunda n. sp. Studies on Neotrojiical Fauna ,ind F.nvi-
ronment 14:21.5-226.

Callardo, C. 1996. Repioduction in Cnpi<liil(i jihilippiana
(Gastropoda, CaKptraeidae) from Southern Chile. Studies
on Neotropical Famia and Environment .31:1-6.

Grmpe G. und E. W'agler. 1940. Die Tiervvelt der Nord- und
Ostsee Band IX bl. Akademische \'erlag.sgesellschaft
Becker and Erier Kom.-Ges. Leip/ng, 147-157.

Heller, J. 1993. Hermaphroditism in molhiscs. Biological Jour-
nal of the Linnean Societ)- 48:19—42.

Hoagland, K. E. 1977. Systematic review of fossil and recent
Crcpidida and discussion of evolution of the CaKptraei-
dae. Malacologia 16:35.3-420.

Hoagland, K. E. and VV. R. Coe. 1982. LanJ development in
Crcpidida maculosa (Prosobranchia: (^repidulidae) Irom
Flonda. The Nautilus 96:122,

Hoagland, K. E. 1983. Ecology and larval development ot Crc-
pidida protca (Prosobranchia: Crepidulidae) troni South-
ern Briizil: a new tvpe of egg capsule lor the genus. The
Nautilus 97:10.5-109.

Hoagland, K. E. 1986. Patterns of encapsulation and brooding
in the Calvptraeidae (Prosobranchia: Gastropoda) Ameri-
can Malacological Bulletin 4:17.3-183.

Hoagland, K. E. and W. Ponder. 1998. Superfaniily Calvptraeo-
"idea 772-774 in Beesley, P L., Ross, G. J. B. an'd Wells,
A. (editors) Mollusca: the Southern Synthesis. Fauna of
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56.5-12.34.

Kiikenthal, W. 1925. Ilandbuch der Zoologie; 5 Band. 6,3-(i7.
Walter de Cruvter and Co., Berlin und Leipzig.

Orton, J. H. 1909, On the occurrence of protandric hermaph-
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46,8-484. '

Parodiz, J, J. 19.39. Las especies de Crcpidida de las costas
argentinas. Physis 17:68.5-709.

Penchaszadeh, P. E. 1974. Ecologi'a del mejillon Mytilus pla-
tcnsis d'Orbignv de bancos circalitorales de la Provincia
de Buenos Aires (Argentina). Tests doctoral, Universidad
de Buenos .\ircs. 1<89 pp.

Ramirez, F, B. H. Roa, C. A. Wron.i and J. I Carreto. 1973.
Plancton v condiciones ccologiciLS en las aguas de la pla-
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Simone" L. R. L., G. Pastorino .md P E. Penchaszadeh. 2000.
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127-141

THE NAUTILUS 115(l):22-28. 2001

Page 22

Revision of the Architectonicidae ( Gastropoda: Allogastropoda)
from the Miocene Chipola Formation, Oak Grove Sand, and
Shoal River Formation of northern Florida, with descriptions of
four new species

G. W. Schmelz

5575 12th Avenue Southwest
Naples. Florida, 34116 USA

ABSTRACT

A revision of the Architectonicidae from the Miocene Chipola
Formation, Oak Grove Sand, and Shoal River Formation of
northern Florida is presented, along with descriptions of four
new species. Two new species are assigned to the genus Helia-
cus, one to the genus Architectonica and a fourth to the genus
Granosolarium.

Additional key words: Architectonica. Heliacus. Granosolar-
ium. Tertian', Neogene, fossil mollusks.

INTRODUCTION

Of the estimated 1100 .species of mollusks believed to
be represented in northern Florida's Miocene Chipola
Formation and Oak Grove Sand deposits onK' about 640
have been described (Yokes, 1989; 1997). Currently, be-
tween 120 and 130 unde.scribed species of mollusks from
the (^liipola Formation, Oak Grove Sand, and Shoal Riv-
er Formation have been (hscovered !))■ tlie antlior in the
Invertebrate Paleontolog)' collections at the Florida Mu-
seum of Natural History' or in private collections. In-
cluded among those species that remain to be described
are a broad range of specimens representing man\ dif-
ferent classes and families. This paper is intended to
examine four of these new species in the family Archi-
tectonicidae.

Bieler (1993) recognized about 140 living species of
Architectonicidae. ApproximateK one third of these ex-
tant species are found in the Atlantic and the remainder
in the Pacific and Indian oceans (Merrill, 1970). Nu-
merous other species have been reported from fossil de-
posits (Gardner, 1947; Ladd, 19S2; Maur\, 1917; Ols.son,
1922; Olsson, 1932; Robert,son, 1973; Woodring, 1959).
Repre.sentatives of this familv usually have flattened,
low-spired shells with heterostrophic protoconchs. Bie-
ler (1993) reported that shells like those of architecton-
icids have been found in fossil deposits dating back to
the Trias.sic. Among extant groups, Bieler (1993) noted
that Psriidnmalaxis and Heliacus appeared hrst iTi the

Cretaceous while Architectonica and other forms ap-
peared in the Eocene. To date, including new forms re-
ported in this paper, a total of eight species of the family
are recorded in the Miocene Chipola Formation, Oak
Grove Sand, and Shoal River Formation.

Gardner (1947) listed sLx members of the Architec-
tonicidae from the Alum Bluff Group. Three were mor-
phologically similar to Architectonica nobihs Roding,
1798. These include Architectonica chipolana (Dall,
1892) from the Chipola River and Tenmile Creek in Cal-
houn County Florida, an unnamed subspecies of Archi-
tectonica quadriseriata (Sowerb)', 1850) from the Oak
Grove Sand in Florida and Georgia, and Architectonica
(juadriseriata waltonensis Garchier, 193fi, from the Shoal
River Formation in Walton Countv', Florida. Of the three
remaining taxa, two belong to the genus Architectonica.
Thev are Arcliitectonica nhear Gardner. 1936, from the
Oak Grove Sand and Architectonica vcrecnndn Gardner,
1936, from the Shoal River Formation, The third species
belongs to the subfamily Heliacinae and was tentatively
identified In' Gardner (1947'), from h\'o "ver\' voung'"
specimens, as Architectonica bisulcafn (d'Orbigny,
1842).

In this paper we look at four new members of the
faniiK .\rchitectonicidae from the Chipola Formation.
One is an uncommon species belonging to the genus
Architectonica that has been collected from onK' two lo-
calities ;i]ong Tenmile Creek. Two of the remaining spe-
cies belong to the genus HeUacits. Of these, one is fairly
common and is well represented in Chipola deposits.
This species is most similar to the "very yoimg ' speci-
mens of A. hisulcata reported by Gardner (1947). The
second species of llehacus is extremeh rare and is
known ouK from the holotxpe. The last representative
belongs to the genus Granosokirium. It is iilso ver\' rare
and is represented by two specimens collected from one
localitx along Teiunile ('reek.

'I'lie author also examined seven specimens of the sub-
species of Architectonica quadriseriata reported by
(iardner (1947) from the Oak Grove Sand, in order to

G. W. Schmelz, 2001

Page 23

^ll 6

^^8

k

Wk^i

A

Figures 1-10. New species of Miocene Architcctonicidae. 1-4. Architcctonica ocuuiiosd new species. 1, 2. Holohpe, UF 9.3992,
maximum height 11. S mm. maximum width 2.3..5 mm. 3, 4. Paratope A, UF 8461.5, maxunum height 9.2 mm, maximum width 18.1
mm. 5-10. Heliacus {Toiinista) calhouncnsis new species. 5, 6, 7. Holotvpe, UF 9.3991; maximum height 8.0 mm, maximum widtli
1.5.0 mm. 8, 9, 10. Parat\pe A, UF 8.5915; maximum height 7.2 mm, maxiuium uiihh 1:3,7 mm.

determine it.s taxononiic .statu.s. Si.\ of these specimens
are the material originalK- stutlied hv Garchier (deposited
at the National Museum of Natural Historx), while one
is from the private collection of Greta and Andy Murray,
Bradenton, Florida.

SYSTEMATICS

SuperfamiK .\rchitect()nicoidea Gray, 1S.50
FamiK' Arcliitectonicidae Gray, 1S.50
Genus Architcctonica Roding, 1798

Architcctonica Roduig, 1798; 78. T\pe species; Tmcliiis ])cr-
spectiviis Linnaeus, 1758, bv subsequent designation ol
Gray, 1847.

Architcctonica i^cnuno.'^a new species
(Figures 1—4)

Description: Sliell size moderate, low-spired, cone-
.shaped. Protoconch .smooth, maxiniuin iliameter
0.94-1.0.3 mm. with 1..5 whorls. Early half protoconch
whorl depressed. Anal keel present. Teleoconch whorls
7, slightly conve.x, separated by deep, narrow suture.
Each teleoconch whorl with 4 prominently sculptured,
beaded spiral cords of ecjual size. Base of sutural groove
formed b\ cord rumiing along peripiieral margin of body
whorl. Umbilicus moderately wide, outlined b\- wide,
denticulate spiral surroimded by deep, spiral groove.
Straight, axial grocnes radiate out from spiral groove and
cross over wide, fairly smooth band, which, in some
specimens, bears faint spiral threads. Two prominent.

Page 24

THE NAUTILUS, Vol. 115, No. 1

raised spiral cords, separated In- spiral groove, present
at peripherv of basal whorl. Innermost cord with well-
developed, beaded sculpture. Outermost cord with less
distinct sculpturing. In most instances, axial grooves ra-
diating Irniii spiral groove do not join beaded spiral cord
on peripherN'. Shape of aperture unknown as it is hadK
damag(Hl on ;dl specimens examined.

Holotype: UF 93992, height 11.8 mm. width 23.5
nun.

Paratypes: Paratope A: UF 84615, Iieight 9.2 mm,
width 18.1 mm, from t\j)e locality; UF 93993-94000, 8
paratvpes, Chipola Formation, north bank of Tenmile
Creek at powerline crossing about 1 mile west-northwest
of mouth of creek at "Baile\'s Ferrv" (SE 1/4 Sec. 12,
TIN, RlOW), Calhoun Countv, Florida (Tulane Univer-
sity' locality TU 830).

Tjpe locality: Chipola Formation, Tenmile Creek,
about 1.25 miles west of Chipola Riser (SE 1/4 Sec. 12,
TIN, RlOW), Calhoun Counts, Florida (Tulane Univer-
.sit^localitvTU 951).

Distribution: A decade of collecting Architcctonica
from numerous Chipola Formation sites and the exam-
ination of hundreds of specimens of Architcctonica at
the Florida Museum of Natural Histon' indicates that A.
geminosa is an uncommon species that is exclusively as-
sociated with the ancient reef environment that existed
along Tenmile Creek in Calhoun County, Florida.

Et^Tnolog>': Alludes to the species raritv and beautiful
sculpture.

Di.scussion: A number of investigators have noted that
living arcliitectonicids have a long-lived veliger stage re-
sulting in a broad geographical range for the adults (Bie-
ler, 1993; Merrill, 1970; Scheltema', 1979). By comparing
modem and fossil forms of Atlantic Architectonicidae,
Scheltema (1979) was able to demonstrate that the ve-
liger stage of Miocene members of this family also un-
derwent extensive migrations. Hence, in order to make
sure die newly tliscovered Chipola species had not al-
ready been described from other Miocene and Pliocene
deposits worldwide, specimens were compared with
those reported in tlie literature from a wide range of
locations including Europe, the Mediterranean, West Af-
rica, Caribbean, Soutli and C;entral America, Pacific
Ocean, as well as other locations in North America.

Architcctonica gcnwiosa bears some similarities to Ar-
chitcctonica ahcnr Gardner, 193(S, from the Oak (iroxe
Sand, but A. alvear does not have well developed scale-
like spiral cords on its dorsal surface and the nigose spi-
ral cord that surrounds the umbilical carina in A. ahcar
is absent in A. geminosa The only other fossil speiies
that bears any resemblance to A. gcnuno.sa is Architcc-
tonica nobihs karsteni Rutsch, 1934, which was dc-
.scribed from Miocene deposits in Venezuela (Rutsch,
1934), central Chile (Frassinetti and Covacevich, 1981)
and southwesteni Ecua(k)r (Marks, 1951). Woodring
(1959) also recorded this latter species from the Pliocene

Chagres Sandstone of Panama. However, A. nohili.s kar-
steni does not have well developed scale-like spiral cords
on its dorsal surface, its umbilical margin is less dentic-
ulate, and both peripheral cords on the ventral surface
are smooth.

A cursory examination of the "Architcctonica quadris-
criata ssp." of Gardner (1947) from the Oak Grove Sand
and Georgia revealed that it represents Arcltitcctonica
cliijiolaiia (Dall, 1892) and should no longer be consid-
ered an undescribed subspecies of Architcctonica quad-
riscriata.

CJenus Hchacus d'Orbignw 1842

Hilieiciis d'Orbignv', 1842: 68; introduced as "division" of So-
larium [= Architectonica]. Tvpe species: Solarium her-
hcrti Deshayes, 1830 [= Heliacus ctjlinclricus (Gnielin,
1791)], by nionotypy, under the incorrect .secondary spell-
ills' "lieberti".

Subgenus Torinista Iredale, 1936

Tiirinista Iredale, 1936: 327. Type species: Torinista papula
Iredale, 1936 = Solarium implrxum Mighels, 184.5, by
original designation.

Heliacus (Torinista) calhouucnsis new species
(Figures 5-10)

Description: Shell small, depressed, with flattened
apex. Protoconch smooth, sunken, luaxiiuum diameter
0.53 mm, with about 1.5 v\horls. Anal keel present. Te-
leoconch whorls 5.5, separated by deep suture. Single
row of well-developed, raised spiral cords runs along
whorls just above and below suture. Two less distinct
spiral cords lie between riiised ones. Spiral cords crossed
liv strong radiid ribs that give them beaded appearance.
Five prominent, elevated, spiral cords extend from outer
edge of last whorl to region halfvvav across base of shell.
First 3 cords keel-like with second one being largest.
Remaining 2 cords less distinct. Three prominently
sculptured cords, two outermost beaded, iiuiermost
strongly denticulate, surround wide undiilicus. .\ll raised
spiral cords along side and base intersecteil bv well-de-
veloped axial ribs, .\perture circular; outer lip crenulate.

Holot>pe: UF 93991, maximum luiglit 8.0 mm. max-
iniiim width 15.0 mm.

Paratopes: I'aiatvpe A: UF 85915, maxiniuni luiglit
7.2 mm, maximum width 13.7 nun, ("hipola Formation,
Tenmile C^reek, about 1.25 miles west of (-'hipola River
(SE 1/4 Sec. 12, TIN, RlOW), Calhoun County, Florida
(Tulane Universitv' locality TU 998); paratope lot UF
95074, 2 shells, (Chipola Formation. Tenmile C-reek,
about 1.25 miles west of Chipola River (SE 1/4 Sec. 12,
TIN, RlOW). C^alhoun County, Florida (Tulane Univer-
sitv localitv TU 951); paratxpe lot UF 91794, 3 shells,
Chipola Formation, east bank of Chipola River, about
1 .000 feet above Fourniile C:reek (SW 1/4 Sec. 29, TIN,
H9\\'), Calhoun Countv, Florida (Tulane University Lo-

G. W. Schmelz. 2001

Page 25

calih- TU 555): parahpe lot UF 84566. 6 sliclls. Cliipola
Formation, Tenmile ('reek, at powerline crossing alioiit
one mile west of C'hipola River (SE 1/4 Sec. 12, TIN,
RIOW), (Calhoun Counts Florida (Tiilane University Lo-
cality- TU 830); parat>pe lot UF 95075. 1 shell. t:liipola
Formation, Tenmile Creek, about 1 1/4 mile.s west of
Chipola Ri\er (SE 1/4 Sec. 12, TIN, RIOW). Callioun
Counts'. Florida (Tulane Universih Localih TU 951);
paratype lot UF 101100. 1 shell, Chipola Formation,
Tenmile Creek, about 1.25 miles west of Chipola River
(SE 1/4 Sec. 12 TIN, RIOW), Calhoim Count\-, Florida
(Tulane Universih' Locali^^' TU 998).

Type locality: Chipola Formation. Tenmile Creek,
about 1.25 miles west of Chipola Ri\er (SE 1/4 Sec. 12,
TIN, RIOW), Calhoun Count\-, Florida (Tulane Unixer-
,sit^■ locality- TU 951).

Distribution: Hcliacus caUwuncnsis is a fairK com-
mon species that is wideK distributed throughout fossil
deposits found along Tenmile Creek, Farle\' Creek, and
the Chipola Ri\er. Thus far there are no reports of it
being collected from Oak Grove Sand.

EtATiiology: Named after Calhoun Counts', Florida.

Discu.ssion: As a small member of the famih' Archi-
tectonicidae. Hcliacus has a more reticulate and stronger
sculpture than those individuals belonging to the genus
Architccfonica (Keen, 1971). Most Hcliacus are 20 mm
or less in size and e.xtant individuals of this genera are
identified b\' their distincti\e spiral operciilum which is
a hom\', pagoda-like structure with a fringed edge
(Keen, 1971). Since the opercula are not preserved as
fossils, other characteristics are used to differentiate fos-
sil species. These features include the tspe sculpturing
on the dorsal and ventral surfaces of the teleoconch
whorls, the presence or absence of an anal keel, the
widths of the protoconch and umbilicus, and the t\pe of
sculpturing surrounding the imibilicus.

Currentlv. Bieler (1993) has identified six subgenera
that belong to the genus Hcliacus. Based upon external
morphological features, die two new species of Hcliacus
described in this paper have been assigned to the sub-
genus Torinistd. According to Bieler (1993). this subge-
neric group possesses disk-shaped teleoconch whorls
with appro.ximatelv five noded, spiral ribs. On the side
of the bod\ whorl, thev also have strong lower peripheral
and infraperipheral ribs surrounding one or tvvo addi-
tional strong ribs.

Hcliacus callwuucusis was misidentified as Arcliitcc-
tonica (Pscuclotorinia) bisulcata d'Orbigny, 1853) by-
Gardner (1947), as she only had access to txvo very'
Noung specimens. An examination of several juvenile H.
caUwuncnsis h\ this investigator (two to three millime-
ters in diameter) revealed that the young do bear a slight
resemblance to the latter species. However, the juveniles
of H. calliouncnsis possess more heavily beaded spiral
cords around the mnbilicus and have fewer and less pro-
nounced beaded spiral threads on the dorsal surface.

A number of species of Hcliacus which appear to be-

long to the subgenus Taritiisla have been reported from
different fossil deposits ((;ar(lner.l948: Jung. 1969;
Ladd. 1982; Mansfield. 1930; Maun. 1917; Olsson,
1964; Petuch, 1994; Pilsbn-, 1922; Woodring, 1959) . Hc-
liacus calhouncusis is most siinilar to Solaiium sfonc-
munac Maurx', 1917 from the Miocene Gatun Formation
of Panama (Woodring. 1959) and (he Pliocene deposits
of the Orcado F"orination in the Dominican Republic
(Maur\-, 1917). However, it differs from this species by
the presence of keel-like spiral cords along its peripher\-,
faint spiral cords on the dorsal surface, and weaker radial
growth lines.

Among extant species H. caUwuncnsis is closeK relat-
ed to H. planispira (Pilsbry- and Lowe. 1932), but is not
flat-topped, has a more well de\eloped suture between
the dorsal whorls, and a wider, more denticulate cord
surrounding the umbilicus.

Hcliacus iToiiuista) cotnfiactus new species
(Figures 11-13)

De.scription: Small, depressed, cone-shaped shell
with angular peripher\-. Protoconch smooth, maximum
diameter 0.64 mm. with 1.3 whorls. First 0.5 protoconch
whorl sunken. Anal keel present. Five teleconch whorls,
first 2.5 whorls flattened, the remainder angled at 45°
toward periphery. Suture distinct, narrow. Five axial
rows of irregular-shapetl beaded cords present on sur-
face of whorls. These include hvo medium-size beaded
cords adjacent to the suture followed h\ a single, narrow-
axial row of beads, a wide trapezoid sciile-hke cord, and
a double row of narrow beads at the peripherx-. An an-
gled periphen is formed b\ 2 peripheral ribs. The area
lietween the peripheral ribs sculptured with trapezoid,
scale-like beads. F'ive prominentK beaded cords sur-
round deep umbilicus, the innermost one strongly den-
ticulate. Aperture oval.

Holot\pe: UF 93987. maximum height 6.8 nun. max-
imum width 1 1.3 mm.

Type locality: Chipola Formation. Tenmile ('reek
about 1.75 miles west of Chipola River (NE 1/4 Sec. 12,
TIN, R10\\'), Calhoun Countx. Florida (Tulane Univer-
sitx localitv TU 546; = USC;S 2212. 'one mile west of
Bailey's Ferry").

Distribution: H. conipactus is a \ery rare species tliat
is onl\ known from the hpe localih'.

Et\-mologA-: .Mludcs to tightK spiraled sculpture on
the bod\ wliorls.

Discussion: Hcliacus C(uni)actns is a uniijui- member
of this genus and iloes not bear a strt)ng resemblance to
am- other species from the (Chipola Formation, Oak
Grove Sand, and Shoal River Formation. From the fossil
record Architccfonica (Pscuclotorinia) liuppyi Jung.
1969, and Architcctonica (Architectonica) fuscicara
McNeil, 1984 are the onlv species which exhibit any sim-
ilarities to H. conipactus The former species is known

Page 26

THE NAUTILUS. Vol. 115. No. 1

J

15

16

• 17 II 18
f

Figures 11-19. New .species of Miocene Architectonicidae. 11-13. HcUacus {Torinista) compactus new species, HoloUpe, UF
9.39S7; nia.\imiim height 6.8 mm, maximum width 11.3 mm. 14-19. Granosolari\im floridanwn new species. 14, 15, 16. Holotvpe,
UF 96.324: maxinnnn height 2.0 mm, maximum width 6.3 mm. 17, 18, 19. ParaKpe A, UF 96.325: maximum height 1.8 mm,
maximum width 4..5 mm.

from several small specimens that were found in the
Pliocene deposits of Matiira Bay in Trinidad (Jung,
1969). Both H. compactus and A. ^itppyi are low spired
shells with with submerged protoconchs that possess five
beaded axial cords on the teleconch whorls and have an
angulate periphery'. However, H. compactus is much
larger, has a much more flattened apex, and the sculp-
turing on its dorsal and \entral surfaces differ consid-
erablv from A. guppi/i. Arcltitcctonico fuscicaca comes
from Ohgocene Mint Spring Formation and the B\Tam
Formation of Mississippi (McNeil and Dockerv, 1984)
and is probabK' the ancestor to the Miocene species.
HcUacus compactus is about the same size and possesses
the same general shape as A. fuscicava, but the spiral
beaded sculpturing on both the dorsal and ventral sur-
face of H. compactus is much larger than the beaded
sculpturing on A. fuscicava.

Genus Cranosolarium Sacco, 1892

Granosolaiitim .Sacco, 1892: .59. T\pe species: Solnnioii iiiil-
legramuii Lamarck, 1822, bv original designation.

Cranosolarium floriduttum new .species
(figures 1^19)

Descripti<ni: Shell small, trapezoid-shaped with bead-
ed spiral sculpture and sharp, beaded peripheral keel.
Protoconch smooth, depressed, maximum diameter
0,54-0.6.5 mm, distinctly heterostrophic, with 2 whorls.
Anal keel present. Teleoconch .15 whorls, deep suture
between each whorl. Opposing sides of each teleoconch
whorl with prominent beaded cord adjacent to suture.

Between the prominent beaded spiral cords on each
whorl, are three, less distinct, tile-like cords. Umbilicus
verv wide. Lower inner peripheral margin of bod\' whorl
extends dowii into umbilicus, which hears two parallel,
strongK' beaded carinae, that spiral toward the inner
ape.x of shell (into umbihcus). Remainder of ventral sur-
face with si.x beaded spiral cords of irregular size. Ap-
erture oval.

Holot\pe: UF 96.324, maximum height 2.0 nun, max-
innnn widtli 6. .3 mm.

Other Upe material: Paratxpe A. UF 96325. maxi-
mum height l.S nun. maxinuun width 4.5 nun. from
txpe locality.

Type locality': (^hipola Formation, north bank of Ten-
mile Creek at powerline crossing about 1 mile west-
northwest of mouth of creek at "Bailev's Ferrv" (SE 1/4
Sec. 12, TIN, RlOW), C;ilhoun Couim, Fk)rida (Tulane
UniversiU- locality- TU 8.30V

Distribution: This appears to be a verv rare species
that has thus far been collected onK irom the ancient
reef einironment of the t\pe locality.

EtynioIog>-: Named after the state of Florida.

Discus.sion: Graiuisolarium jiorhldiuiiu rcadiK stands
out from the rest of the Architectonicidae found in the
(!hipola Formation. In general, it has the overall body
form of the genus Architectonica but differs from this
group In its broad umbilicus and prominent peripheral
keel. Hielcr (199.3) assigned members of the .\rcliitec-
tonicidac with these characteristics to the genus Gra-
tiosolarium.

G. W. Schmelz, 2001

Page 27

Granosolfiriuiu florichiiium bears some similarih' to
Architcctonicii (Pscudotorinio) cuprcpcs Woodring,
192S, from die Pliocene of Jamaica. Both are about the
same size, but G. floridamim is more dorso-ventralK'
compressed, the sutures on the dorsal surface are more
distinct, and it possesses fewer beaded spiral cords
around the umbilicus. Granosolahum floridanuin is
somewhat similar to Granosolaiium aspcnan (Hinds,
1844) an e.xtant form which is listed by Bieler (1993)
from a number of diverse locations including Australia,
New Zealand, Indonesia, and West Africa.

Granosolarium aspcnim has also been reported from
the middle Miocene fossil deposits of Australia (Garrard,
1961) and from the Pleistocene deposits ot New Heb-
rides (Ladd, 1982). However, G. floridanuin differs from
G. aspcnim in that it possesses tile-like sculpturing on
its dorsal surface and paired, strongK' beaded, carinae
around the outer margin of the umbilicus.

Four species of Granosolarium ha\e been described
from Eocene deposits in the Ne\\' World, but the\' are
all larger and ha\e finer dorsal and \entral sculptural
features than those in G. floridamim. Two were collected
from the MoocKs Branch Formation in Louisiana and
described bv Palmer (1947). Thev are Architcctonica
(Granosolarium) ornata jacksonia Palmer, 1947, and Ar-
chitecfonica (Granosolarium) mcckana siihsplendida
Palmer, 1947. Architectonica aldrichi (Dall, 1892) was
collected from the Lower Claiborne of Mississippi and
Architcctonica clahorata was obtained from the lov\er
Claiborne and Gosport Sand of Alabama (Palmer, 1937).

ACKNOWLEDGMENTS

The author extends a special note of thanks to Roger W.
Portell for allowing examination of Chipola Formation
specimens under his care from both the Florida Muse-
um of Natural Historv and Tulane Collections, as well
as providing assistance with all photographic work and
reviewing earlier versions of this manuscript. Additional
appreciation is extended to Warren C. Blow, United
States National Museum of Natural Histon; for the loan
of specimens from the Gardner Collection, Andrew and
Greta Murrav for the loan of Oak Grove Sand speci-
mens, Cecil Sexton and Burt Haves for granting the au-
thor permission to collect on their property' and to Dick
Petit for his assistance with the acquisition of relevant
literature.

LITERATURE CITED

Bieler, R. 1985. Die Gattiingen der Arcliitectoiiicidae (Gastro-
poda: Allogastropoda). Teil 3: Architcctonica. Nipternxis,
Hvliacus. Fjisolariiun ,\rcliivtiir MdlliiskciikuiKlc 1 l(r89-
117,

Bieler, R. 1993. Architectoniddae of the Indo-Pacific (MoJIiis-
ca. Gastropoda). Gustav Fischer Verlag, Stuttgart. 376 p.,
3 pis.

Frassinetti, D. and V. Covacevich. 1981. Arciiitectonicitlae en
la Formacion Navidad, Miocene, Chile Central Parte II.
Architcctonica (Architcctonica) nobili.'i kar.'itcni Rutsch.

1934. Boletin del Museo National dc Historia .\atnral 38:
147-154, 4 figs., 1 tab.

Gardner, ], 1947. The niolluscan fauna of the .\lum Bluff
Group of Florida. Ft. S: Ctenobranchia (remainder), As-
pidobranchia, and Scaphopoda. United States Geological
Suney, Professional Paper 142-H:493-656, pis. 52-62.

Gardner, J. 1948. Mollusca from the Miocene and lower Pli-
ocene of X'irginia and North Carolina. Ft 2. Scaphopoda
and Gastropoda. United States Geological Sur\ev, Profes-
sional Paper 199-B:179-279, pis. 24-38.

Garrard, T. A. 1961. Mollu.sca collected by .Vl.\'. •'Challenge"
off the east coast of Australia. Journal of the Malacological
Socieh' of Australia l(5):2-;37, pis. 1-2.

Grav, J. E. 1847. A list of the genera of Recent Mollusca, their
sviionvma and tvpes. Proceedings of the Zoological Soci-
et\- of London 15:129-219.

Jung, P. 1969. -Vliocene and Pliocene mollusks from Trinidad.
Bulletins of .\nierican Paleontologv' 55(247>:2S9-657, pis.
1.3-60.

Keen, A. M. 1971. Sea shells of tropical West America. 2nd
ed. Stanford Universih- Press, Stanford C.\, xiv + 1064p.

Ladd, H. S. 1982. Cenozoic fossil mollusks from western Pa-
cific islands; Gastropods (Eulimidae and Volutidae
through Terebridae). United States Geological Survey,
Professional Paper 1171:iv + 100 pp., pis. 1—11.

MacNeil, F S. and t). T. Dockerv III. 1984. Lower Oligocene
Gastropoda, Scaphopoda, and Cephalopoda of the \'icks-
burg Group in Mississippi. .Mississippi Department of
Natural Resources, Bureau of Geologv- Bulletin 124:1-
41.5, pis. 1-72.

Mansfield, W. C. 1930. Miocene gastropods and scaphopods of
the Choctawhatchee Formation of Florida. Florida Geo-
logical Sui-vey Bulletin 3:1-185, pis. 1-21.

Marks, J. G. 1951. Miocene sfratigraphv and paleontologv- of
southwestern Ecuador Bulletins oi .\merican Paleontol-
og>-33 (1.39):1-162, pis. 1-9.

Maurs', C. J. 1917. Santo Domingo hpe sections and fossils.
Part 1: Mollusca. Bulletins ol .American Paleontologv' 5
(29):1-251, pis. 1-39.

Merrill, A. S. 1970. The family Architectonicidae (Gastropoda:
Mollusca) in the western eastern Atlantic. Unpuhl. Ph.D.
thesis, Universitv- of Delaware: .3.38 pp. 42 pis. (University'
Microfilms International, Inc., Ann Arbor, No. 71-6444.)

Olsson, A. A. 1922. The Miocene of northeni Costa Rica with
notes on its general stratigraphic relations. Pt. 1: Bulletins
of American Paleontologv- 9 (.39):1-167, pis. 1-15.

Olsson, A. A. 1932. Contributions to the Tertian,- paleontologv-
of northern Peni. Pt. 5. The Peruvian Miocene. Bulletins
of American Paleontologv- 19 (68):l-264, pis. 1-24.

Ols.son. A. A. 1964. Neogene mollusks from northwestern Ec-
uador Paleontological Research Institution: 256 p.. 38 pis.

Palmer K. \'. W, 1937. The Claiborni.m Scaphopoda. Ciastro-
poda and Dihranchiate Cephalopoda ot the soutlieni
United States. Bulletins of American Paleontologv- 7 (32),
pt. 1, 1-.548, pt. 2, pp. .549-7.30 pis. 1-90.

Palmer K. V. W. 1947. Univalves and index: p. 207-563. pi.
26-56. 62-65, In: Harris. G. D. and K. \'. W. Palmer (eds.)
The Mollusca of the Jackson Eocene of the .Mississippi
Emhavment (Sabine River to the .■\labama River). Bulle-
tins of American Paleontologv .30( 1 17): 1-5:53. pis. 1-65.

Petuch, E. J. 1994. Atlas of Flonda Fossil Shells (Pliocene and
Pleistocene marine gastropods) Spectrum Press, Evans-
ton, .394 pp., 100 pis., 20 text-figs.

Pilsbrv-, H. A. 1922. Revision ofW W. Gabb's Tertian.- .Mollusca

Page 28

THE NAUTILUS, Vol. 115, No. 1

of SaiiU) Doinini^o. Proceedings of the .\cadeinv of Nat-
ural .Sciences of Philadelphia 7.3;:3()5-t:35. pis. 16-47.

Pilsbn-, H. A. and H. \. Lowe. 19.32. West Me.\ican and Cen-
tral American inollusks collected by the H.N. Lowe,
1929-3L Proceedings of the .\cadeniv of Natural Sciences
of Pliiladelphia 84: .3.3-144, pis. 1-17.

Robertson, R. 1973. On the fossil histoiy and intrageneric re-
lationships of Philippin (Gastropoda: Architectonicidae).
Proceedings of the Academy of Natural Sciences of Phil-
adelphia 12.5:37-46.

Roding, P. F. 1798. Museum Bolteniannm sive catalogus ci-
mehonmi e tribus regiiis naturae. . . pars secunda. . . J. C.
Trapp, Hamburg, viii 4-199 pp.

Rutsch, R. 1934. Die Ga.stropoden aus dem Neogen der Punta
Gavilan in Nord-Venezuela. Abhandlungen der Schweiz-
erischen Palaeontologischen Ge.sellschalt 54-.5.5: 1-169.
20 figs., pis. 1-20.

Sacco, F. 1892. I molluschi dei terreni terziarii del Piemonte
e della Liguria, 12 (Pyramidellidae [fine], ringiculidae, So-
lariidae e Scalariidae [aggiunte]). C. Clausen, Torino, 86
pp.. 2 pis.

Scheltema, R. F. 1979. Dispersal of pelagic laivae and the zoo-
geography of Tertians' marine benthic gastropods. In: Grav,
|. and h.. J. Boucot (eds.) Historical Biogeographv, Plate
Tectonics and the Clianging Environment. Oregon State
Universitv- Press, CorvaUis, pp. .391—397, 6 figs.

Yokes, E. H. 1989. An overview of the Chipola Formation,
northwestern Florida. Tulane Studies in Geolog\- and Pa-
leontology 22:1:3-24.

Yokes, E. H. 1997. Notes on the fauna of the Chipola For-
mation— XLin. Additions to the molluscaii fauna since
1947. Tulane Studies in Geolog\' and Paleontolog)' 3:211-
216.

Woodring, W.P. 1928. Contributions to the geolog\' and pale-
ontologv of the West Indies. .Vliocene mollusks from Bow-
den, Jamaica. Part II: Gastropods and discussion of re-
sults. Carnegie Institute of Washington Publication 38.5:
i— vaii, 1-.564, pis. 1-40.

Woodring, W. P. 1959. Geologv' and paleontologx ol Ciuial
Zone and adjoining parts of Panama. Descrijition of Ter-
tiaiT mollusks (Gastropods: Yermetidae to Thaididae).
United States Geological Sunev. Professional Paper 306-
B:l-2.39 pp.. pis. 24-.38.

THE NAUTILUS 115(l):29-34. 2001

Page 29

On the publication date, authorship, and type species of
Umbracuhnn and TijJodina (Gastropoda: Opisthobranchia:
Tylodinoidea)

Angel Valdes

Dopaitnu'iit of Iinertebrate Zoologv

and Geolo!^'
Golden Gate Park
California Aeadeniv of Sciences
San Francisco, Ca'9411S USA
avaldes@calacadeniy.org

ABSTRACrr

The genns name Umbrncidmn Schninaclier, 1817 is a senior
svnionvm of Umhrclhi Lamarck, 1819, (liistroplax de Blains-ille,
l'819.'0//i/;n'//(( de P^enissac, 1821, Umbclhi delle Chiaje, 1831,
and Ojjcrciil/itiim Morch, 1852. Other names previously intro-
duced for this genus are not available in the meaning of the
International Code of Zoological Nomenclature. Several au-
thors have alternatively considered Patella sinica Gmelin, 1791,
or Patella iiinhracitla Lightfoot, 1786, to be the type species of
Umbraciilum. However, since none of these species was listed
in the original description of this genus, they are not eligible
to be the hpe species. Acardo iimbella Lamarck, 1801 [= Um-
braciiliim iindjraciilitin (Lightfoot, 1786)] is here selected to
be the type species of Lhnbraculum. In addition, a valid des-
ignation of tApe species for Umbrella, Ombrrlla. and Opercu-
latum could not be found in the literature; Umbrella iiidica
Lamarck, 1819, is here selected to be the type .species of these
genera. Thus, these three names become objective syiionyiiis.
The genus name Tijludina Rafinesque, 1814, type species Tij-
lodina piincfidata Rafinesque, 1814 [= Ttjlodina pervema
(Gmelin, 1791)], by nionotvpv, was published in 1814 and not
in 1819.

Additional key words: historical re\ision, nomenclature, type-
species designation.

INTRODUCTION

The genera Uinhroculuni and Ti/lodiiui imlutle relati\ely
\vell-kiioy\n opistliiihranch mollusks, recently rede-
scrihed liv W'illaii (1987). Thev appear to he closely re-
lated opistlidliranchs, classified yyithin the Umhraculo-
idea Dall, LSS9 (Willan, 1987), or more correctK- T\lo-
dinoidea J. E. Gray, 1847 (Willan, 1998). Both taxaare
characterized hv a dorsal, external shell, resembling an
umbrella or C'liinese hat. Members of Uiiil)riiadiiiu and
Ttjlodina have been known since the late 17()0s. hut so
far only a few species belonging to these genera have
been described. According to Bum (1959) there is only
one valid species of Umbracuhnn distributed througliout

all tropical and subtropical seas, whereas Thompson
(1970) distinguished an Atlantic and an Indo- Pacific spe-
cies, and Marcus (198.5) recognized a third species from
the Caribbean Sea. Thompson (1970) suggested that all
species of Tijlodina should be merged into a single one,
whereas Willan (1987) considered that this genus con-
tains five valid species. Lhnhracuhun and Tijlodina are
also known from the fossil record (\'aldes and Lozoviet,
2000), and several fossil species of Vmhraculum have
been described. Another related genus is Anidoli/ta Wil-
lan, 1987, txpe species Auidoh/ta diichcni Lcnen, 1846,
which contiiins less than five valid species (Waren and
di Paco, 1997). Two other genera previously assigned to
the Tylodinoidea are Bctiinia Jousseaume, 1883 and
Spiiicclla Rang and Des Moulins, 1828 (Pnivot-Fol.
1954; Willan, 1987). Bcrtinia lias been recently trans-
ferred to the Patellogastropoda (see Kasc and X'aldes,
1997), and the systematic position of Spiiicclla rem<iins
uncertain, but this latter genus probably represents an
opisthobranch mollusk (Vakles and Lozouet. 2000).

A review of the literature shows a large degree of dis-
agreement among different authors regarding the \alid
name and type species of Ihnhracidniii and the date of
publication of Ti/lodina. The present papi'r attempts to
determine the \alid name, exact date of publication, and
type species of these ty\o genera and their .s\iion\ins. In
order to reconstnict the nomeiiclatural histon of Um-
hracuhun and Tijlodina. this paper reviews the original
descriptions of these ty\(> genera as well as other related
literature.

DISCUSSION

Species of Tylodinoidea were named and described lor
the first time in several classic papers. These animals,
known only from the shell, were thought to be patello-
gastropods (Cnielin, 1791; Bosc, 1802). or a single valve
separated from species of the bivalve genus Acardo
Commervon /;/ Brugiiiere, 1789 (see Lamarck, 1801).

Page 30

THE NAUTILUS, Vol. 115, No. 1

Figures 1-3. Classic illustrations of Umbraculum. 1. Marti-
ni's (1769) 'Parasol Chinois'. 2. Dorsal view of Chemnitz's
(1788) 'Umbella Chinensis'. 3. Ventral view of Chemnitz's
(1788) 'Umbella Chinensis'. \\\ illustrations originally in color.

Mo.st of the classic descriptions of species and genera of
Tylodinoidea are not consistently binominal, and there-
fore most of the names introduced at that time are not
available (International Conmiissicjn on Zoological No-
menclature, 1999: Article 11). Davila (1767) and Martini
(1769) described species of Umbraculum for the first
time, under the names 'Lepas Umbella Chinensis' and
"Parasol Chinois' (Figure 1) respectixeh'. Chemnitz
(1788) compiled early descriptions of T\lodin(jidea un-
der the name 'Umbella Chinensis' (Figures 2-3), includ-
ing "Operculatum laeve' of Linnaeus (1753), Favart
d'Herbign\'s (1775) Parasol (>hinois', Patella paira Da
Costa, 1778 and de Favanne's (1780: 1784) Parasol Chi-
nois'. All these references appear to be .species of Um-
braculum except Da Costa's (1778) Patella parva, which
is a sviionym of the patellogastropod Tectum virginea
(Miiller, 1776). The works ol Martini (1769) and Chem-
nitz (1788) have been placed in the Official Index of
Rejected and Invalid Works in Zoological Nomenclature.
Lightfoot (1786) introduced for the first time a bi-
nominal name for a species of Umhraeuluni. Patella uiii-
hracula Lightfoot, 1786. Curiously, Patella timbracula
Lightfoot, 1786 was not listed by Sherbom (1902), who
only alluded to Gmelin's (1791) descriptions of species
ol T\lodinoidea: Patella sinica for Umbraculum, and Pa-
tella pcnersa for Ti/lodina. Dance (1962) revised Light-
foot's (1786) work, which meets the criteria of publica-
tion of the Code (International Commission on Zoolog-
ical Nomenclature, 1999: Article 8).

The V.\lid N.\me .\nd Type Species ok Umbr.\cvlvm

The \emacular name Umbella' was used in classic lit-
erature to designate species of T\lodinoidea (Davila,
1767; Chemnitz. 1788). Lamarck (1801) described a new
species of mollusk, Acardo umbella Lamarck. 1801,
based on vernacular references bv Davila (1767) and
Martini (1769). These tsvo references are descriptions
and drawings of shells of Umbraculum, and therefore
the species Acardo umbella Lamarck, 1801, belongs to
this genus. One of these references. 'Umbella Chinensis'
of Chemnitz (1788), was published in binominal form bv
Lamarck (1801) as Umbella chineiisii. ApparentK'. La-
marck (1801) was the first author to name Umbella in
the binominal form, but he introduced this name in the
SMion\m\' of Acardo, therefore rendering it unaxailable
(International Commission on Zoological Nomenclature,
1999: Article 11.6). Gray (1847) considered that 'Um-
bella' was originalK' and validl\' introduced by Chemnitz,
and selected 'Patella umbellata' to be the t\pe species
of this genus.

Lamarck (1801) believed that Acardo represented a
bivalve mollusk, and that Umbella chinensis was de-
scribed bv Chemnitz (1788) based on a single valve of
his new species Acardo umbella Lamarck. 1801, How-
ever, a re-examination of the description and illustrations
by Chemnitz (1788: 341, pi. 169, figs 1645-1646), clearly
shows that 'Umbella Chinensis' is a species of Tylodi-
noidea (Figures 2-3). This was recognized hv Schu-
macher (1817) and Pilsbrv (1895-96). The genus Acar-
do, Upe species Acardo ciustulaiius Comnier^on in
Bruguiere, 1789, was later found to be an epiph)'sis of
a whale, not a mollusk (Deshaves, 18.30: Gra\-, 1847).

ApparentK, Lamarck (1812) recognized that his Acar-
do umbella was not a valve of Acardo, but a different
genus of gastropod mollusks, and introduced for the first
time the genus name 'Ombrelle', with a verv short de-
scription and no specific names associated with it. How-
e\'er, Lamarck's (1812) work is not consistentK' binom-
inal and does not meet the criteria of availabilit)' of the
Code (International Commission on Zoological Nomen-
clature, 1999: Article 11). Years later, Lamarck (1819)
again used the name 'Ombrelle' for this genus of gastro-
pod mollusks, this time also Latinized as Umbrella. At
the same time Lamarck (1819) described two new spe-
cies. Umbrella iudica Lamarck. 1819. and Umbrella med-
iterranea Lamarck. 1819. and regarded Gastroplax de
Blmnville. 1819, as a .svnomTn of Umbrella. De Blaimille
(1819) based Ga.stroplax on a single species, 'Patella om-
bracula Chemnitz', with valid binominal name Umbra-
culum umbraculum (Lightfoot. 1786), of which it is the
t\pe species In* monotvpv Lamarck's (1819) paper is
consistently binominal, and Sherbom (1925) and Neave
(1939) regarded it as the first valid introduction of the
genus name Umbrella. In addition, Lamarck (1819) de-
scribed for the first time the soft parts of a representa-
ti\c of this genus, a task later completed by Moquin-
Tandon (1870). In the following years, and during most
of the 19th (-entun, the genus name Umbrella gained

A. Valdes, 2001

Pai^e 31

unhersal acceptance. Examples of usage of this name
are the papers hv Menke (1828). Rang (1829), Deshaves
(1830), Phihppi (1836), Sowerbv (1839), Cantraine
(1841), Herrmannsen (1846-49), 'Soulevet (1852), Fi-
scher (1880-87), Vavssiere (1885) and Locard (1892).
Most of these authors considered Lamarck's (1812) work
as the first valid description of this genus, and subse-
quently Herrmannsen (1846-49) [1849] selected 'Patella
umbrella Chemn.' as t\pe species of the genus. The first
valid introduction of the genus UtnbrcIIa is Lamarck's
(1819). 'Patella umbrella Chemn.' is not a valid species
and was not originalK' listed in the description of the
genus. Therefore, Herrmannsen's (1846—49) [1849] is
not a valid tspe-species selection for Umbrella. I have
not been able to find in the literature a \alid designation
of t\pe species for Uinbrclla Lamarck, 1819. thus Um-
brella indica Lamarck, 1819 [= Umbraculiim unibra-
ciiltim (Lightfoot. 1786)] is here selected to be the tspe
species of the genus.

Schumacher (1817) described the genus Umbraculiim
based on references to Martini's (1769) 'Parasol Chinois',
de Favanne's (1780; 1784) "Parasol chinois', Chemnitz's
(1788) 'Umbella Chinensis', Acardo iimbcUa Lamarck,
1801, Patella umbellata Bosc, 1802, and Acardo orbi-
cularis Megerle von Miihlfeldt, 1811. Subsequently, Pils-
bry' (1895-96) [1896] selected Umbraculum sinicum
(Gmelin, 1791) to be the t\pe species, and at the same
time considered that several of the species cited in the
original description of Umbraculum were .s\nonvms of
17. sinicum. Otlier authors (Thompson, 1970: Marcus,
1985; Willan, 1987) considered Patella umbracula Light-
foot, 1786, to be the tNpe species of Umbraculum. Ac-
cording to the Code (International Commission on Zoo-
logical Nomenclature. 1999: .\rticle 67.2). since Umbra-
culum .finicum (Gmelin, 1791) and Patella umbracula
Lightfoot, 1786, were not included in the original de-
scription of Umbraculum. neither of these two designa-
tions of tvpe species is valid. I have not fomid in the
literature a valid selection of t^pe species for Umbra-
culum; accordingly, Acardo umbella Lamarck, 1801,
which was included in the original description of L'//i-
bracidum. is here selected to be the bipe species of this
genus. According to Pilsbr>' (1895-96), A. umbella is a
junior svnomTn of U. sinicum. which is also a junior sMi-
on\m of U. umbraculum (see Willan, 1987). There is no
(juestion that Umbraculum and Uittbrella were described
with the same features, and most authors regarded botli
names as sMiomins (see Willan, 1987).

Pilsbr\- (1895-96) noticed for the first time that Um-
brella was validK' introduced b\- Lamarck in 1819, and
not in 1812, and therefore that Umbraculum was a se-
nior SMionym of Umbrella. In the following years, ;ill
authors followed Pilsbry's opinion, and the genus name
Umbrella has not been used as valid for more than one
hundred years. The latest papers I found using the name
Umbrella as \-alid are those b\- Va\ssiere ( 1885) and Lo-
card (1892). Since Pilsbry's (1895-96) paper, the genus
name Umbraculum has gained universal acceptance and
it is now in constant and exclusive use.

Other Svncjwms of Umbraciiam

Some of the vernacular or unavailable genus names in-
troduced for species of Umbraculum in the classic pa-
pers mentioned above, were later latinized or became
available bv other means. These names are thus junior
svnon\TOS of Umbraculum. Fenissac (1821), latinized for
the first time the name Ombrella, based on the French
spelling 'Ombrelle', also used by Lamarck (1812; 1819)
for Umbrella. Delle Chiaje, 1830-31 [1831] again laHn-
ized the genus name Umbella. this time not in svnon\inv,
and therefore he is the author of the name. Miirch
(1852) made available the genus name Operculatum for
the first time, which is a binominalization of 'Opercula-
timi' Linnaeus, 1753. In the original descriptions of both
Ombrella and Operculatum several species were listeil
and no t\pe species was designated. I ha\e selected Um-
brella indica Lamarck, 1819, which is mentioned by Fer-
ussac (1821) and Morch (1852) to be the t\pe species
of both genera.

Willan (1987) suggested that Spiricella could be a s\n-
onvm of Umbracidum. but this genus is clearK distinct
and probably not even a member of the T\ lodinoidea
(see Valdes and Lozouet, 2000).

The Date of Pubfkatiox of TYLoni\.\

The genus name Ti/lodina was originalK' and \alidly in-
troduced b\- Rafinescjue in 1814 and not in 1819, as sug-
gested by some authors (e.g., Pilsbr)-, 189.5-96; Willan,
1987). Several sources (Herrmannsen, 1846—49: Sher-
bom, 1925; Neave, 1939) already pointed to the work
bv Rafinesque (1814) as the first \alid introduction of
the name Ti/lodina.

Pilsbr)' (1895-96), and subsecjuently other authors,
considered that the genus Tijlodina was introduced for
the first time bv Rafinescjue (1815), who mentioned the
name Tijlodina without a description. Accorcbng to Pils-
bn- (1895-96), Tijlodina Rafinesque, 1815, is a iiomen
nudum. Later, Rafinesque (1819) published a full de-
scription of Tijlodina. which was regarded b)' Pilsbr)'
(1895-96) as tlie valid original description of the genus.
However, Rafines(jue (1819) mentioned: "Since the 12th
and latest issue of my Journal cncijclopedique de la Si-
cilie [Rafinesque, 1814] is not in Paris, and it w^as almost
completeK destroxed tluring the two consecuti\e ship-
wrecks that I suffered, I am sending you the characters
of eleven genera of mollusks and poKps. among the 36
new genera that it cont;iins, begging you to publish them
again ". It is evident then, that the genus Tijlodina was
actualK- first described in 1814, and that most copies of
the 12th issue of this paper were lost in the mentioned
shipwrecks.

I found a copy of Rafinesque's (1814) work at the li-
bran- of the Museum national d'Histoire naturelle in
Paris, but as Rafinesfiue commented in 1819, i.ssue 12
(volume 2) is missing. However, there are still copies of
that work available in public libraries in the United
States. Fitzpatrick (1911) described in detail Rafin-

Page 32

THE NAUTILUS, Vol. 115, No. 1

e.sques (1814) work and mentioned the 12tli and la.st
i.ssue of \()liime 2 (pp. lfiI-196). Tlii.s is.sne include.s a
chapter entitled "Soniiolo^ia. Definizioni di .36 nuovi
Generi di Aniniali niarini della Sicilia, pp. 161-166" that
contains tlie original description of Tt/lodiiui. A part of
this chapter, including onl\' 1 1 of the 36 originalK' de-
scribed genera, was published again h\ Rafinesque
(1819), repeating the description of Ti/loclina. According
to Fitzpatrick (191 1) there are full copies of Rafinesque's
(1814) work at the Librar\' of the Academy of Natural
Sciences of Philadelphia and the Library of the Histor-
ical Societs' of Penns\lvania. Another cop\', lacking issue
12 of vohnne 2, is at the Library- of the Congress. There
are more copies lacking several issues in other public
libraries in the United States. Sherboni (1925) cited Raf-
inesque's (1814) paper and mentioned the page number
with the original description of Ti/lodiiui (p. 162), so it
is ver\' hkely that he also saw one of the remaining com-
plete copies. UnfortunateK, I had not direct access to
an\' of Rafinesque's (1S14) complete copies of this work,
which, due to its raritw are no longer available for loan
or reproduction.

Binney and Tryon (1864) compiled the papers by Raf-
inesque, but thev did not include the paper of 1814 on
Ti/Io(lina. onl\- the papers of 1815 and 1819. This was
probabl)' the reason why Pilsbiy (1895-96) and subse-
quent authors erroneously determined 1819 as the orig-
inal publication date of Ti/lodino.

l<Hiiiitisia di Monterosato, 1884 (hpe species Ti/lodiiui
citrina |oannis, 1853, b\- original designation), and Ti/-
lodiiuila Mazzarelli, 1897 (hpe species Tylodinella trin-
chesii Mazzarelli, 1897, by monotypy), are junior s\tio-
nvms of Tijlodina (see Willan, 1987). The genus Par-
mophonis de Blainxille, 1817 (hpe species Pannophoni.s
elongatus de Blain\ille, 1817) is a .synonym of Scutus de
Montfort, 1810 (Fissurellidae) and not a .s\Tion\'m of Ti/-
lodina as suggested bv 'Willan (1987). When introducing
this new genus, de Blainville (1817) mentioned that Par-
tnophonis is just a translation into Greek of the name
Snilits.

TiiF TiTK spFcirs oi' T\iA)ni\.\

Rafinesque (1814) introduced the gemis Ti/lixliiui based
on a single species, Tt/lodina pttnctulata Rafinesque,
1814 (described in the same work), which became the
t\pe species b\- monot\p\- (see Sherboni, 1925). Willan
(1987) erroneously stated that Pilsbn' (1895-96) selected
Tijlodina citrina Joannis, 1834, to be the type species oi
Tijlodina h\ sub.sequent designation. The species name
T, citrina is not mentioned in an\ of l^afines(jue's papers
deahng with Tijlodina (1814; ]8i5: 1819) and therefore
it is not eligible to be the type species. Moreover, Pilsbn
(189.5-96) did not select a t\pe species for Tijlodina.

Pilsbrv' (1895-96) conunented that it w-as uncertain
which spt>cies of Tijlodina was kncmni to Rafines(jue.
Pilsbr)' (1895-96) considered several valid species of this
genus to coexist in the Mediterranean Sea. In light of
Willan's (1987) review of Tijlodina it appears that there

is onK oni- valid Mediterranean species, Tijlodina pcn-
crsa ((Jmelin, 1791), which is most likelv a senior s\ii-
onvm of T. piinctidata.

CONCLUSIONS

After the critical review of the pertinent literature, the
valid name, e.xact date of pul^ilication, and hpe species
of the genera Unihracidiim, Tijlodina. and their junior
sMionvms has been determined. In order to clarifv the
results of this work a summary svTionvTny is presented
here.

Vinhnindiun Schumacher, 1817: 177-179.

Type species: Acardo umbella Lamarck, 1801 [= L'//i-
hracidiun umhraciduni (Lightfoot, 1786)], here desig-
nated.

-I- Umhirlla Lamarck, 1819: .3.39-.34.3. T\pe species: Um-
hrrllii indira Lamarck, 1819 [= Vmbracnlum nmhra-
culum (Liglitfoot, 1786)], here designateil.

+ Gn.stn>j)lnx de Blain\ille, 1819: 182. T\pe species: Uin-
hmculum iimbraailxim (Lightfoot, 1786), by original
designation, originally cited as Patella ombracula
Chemnitz'.

+ OinhrcUd de Fenissac, 1821: x.\i.\. T\pe species: Lhn-
hirlld inilini Lamarck, 1819 J= Umbracuhun mnbiri-
cidmit (Li^littoot, 1786)], here designated.

+ Umbcllfi Delle Chiaje, 1822-1831 [1831]: 2()(). 209.
213. T\pe species: VmhcUa inediterranca (Lamarck,
1819), bv monotvpy.

+ Operadatiini Morch, 18.52: 137. Tvpe species: I'inhrrlhi
indica Lamarck, 1819 [= Umbraculum umbrnciilum
(Liglitfoot, 1786)], liere designated.

Tijlodina Rafinescjue, 1814: 162.

Type specie.s: Tijlodina punctidata Rafinesque, 1814
[= Tijlodina pcncr.sa (Gmelin, 1791)], bv monotspv.

-I- jofiiinisin di Monterosato, 1884: 149. Tvpe species: Ty-
lodina citrina Joannis, 18.53 [= Tijlodina perversa
(Cimelin, 1791)], bv original designation.

-I- TijlodincUa Mazzarelli, 1897: 596-600. T\pe species:
Tijhxiiiicllti trinrhc.sii Mazzarelli. 1897 [= Tijlodina
pcncr.sa (Gmelin, 1791)]. bv monohpv

.\CKNOWLEDGMENTS

I am \en gratehd to Lam ('urrie and Patricia Shea-
Diner, of the Libran of the (California ,\cademv of Sci-
ences, for their invaluable help in compiling the litera-
ture necessarv for the completion of this paper.

LITERATURE CdTED

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A. Valdes, 2001

Page 33

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THE NAUTILUS 115(l):35-36. 2001

Page 35

Notes

A note on Liicina miilfilineafa 'Tuomey and Holmes" (Biv
Lucinidae)

Richard E. Petit'

Research Associate
Department of Invertebrate Zoology
National Mnseuni of Natural Historv
Smithsonian Institution
Washington, DC 20560-0118 USA

Lib.'arv

APR 1 ^ 2001

Wov'~

The species name multilincata "Tuonie\- and Holmes" is
in current usage for a species of Parvilucino that occurs
in the Pliocene and Pleistocene of the Carolinas and
Florida and in the Recent Fauna of the western Atlantic
from the Carolinas to Brazil.

Tuomev and Holmes (1S56: 61, pi. 18, figs. 16. 17)
identified this species as Lucina multistriata Conrad
(1844: 307). The\' did not cite the original description
but cited Conrad's later usage (1845[in 18.38-61]; 71, pi.
40, fig, 6). UnfortunateK', in citing Conrad the\' mis-
spelled the species name as multilincata instead of mul-
tistriata. As Conrad is not onl\' shown as author of the
binomen, both in the text and on the plate caption, but
a reference to his work is also given, it is clear that mul-
tilincata is an incorrect spelling. Holmes (18.58: 29. pi.
6, fig. 6) repeated die error, using the same spelling, and
crediting authorship to Conrad.

The first person to credit this name to Tuome\' and
Holmes was Dall (1903: 1384) who recognized that the
shell figured b\' Tuome\- and Holmes was distinct from
L. multistriata Conrad and cited it as Phacoidcs (Par-
vilucina) multilincata Tuomev and Holmes. He placed
in svnon\'m\- the Recent Phacoidcs {P.) crcnclla Dall,
1901. Gardner (1944: 79) followed Dall and added the
comment that the misspelling bv Tuomey and Holmes
was a "happv blunder" as the species thev figured was
not conspecific with L. multistriata Conrad. The validity'
of Dall's action has ne\er been fjuestioned. However, a
misspelling of an attributed name has never made sucli
misspelling an available name. In both the current Code
(Intemationiil Commission on Zoological Nomenclature,
1999) and the one just superceded (International Com-
mission on Zoological Nomenclature, 1985), this is ad-
dressed in Article 33, Among the man\' workers who
have followed DaU and Gardner in attributing this name
to Tuome\ and Homes, alwa\s incorrectK' dated, are Ab-
bott (1974: 459), Bretsk-\- (1976: 263), R'ios (1994: 252),
and Turgeon ct al. (1998: 38). Campbell ( 1993: 28, figure
63), in reporting the species from the Pliocene of \'ir-

' Mailing address: 806 St. Charles Road, N^
SC 298.52-2846 USA.

)rtli .VIvrtle Beach,

ginia, correctK' dated Tuome\- and Holmes but listed no
SMiomms except for a typographical error bv Cooke.
19.37,'

It appears that the earliest available name for the spe-
cies figured b\- Tuome\- and Holmes and later authors
as Lucina multilincata Tuomes and Holmes, is Panilu-
cina crenella (Dall. 1901: 825.' pi. 39. fig. 2). Dalls orig-
inal figure is reproduced bv Abbott (1974: 458, fig.
5290)."

In recent correspondence with Drs. E. V. Coan and
L. D. Campbell concerning this note, thev have ad\ised
that the svstematics of Parvilucina of both the eastern
Pacific and westem Atlantic are in need of revision. The
comments of both on this short note are acknowledged
with appreciation.

LITERATURE CITED

Abbott, R. T. 1974. American Seasheils. 2"' edition. Van Nos-
trand Reinhold, New York, 66.3 pp., 24 pis.

Bretskv, S. S. 1976. Evolution and classification ot the Lucin-
idae (Mollusca: Bi\ai\ia). Paleontographica .\mericana
8(50):219-3.37. pis. 2.5-36.

Campbell, L. D. 1993. Pliocene molluscs from the Yorktowii
and Chowan River Formations in Virginia. Virginia Divi-
sion of Mineral Resources Publication 127:i-\ii. 1-2.59.

Conrad, T. A. 1838-61. Fossils of the Medial Tertiarv- of the
United States. Dobson. Philadelphia. x\i + 89 pp., 49 pis.
[.\vi + .32. pis. 1-17. 1838: 3;3-,56, pis. 18-29, 1840; .57-
80. pis. ,30-45. 1845; 81-89. pis. 46-49, 1861], [Reprinted
1893 by W. H. Dall. Philadelphia]

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tv-nine new .Miocene, and one Eocene fossil shells of the
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fomiation near Mvrtle Beach. S.C. Journal of the VViLsh-
ington Academy of Sciences 27(l):l-.5.

Dall. VV. H. 1901. Sviiopsis of the Lucinacea and of the Amer-
ican species. Proceedings of the United States National
Museum 23( 1237):779-.S.33, pis. 39-42.

Dall, W. H. 1903. Contributions to the Tertiary fauna of Flor-
ida with especial reference to the silex beds of Tampa and
the Pliocene beds of the Caloosahatchie Ri\er Part VI.

Page 36

THE NAUTILUS, Vol. 115, No. 1

Concluding the work. Tran.sactions of the Wagner Free
Institute of Science 3(6):viixiv, 1219-1654, pis. 4.S-60.

Gardner, J. 1944. Mollusca from the Miocene and Lower Pli-
ocene of \'irginia and North Carolina. U. S. Geological
Survey Professional Paper 199-.\:1-17'S. [imprinted date
1943; actual publication date Januaa- 2S, 1944].

Holmes, F. S. 18.5<S-6(). Post-Pleiocene fossils of South Caro-
lina. Russell and Jones, Charleston, xii + 122 -I- v pp., 28
pis. [1-64, pis. 1-10, 1858; 65-98, pis. 11-14, 1859; i-xii,
99-122, pis. 15-28, i-v. I860].

International Commission on Zoological Nomenclature. 1985,
Inteniationiil Code of Zoological Nomenclature, Third
Edition. International Tnist tor Zoological Nomenclature,
London, Berkele\', and Los Angeles. x.\ + 338 pp.

International Commission on Zoological Nomenclature. 1999.
International Code of Zoological Nomenclature, Fourth

F.dition. International Trust for Zoological Nomenclature,
London, .x.\L\ + 306 pp.

Rios, E. de C. 1994. Seashells of Brazil, 2'"' edition. Funda^ao
Universidade do Rio Grande, Rio Grande, 368 pp., 113 pis.

Tuomey, M. and F. S. Holmes. 1855-57. Pleiocene fossils of
South Carolina. Russell and Jones, Charleston, .wi -I- 152
pp., .30 pis. [l-.3(), pis. 1-12, 18.55: 31-144, pis. 1.3-28,
1856; 145-152, i-\-vl pis. 29-30, 1857.] [Reprinted 1974,
Paleontological Research Institution, Ithaca, N.Y.]

Turgeon, D. D., J. F. Q\iinn, Jr., A. E. Bogan, E. V. Coan, F.
G. Hochberg, \V. G. Lyons. R M. Mikkelsen, R. J. Neves,
C. F E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.
G. Thompson, M. Vecchione, and J. D. Williams. 1998.
Common and scientific names of aquatic invertebrates
from the United States and Canada. Mollusks. Second
Edition. American Fisheries Society, Special Publication
26, Bethesda. ix -I- 526 pp.

A replacement name lor a New Caledonian CalUostoma species
(Gastropoda: Trochidae)

Bruce A. Marshall

Museum of New Zealand Te Papa

Tongarewa
PO. Box 467
Wellington
NEW ZEALAND

A.S a result ot protedural omission during preparation of
a recent paper (Marshall, 1995), I overlooked the fact
that a specific epithet chosen for a new CaUtostoma spe-
cie.s was already in use in' Quinn (1992).

CdUtostoma (Benthastrlcna) kdiitiknniiu new name
Calliustunm {Bcnthastclcna) coruntitiim Marshall, 1995; 409,

fig. 49—51, 128, 156. Not Calliu.stonia coronatiitn Quinn,

1992.

The replacement name is lor the indigenous people of
New Caledonia and Vamuitu.

LITERATURE CITED

Marshall. B. A. 1995. Calliostomatidae (Gastropoda: Trochoi-
dea) from New Caledonia, the Lo\alt\' Islands, and the
northern Lord Howe Rise. Resultats des Campagnes MU-
SORSTOM 14. Memoires de la Museum National
d'Histoire Naturelle 167:381-458.

Qviinn, J. F 1992. New species oi Calliostanui Swainsou. 1840
(Gastropoda; Trochidae), and notes on some poorK known
species from the western .\tlantic Ocean. The Nautilus
106:77-144.

Erratum

In the section "Txpe material" in Simone <•/ ai (2000: L39) we inacKertentk printed an incorrect parahpe number.
The USNM paratvpe numlier therein should he 880725, not 20KiO()9 as originalK' stated.

LITERATURE CITED

Simone, L. R. L, G. Pastorino, P. E. Penchaszadeh. 2000. Ci<-))iihihi ciiicntnui iCJastrojioda; CaKptracidae), a new species horn the
littoral of ArgenHna. The Nautilus 114:127-148.

THE NAUTILUS 115(1):37, 2001

Page 37

Book Review

Panamic Province MoUiiscan Literature:
Additions and Changes from 1971 through
2001

Sko<iluiuI, C. 2001. Panamic Province niolluscan litera-
ture: Additions and changes (rom 1971 tlirough 2001. 1.
Bivalvia. 2. PoKplacophora. The Festi\ns 32 {supple-
ment):i-v-l- 1-119 (Bivalvia); 1-20 (PoKplacophora).

If the concept of "deliniti\e" could he applied in sci-
ence, M\Ta Keen's (Keen. 1971) second edition of Sea
Shells from Tropical West America would he it. Includ-
ing in excess of 3.300 species and spanning 1064 pages,
the volume covered e.xliaustivelv the Panamic malaco-
fauna (25°N-6°S) as understood in 1971, packing more
information per species than what is the norm for books
of its kind. It also brought with it a wealth of biblio-
graphic references. But 30 \ears have now passed and
nothing in science is final and conclusive. Thanks to the
work of a large number of West Coast malacologists,
some of them Keens own graduate students, manv new
t;«a were discovered, the identitv of others reevaluated,
geographic ranges and the phvlogenetic relationships
among moUusks better defined. Through comprehensive
research of the literature, Carol Skoglund has endeav-
ored to bring up to date information on Panamic mol-
lusks in a series of papers previousK published in the
San Diego Shell Club publication The Festivus (e.g.,
Skoglund. 1989; 1991).

The two-section, non-illustrated volume is informa-
tive, well researched, and as Keen's book did before it,
vields a huge amount of bibliographical references. After
iill, this is the main proposal of Skoglund's new volume,
to provide an update on taxonomic works on Panamic
bivalves and chitons published since 1971. But the vol-
inne is much more than just a listing of references on
Panamic Hterature. Species are hsted according to cur-
rentlv accepted arrangements: the supraspecific taxtjno-
mv of the section on bivalves is stnictmed after the su-
perb work bv Coan, Scott and Bernard (2000). Changes
in ta.xonomic, nonienclatural, or geographical status in
species previouslv treated in 1971 are denoted by Keen's
original species number (but not necessarily in the same

sequence), given in a left-indented column before each
entrv. This svsteni facilitates cross-referencing and is
particularlv useful when species are moved to another
genus or familv. New names introduced since 1971, ei-
ther as new taxa or imder new combinations, are indi-
cated bv boldface printing. The treatments are uniform,
with topics under each specific entrv following an estab-
lished sequence. The work is plentv in annotations, and
no deletions or additions are left unexplained. A taxo-
nomic index, much desired in this tvpe of work, is pro-
vided; a spot check showed it to be complete and vir-
tuallv free of t\pographic or transcription errors. The
same is true for the remainder of the volume, and both
author Skoglund and TIte Festivus editor Carole Hertz
are to be congratulated on the task.

Although I am far from being a specialist on Panamic
bivalves and chitons. I am impressed with the profusion
of information and the consistencv present in Skoglund's
work. A similar update on the Panamic gastropods would
be most welcome!

The cost within the US is $22 postpaid, overseas $25
(surface) or $30 (air mail). Contact the San Diego Shell
Club, c/o 38S3 Mt. Blackburn Ave., San Diego, CA
92111, USA, or e-mail Carole Hertz (cmhertz@pacbell.
com) for further ordering information.

LITERATURE CITED

Coan. E. V, P. V. Scott and F. R. Bernard. 2000. Bivalve sea-
shells of western North .America: Marine mollusks from
Arctic Alaska to Baja California. Santa Barbara Museum
of Natural Historv Monographs number 2, 764 pp.

Keen, A. M. 1971. Sea shells of tropical West America: Marine
mollusks from Baja Caiif'oniia to Peni. 2'"' edition. Stan-
ford UniversiK Press, Stanford, xiv + 1064 pp.

Skoglund, C. 1989. Additions to the Panamic Province gastro-
pod literature - 197f through 1988. The Festivnis 21(9):
7^8-91.

Skoglund, C. 1991. Additions to the Panamic Province bivalve
(Mollusca) literature 1971 to 1990. The Festivus 2,3 (sup-
plement): 1-74.

Jose H. Leal

The Bailev -Matthews Shell Miisrum
.307.5 Sanibel-Captiva Road
Sanibel, FL 33957 USA
Jleal@shellmuseiim.org

THE NAUTILUS 115(1):38, 2001

Page 38

Notices

THE 2001 i; I ABBOTT VISITING CURATOHSIIIP
The Bailey-Matthews Shell Museum is pleased to imite applications for the 200] R. T Abbott Visiting Curatorship.

The Curatorship, established originallv in accordance with the wishes of the late Dr. R. T Abbott, Founding
Director of the Shell Museum, is awarded annualK to enable malacologists to visit the Museum for a period of a
week. Abbott Fellows will be expected, by performing collection-based research, to assist with the curation of
portions of the Museum's collection and to provide one evening talk for the general public. The Museum collection
consists of marine, freshwater, and terrestrial specimens. A large percentage of our hoklings have been catalogued
tlirough a computerized database management system. A substantial portion of the time will be available for
research in the collection, but field work in southwest Florida can be arranged. The R. T Abliott Visiting
Curatorship is accompanied by a stipend of $1,500.

Interested malacologists are invited to send a copv of their curriculum vitae together with a letter detailing their
areas of taxonomic expertise and research objectives, and provide a tentative subject for their talk. Send materials
to:

Dr. Jose H. Leal

The Bailev-Matthews Shell Museum

R O. Box 1580

Sanibel, FL 33957

jleal@shellmuseum.org

Applications for the 2001 Visiting Curatorship should be sent no later than Mav 15, 2001. The award will be
announced by late June. Questions about tlie Visiting Curatorsliip should be sent to the e-mail address above, or bv
phone at:

(941) 395-2233; fax (941) 395-6706

/

This pul)lication is sponsored in part bv

the State of Florida, Department of State,

Division of (Cultural Affairs, and

the Florida Arts Council

APR i ^' zm

Waocs'..^, .

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rHE NAUTILUS

Volume 115, Number 2
August 6, 2001
ISSN 0028-1344

A quarterly devoted
to malacology.

EDITOR-IN-CHIEF

Dr. Jose H. Leal

Tlie Baile)-Matthews Shell Museum
3075 Sanibel-Captiva Road
Sanibel.FL 33957

MANAGING EDITOR

Christina Petrikas

The Bailey-Matthews Shell Museum
3075 Sanibel-Captiva Road
Sanibel.FL 33957

EDITOR EMERITUS

Dr. M. G. Harasewych
Department of Invertebrate Zoology
National Museum of
Natural Histor)'
Smithsonian Institution
Washington, DC 20560

CONSULTING EDITORS

Dr. Riidiger Bieler
Department of Invertebrates
Field Museum of
Natural History-
Chicago, IL 60605

Dr. Arthur E. Began

North Carolina State Museum of

Natural Sciences

Raleigh, NC 27626

Dr. Pliihppe Bouchet
Laboratoire de Biologic des
Invertebres Marins et Malacologie
Museum National d'Histoire Naturelle
55, rue Buffon
Paris, 75005 France

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

Dr Eileen H. Jokinen
8234 E. North Shore Road
Sault Ste. Marie, MI 49783

Dn Douglas S. Jones
Florida Musemn of Natural Histon-
University' of Florida
Gainesville, FL 32611-2035

Dr. Harry G. Lee

1801 Barrs Street, Suite 500

Jacksonville, FL 32204

Dr. Charles Lydeard
Biodiversit)' and Systematics
Department of Biological Sciences
University' of Alabama
Tuscaloosa, AL 35487

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

Dr. Paula M. Mikkelsen

Department of Living Invertebrates

The American Museum of Natural

Historv

New York, NY 10024

Dr Diarmaid O Foighil

Museum of Zoology and Department

of Biology
Universitv of Michigan
Ann Arbor. MI 48109-1079

Dr. Gustav Paulay

Florida Museum of Natural Histoiy
Universitv of Florida
Gainesville, FL 3261 1-2035

Mr. Richard E. Petit

PO.Bo.\30

North Myrtle Beach, SC 29582

Dr. Gar)' Rosenberg
Department of Mollusks
The Academy of Natural Sciences
1900 Benjamin Franklin Park-yvay
Philadelphia, PA 19103

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

Dr. G. Thomas Watters
Aquatic Ecology Laborator)'
1314 KinnearRoad
Columbus, OH 43212-1194

Dr. John B. Wise

Houston Museum of Natural Science

Houston, TX 77030-1799

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THEfcNAUTILUS

CONTENTS

\ oliniic 113. Siinihcr 2

Aiioiisl 6. 2001

ISS\ 0()2S-]3H

Patricia Mil()sla\icli
Pablo E. Pcnchaszadeh

A(li'l|)lii][ili,i'4\ .uiil t.iiiiiilialisiii (liiriiiLi; caiK (livflupiiiriil

(il C'nifihiiliiiii (luricnla (Ciniflin. 171)1) ((;astn)[)(i(la:

( ^aKptiacklac! Irom tlie Wiic/iiclaii Cariljlicaii

39

Cal•^• W. Schiiiclz

Till' laiiiiK Mitridai' l( ^astrnpi.da* in llir Enwcr Miin-cnc
( 'lii|iiila I'liniiatiiiii iil iiortlierii Flmiila

45

Cristian F. Itiiarte

I'isiiliiiiii chujuiiditinii new spctirs Irniii Santa ( .'ni/ dr la
Siena. |-i(ili\ia (l^i\al\ia: Spliacriitlac)

50

Francisco J. Garcia
Jesus S. Troncoso

Fdnniiiiis clcii/ilcxidr. a new species (( )pisllinliiaiieliia:
Aeiilidiiilae) liiiiii llie eastern Pacific Ocean

00

SiKia Macia

Fiipiilatinn iKiiaiiiics and <j;ni\\tli rate ul tlie tiirliinid

^astrnpod IJthopoiiui (i)itrnc(nnnn (Miilliiseai in 15isca\ne

Ba\. Florida, L'SA ' 62

John B. ^^isc

AnatoiiA (il Hddiicn jadisi lOlssun and \lc(.iiit\. 11)58)
(Heteriiliiaiieliia: l'\ lainidelliclaei liiini the western
Atlantic, with ciiniparisons to cither species in the 'j;enns

fiS

Notices 76

t/annc Eiologica' LabCHic,
WooCs Hole Oceanographic Insiiiuiioi^

I thr^ry

AUG 2 0 2001

Wooos no'e. iViM ivS43

THE NAUTILUS 115(2):.39-44. 2001

Wl'M- .'W

Adelphophag)' and cannibalism dnring earlv development of"
Criicibuhim auricula (Gmelin, 1791) (Gastropoda:
Caly][itraeidae) from the Venezuelan Caribbean

Patricia Miloslaxicli

Uni\'('isii!ad Sinujii BoliMir
Departaiiiento de Estudios

Aiiihientiiles
Apartado Postal <S9.000, C:aracas lOSO
VENEZUELA

Pablo E. Penchaszadeh

Museo Argentino de Ciencias
Naturales-CONICET-UBA

A\; Arit;el Clallardo 470

CUdo n|H BiiciDs Aires

ARCENTINA

and

UniMTsidad SiiiiDii 15(}li\ar

ni'partaiiieiitd de Kstudicis
Aiiiliieiitales

Apartado Postal SUIHKI, Caracas HlSd

\ENEZUELA

ABSTRACT

CiiK iliiihiiii iiiiriiiilii ((wiieliii, 1791) was loiiiid liMiii; attaelicd
to roekv substrates at Isla Caribe, Estado Sucre, N'euezuela.
between 0.5 and 1 m depth. The shell of sexualK mature te-
males ranged troni 12 to 2.3 mm in diameter and (i to 10 mm
in height. These females brooded between 4 and 20 fj^i^ cap-
sules in the mantle ca\itv. Each egg capsule luid a stalk .md
the stiilks were joined at the point of attachment to the sub-
strate. Each egg capsule measured between l.(i ami 3.5 mm
in length (without the stalk) and between 1.0 and 2.S nun in
width. The number ol nncleaxed eggs per capsule \aried be-
tween 55 and .305 and measured approximatcK 220 (xni in
diameter. A\\ eggs started development and complett'd the hrst
four divisions, but onK 7 ± 4 7c (between 3 and 24 eggs per
capsule) continued to develop. The remainder was ingested bv
the embnos as nurse eggs. At the end of this period ot adel-
pliophagv, the cmbr\os started to cannibaUze each other, after
whicliouK I to I 1 (lubAOS W'ere left in each capsule jmcniles
wllli a well-developed loot and short cephalic tentacles craw led
uiil ol llie capside. Cilian movement was obsened in the re-
gion where the velum had been resorbed. The shells are
brown, calcified, and measure about 7.30 jj.m in length. Both
intracapsular feeding strategies, adelphophagv and cannibalism
have been previonslv reported in the lamilv C^alvptraeidae.
Both strategies are here reported tor the first tinie in a species
of the n;emis Cnirihnlmn

Ailililiiiiuil kill iiiirds (,'aeno'j;astropoda. egg capsules, embiA-
onic nutrition. repro(.luctiou. nurse eggs, Venezuela.

INTRODUCTION

C;asti(i[i()(ls (il the laiuilv ( 'alvptraeidae are cluiiaeteii/etl
bv tlie production of eggs contained in nienihranous egg
capsules that are attached to hard substrates and ari'

brooded betxveen the neck .uid pi"o|iodiuui of the le-
inale. The spawn is composed ot si'veral sac-shaped egg
eapsides joined to each other at tlie iiase of a stalk. Each
sac is composed of a thin im-nibrane and the stalk is a
continuation of tliis niiMubrane. Within genera, some
species produce large eggs witli large amounts of volk
that hatch as crawling juveniles. Other species procluee
smaller eggs that hatch as planktonie veliger lanae and
others complete inli'acapsnlar develo[)ment bv feeding
on nurse eggs (Hoagland, 19Sfi).

The genus Crcpicliila Lamarck. 1799. is the best stud-
ied in the tarnilv. Hoagland (f9S6) reviewed encapsula-
tion patterns and brooding ol about 20 sjieeies of f";r-
jihluUi and fountl that egg di.imeler is veiA variable.
ranging from L30 to 440 (jim. Ilalclilings can be plank-
totronhic veligers, pi'divcligers oi' crawling juveniles. E.\-
tra-endm'onie nutrition, when present, can be in the
form of nurse eggs or cannibalism among sibling I'ln-
biAos. .\d(dpliopliagv has been reported in .it least 7 Crc-
pUliihi species and cannibalism oi damaged or abnormal
embnos is widespread in the gemis (Hoagland. I9Stt).

The genus ('nicihiihiiii Sclumiacher 1S17. is charae-
teiized bv cap-shaped shells with a completi' cup-like
support, which is attached in its base or along one side.
.\bbott (1974) reported 12 .\meriean species, 10 in the
Pacific from (.'alih)rnia to (liile and 2 .-Xtlantic .species,
Cnicibiiliiin iniriciilfi li'om South ( 'aroliua to Te.xas and
West Inilies to Brazil and C.nirihulinn strintiiiu Sav.
IS24, from Nova Scotia to both sitles of Florida. Di'az
and Pusaua (1994) ri'ported .3 species in the Colombian
Caribbean, C. iiiinciila. C. iiKirrnsc Weisbord, 1962. and
C. j>laiiiiiii Schumacher IS 17. Reproduction of species
in the genus (Uiirihuluiu is less known. Oie ( 193S) stud-
ied the se.xual phases oi Cnicibiiliiin spinosuin Sovverbv.

Page 40

HE NAUTILUS, W

15. Xo. 2

'I able 1. ('niiihtilnin tiiirifiil/i. (.'lianictcristifs ol oiiibnos
(liiiiii',; iiiliac.ipsiilar di'vclopiiK'nt. Xalucs represent mean -
SI) iiiniil)irs in parentheses indicate range.

I'"igui"e 1. Adnlt shell i>l Cniiihiiliiiii iiiiiu'iihi Diirsal and
\entral \ie\v. Scale bar = 13 nnii.

1<S24, from tlie coast of Sontlicin Cialifornia. Dexeiop-
inent oi Cni<ihiiliiin sculclldhiiii W'ooil. 1828. from tlic
Persian (hdf was studied In Tliorson (1940). Bandel
(1975) described the protocoiicli tA Cnicibiihim aiirifiila
(Gmelin, 1791) from the Coioiiiiiian Caribbean. Fen-
chaszadeii (1984) studied some aspects of the intracap-
sular de\elopment of Cnicihtiliiin iiuiri'iisc Weisbord.
1962, from Venezuela. Hoagland (198fi) ga\e inh}iiiia-
tiou on some reproductive as[)ects ol ('nirihiilniii prr-
soiiiiluin Keen. 1958, Cnicihiilinn sciilillaliini. C'niri-
huliiin spinostim. and Cnicihiiliiin lunhrclla Deshayes,
1830, from the Pacific coast of ( Central .America.

Most Cntcibiilinn species ineulioiK'd abo\e ha\'e
plaiiktonic de\elopment and hatch as \-eliger lanae (C
jHTsiniatiim, C. scutclliifuiii. C. spiuosmn and C uin-
hrclla reviewed bv Hoagland. 1986) or veliconch (C aii-
rirulti, according to Bandel, 1975). Up to now, the onl\
species knowni to lia\e non-planktf)nie direct dexelop-
ment is C. marcnsc ( Penchas/.atleli, 1984). Neither nurse
eggs nor cannibalism among embnos ha\'e been report-
vi\ in an\ ol the Cnicihiihiin species.

Herein we describe the egg capsules, intracapsular
de\('lopment of the embr\'os, and hatching mode ol C.
iiuiicida from Isla Caribe, luealed on llic northeastern
coast of Venezuela. The iclalinnship between female
size and the number and sl/e ol egg eajisules. size ol
the eggs and size ol hatchlintis was also deteiiiiined.

M/\TER1ALS AND .MKIIIODS
Si'i:c:iMr..\s

Brooding specimens were collected in March 1994 and
Februan 1995 at Isla Caribe, (ihacopala, northern .-Krava
Peninsula, Estado Sucre, \ene/n(la (10°42'ir'N,
fi3°52"57"Wi between 0.5 and I ni depth, '{"here, indi-
viduals ol Cnicihiilion aiirirnlii were found attached to
large flat rocks set in a patchv habitat of sand and Inille
griLss (Thalassia tcstii(liiiurn). Thev were easiK idenlilied
by the edge of the imier cup. which is enlireK Irei' (fig-
ure 1) and the outer shell, which is beige with coarse
rililxins (descriptions in .Abbott, 1974. Diaz and l'n\ana,
1994). Some of the indi\iduais were fixed iu a glucami-
ne-acetate (G.\) buHer contaiuinsi 4-6 '7( formalin (Mii-

st,

(.'haraeteristie>>

.Si/.e : p.m)

Egg

■T

I'.nilnro

l'"nil)r\(i

4

l*'.arl\ \('li<ier

\ 'e litre r

l'edi\i'li''er

Hatch

Unclea\ed vellnw egg.

Developing emhnos l)e<ri]i to
int^est nnrse eggs.

Nurse eg^ intrestion ends.
Cannibalism amoni; sihlini;
embrsos begins.

(aninibalisni ends. EarK
\cliger with round \-elnni.
small foot, e\es, short ,ind
thick cephalic tentacles,
organic matri.x of shell.

N'ehger with velum measuring
250 (jLui in diameter,
pigmented with wliite
spots, non-calcified proto-
conch, enlargement of foot.

Preliatchling with reduced
\eluni (less than 200 (jlui',
lonii and thin cephahc
tentacles, long foot with
edge pigmented (measures
1 mm), calcification ol shell
(veUowish) begins. Packed
N'olk in the p()Steri<ir repon
ol the animal

(.iawlinij |u\t'nile. absence ol
\clnm. cilia remain, loot
with two lobes in the
anterior part, caleifietl
brown shell, little to no \olk
resenes.

218.1 ± 16.0

( 195-236)

n = 99

362.4 ± S2.4

(236-472)

n = 99

615.7 ± 82.9

(472-786)

n = 105

701.0 ± 78.5

(511-786)

n = 69

726.1 ± 61.2
1629-825)
n = 64

740.5 ± 69.6
(629-825)

728.0 ± 70.0
(680-840)
n = 10

oski\ii'h and j'enchas/atleh. 1997' and ollieis wc-ie kept
alive ill tanks at 25-27° C and 35 Vcc saliiiil\ with aer-
ated, iion-eirciilating seawater.

\ouc1km- adult material ol C. iiiiiiciild i entire individ-
uals fixed in formalin and presened in ethanol) has been
deposited in the Museum of Comparative Zoolog\. Har-
vard Universitv. (Cambridge. Massachusetts, catalogue
number MCZ 304128.

I)l V I IDrNU'.NT

\\(' sindied 30 leiiiales ol C iiiiriciilii. Ill I'ach spawn we

observed: ( I ' the n Iier and si/e of egg capsules

brooded per leniale. i2' the nimiber and size ol eggs
and dcvclo|)ing eiiibrvos widiin ihe capsiili'. I'll the dil-
ferenl slau;es of dev (■lopinenl. and il' the hatching
mode. The .>0 females produced a total ol '■''t~-'i egg ea|i-
siiics; of these we raiukuiilv chose 125 egg capsules (be-
tween I and 10 from each female). Egg capsules were
se|)araled liiiiii the cluster, classified according to stage
ot di'velopnieiil and kept ill .lO ml beakers inside tanks

Patricia Miloslaxicli and Fahlo E. Pencliaszatli'li. 2001

Page 41

with aeratc-d and Millipore-filteivd (W'liatnian CI-' IV'
Sfawatcr. Filtered seaw ater was renewed daik. 1''.^^ cap-
sules were inculiated in tlie tlark at 2(i ^( .' in a I'reeision
(SI SI incnhator.

Ohsenations of ]\w and presencd malciial was duiu'
witli a ZKISS dissecting microscope and a ZEISS com-
pound microscope: measnreivients weic taki-n witli an
ocular micrometer. Results aii' leported as mean ± stan-
daid de\ iaiion,

RESULTS

The shell diameter anil height ol se\uall\ mature le-
males of C. auricula ranged from 12 to 23 nun ( IS. 4 ±
2.6 mm, n = 30) and from 6 anil 10 nun (T.fi ± 1.4
www. w = 16) respecti\el\-. A significant coirelation was
lomril l)et\veen female shell cliaTuetei- and capsule size
(Pearson correlation coefficient r = 0.54. p < 0.01). that
is. larger females brooded larger egg capsules. Howexcr.
no significant correlation was foimil hetA\eeu female di-
ameter and the numher ol capsules, eggs per capsule,
nnmlx'r ol hatchliugs. nor hatehliug size.

Di;\ Ki,()i>\ii;\r

Females hiooded hetween 4 and 20 egg capsules m llie
mantli' ca\it\ (12.4 ± 4.4. n = 30. Egg capsules hail a
triangular shape, with one corner extendiTig to form a
stalk, and were attached to the substrate at the base of
this short stalk. The stalks are flattened onK al the base
ol the capsule and the rest is thread-like, I'lic (.'apsnle
walls were thin, fragile, transparent (figuri' 2), and the
egg capsule was turgid. No exit plug was obsencd. Ivgg
capsules measured bet^\■een 1.6 and 3.5 nun lengthwise
(without the stalk) (2.5 ± 0.3 mm, n = 125) and 1,0 to
2.S nun wide (2.1 ± 0.3 nun. n = 125).

The number of eggs per capsule \aried from 55 to
305 (161 ± 56. n = 56). Unclea\ed eggs were \ellow
and measun'd approximately 220 (xni in diameter (figmc
2). All eggs undeiwent the first four clea\ages, 93'7( ai-
rested dexelopment at this stage and were ingested b\
the lemaining de\eloping cmbixds (between 3 and 24
per egg capsule, 11.0 ± 6.0, n = 21) as nurse eggs.
Within the capsules from a single female. embiAos de-
\eloped ssnchronousK'. Gastrnlation is accomjilished
when tlie ectoderm undergoes epiboK from the animal
pole ami en\elops the other cells of the enibixo, l)e-
\elopiug embixos measmed about 362 jjim when th<'\
started to ingest the whole niu-se eggs, the\ weie o\al-
shaped, \ellow, with no \elar lobes nor shell (figure .3).
M the end ol the ingestion stage, these emimos mea-
sured about 616 |jLm in length; tliev were \ellow and the
nurse eggs were tightK packed in their interior. M this
point of their dexelopment, the embnos started to lau-
nilialize each other. It was common to find two embnos
feechng u[)on a third one (figure 4). Alter (he tannibal-
ism stage, 1 to 11 embnos lemainrd m each capsule
(7.0 ± 3.0, n = 21) and de\elopi'd the \elar lobes. The
early intracapsular veliger measured about TOO [xm and

was chaiacleri/ed b\ a small round velum, a sn\all loot.
e\es. \en short and thick ci'phalic tentacles and sin-
roimded b\ an organic matrix (figure 5). A fragile shell
nieasuiing aroimd 72() fj.m developed from the organic
matrix. The develojied velum measured 250 fim in di-
ameter and liad white spots (figure 6). The intracapsular
pediveliger slau;c had a reduced velum measuring less
tli.m 200 |jLm iu dianielei-. Iiad elongated and thin ce-
phalic tentacles, and a long foot (about 1 umi) with
blown lines aioiiu the edges. When cakificatiou started
(figure 7). the pediveliger shi'll measured about 740 |jLni.
Pediveligcis were not able to swim wlien excapsniated
into seawatei'. A lew davs prior to hatching i6 to 7 da\s),
volk was obserxi'd inside ol the pediveligers. These volk
resenes disappeai'ed almost con ipleteK b\- hatching, and
onlv a small vellow s|)ot m the |)osti'rior region oi the
ammal remained. Hatching occnrri'il as crawling ju\(>-
niles with a brown cakified shell measuring 730 |xni in
length, the slicil w.is velativelv smooth with thin hori-
zontal lines separali'd 20 |j.m liom each other, juveniles
had a vvell-deveiope<l loot with a small and transparent
operculum, shoit cephalic tentacles anil novelum. How-
ever, cilian movement was obsened in tlie region where
the velum liad becTi lesoiiied itabli* 1).

i5is(:ussi()\

rhe relationship In-tween leinale size and some repro-
ductive parameteis such as i-apsule size, number ol eggs
and developing embnos within each capsuk-. and liati-h-
ing size of the iuveniles has l)ccn rcjiorted in seveial
lamilies of marine gastropods uSpiglit ct ill.. 1974. in
Mmicidae: Robertson. I9S3. in E[)itouiiilae: Miloslavicli
and Dntiesne. 1994. in Huccinidae). These studies in-
dicate that laiger lemales prodnie bigger egg capsules
that contain more eggs and more and i)igger hatchlings
than smaller lemales. In Crucihuhun auricula we found
that larger females brood larger egg capsules than small
lemales: however, these egg capsules do not i-outaiu
more eggs nor produce more or bigger hatchlings, there-
lore the reproductive outcome ol both small and large
lemales is the same, ('lia])arro ct al. (1999) found no
lelatiouship between leniale size and the numljer of de-
veloping embnos (A ('rcj)i(lula dilatiila Lamarck. 1S22.
however, the mean size of the juveniles at hatching did
increase with the shell length oi the lemale, because
more uuise eg^s weie available to an embno Iroui a
larm'i' li'iiiaie than to an embno Ironi a smaller one.

In lalvptr.ieids, the nuTiiber ol egg capsules per brood
is iisnalK related to lemale siz.i' (see table 3 in Iloagland,
19S6), Cnicihulinu auricula is one of the smallest spe-
cies of Criiiihuluui. 4lie nnmbci' of egg capsules (mean
12 egg capsules per brood' in this species is one of the
lowest amon<4 the live speiies ol the genus for which
there is information on reproduction published. It is
similar to that of (,'. uuircusc (about 14 egg capsules) but
it is lower tliiiu (,', sculcllaluni. (.'. spinosum (both with
a mean ol 20 ca])sules). anil C. pcrsonatum (30 cap-
sules!. I'lie piodnction of hatchlings of C. auricula is also

Paire 42

THE NAUTILUS, \ol. 115, No. 2

Figures 2-7. Iiilrac,i|)siiliir developnient a\' Cniciliuliiin (iiirii-iilii. 2. (icncnil xicw ol the (■";>; capsules with uiic-k'a\X'tl vSii^ (stasle
1). SfiiU^ bar = 1 .5 nun. 3. Mijg capsnie containint; both dcNclopini; cnihnos ilarjir! and nursi' Ci^ifs (small) lstam- 2: adclpliopluii^ ).
Sc;ile bar = fiOO |j.ni. 4. Canniljalisni anions siblinsj cnibnos (staijc 3). Scale bar = 200 |xni. 5. Ivirly \eliger inside egg capsule
(stage 4). Scale bar = fioO (j.ni. 6. Wliger with non-ealcilied protoconcli (stage .5). Scale bar = 250 jj.ni, 7. Prehatching (stage 6),
Scale bar = loO \xm.

Fatrifia Miloslax ieli and Fahlo E. PciR-liaszadeh, 2001

Page 43

Table 2. Brood characteristics and total production of li.ilcliliii'^s witliin the neims Cnirihuliiiii iNA = nol .i\ailalil(

Species

K'«

size

(j.nit

Xo.

capsules

Xo.

halclilin

apsule

jj Total hatchlinii
production

Hatchlin<j size
(jxm.i/niode

Source

C IIUIITIIsi

XA

1)

14.5

203

1020-1060

Penchasza

leh. 19S4

C. pcrsDiKitiim

XA

:59

275

S25()

Crawler
320

Iloanland,

I9S6

C. saifclhitun)

XA

20

200

400(1

\eliuer
XA

Iloaiiknul.

19S6

C. spiiiiisiiiii

NA

2(1

200

4000

N'elisjer

2S0

\i -liner

440

\eli14er

72S

Crawler

Hoanland.

lOSfS

C. niiihrillii

NA

31

150

4(i50

Hoagland.

19S(i

C. (iiiriciilii

218

12

I

84

This work

ver\" low (about 84 crawiine; jmcniles), followed li\ C
marense with a total production of 200 crawling juve-
niles. The other Cnicihiilinu species produce veliger lar-
vae in numbers o\er 4000 (Moagland. 1986) (table 2),

The size oi the uiicleaxed egg is \er\' important be-
cause it determines the amount ot \olk asailable for tlie
embiyo to dexelop when other extraembrvonic food
sources such as niu'se eggs, cannibalism among siblings,
or intracapsular fluid, are not a\ailable. The egg of Cni-
rihtiliim auricula is within the size range of caKptraeid
.species in the genus Civpidula. whicli \ai-\ betA\een 150
and 420 |xm (Hoagland, 1986). Witliin the genus Cru-
cibuluni. the egg size has been pre\ioiisl\ reported oiiK
for C spiuosuni (from 170 to f90 |xni; Coe, 1938).

\\ithin calvptraeids and particularK in the genus Crc-
pidula. iwt) feeding modes during intrat-apsular de\el-
opment ha\e been described: adelphophag\, or ninse
egg ingestion, and cannibalism: liotli were obserwd in
C. auricula. Hoagland (1986) reviewed the patterns ol
encapsulation and brooding in calvptreids and reportitl
no nurse eggs in the five Cnicihulum species for whicli
we have data (C. pcrsonatuui. C. scutcllatuin. C. spiiios-
um. C. uiuhrcUa. and C. marcnsc). However, Coe (1938)
reported that a considerable proportion ol tlic <'ggs of
C. .spinosuin fail to complete tle\elopnicnt, as the iiimi-
ber of veliger larvae that hatch is much lower than the
original number of eggs. This difference between n\im-
ber of eggs and number of hatching veligers could be
due not oiiK to mu'se eggs but also to embnos ingested.
Thorson (1940) reported that the large size of the eni-
brvos at hatching (0.7 mm) in C. scutcllatuin from the
Persian Gulf was pi'obablv due to cannibalism and that
mu'se eggs do not seem to occur If properK idi'iitificd
In Thorson, this species woidd have a \er\ wide geo-
graphic range e\en for a species with planktonic de\cl-
opment (Hoagland, 1986), since Keen (1971 ) ami .-Xbbott
and Dance (1982) reported it as a tropical western
.\meiicaii species ranging from .Mexico to Ecuador Pi-n-
chaszadeh (1984) stated, for C lunrcnsc. that presum-
ably neither nurse eggs nor cannibalism are inxoKcd
during development given the uniformitv of shell size at

hatching and bec-aiise inide\cloped eggs were not ob-
sened.

Bamlel (197.5; 19761 reported that C. auricula from
the Colombian Caribbean proiluces 10 to 12 vcllowish
eggs and tliat ;dl develop into a veliconch possessing a
functional foot and a large velmn. His observations were
probablv carried out after the developing embnos fin-
islied eating the nurse eggs, which could h,i\c led liiiii
to misinteipii't the embnos as eggs and to not obsene
the 150 nurse eggs per capsule we obsened (93 7c nurse
eggs), -\notlier possil)ilitv is that he v\(n'kcd with a dif-
ferent species. In the Colombian CJaribbeau, Di'az and
Pin ana ( I994i have reported three Cnicihulum species:
C. auricula. ('.. luarcu.sc. and C. planum, the first hvo
are relativcK c-ommon and the third one is uncommon.
The shell of C. marcn.sc is simikir in size to C auricula
(15 mm in diameter) but is i-haracterized bv a ciined
apex: the shell of C. planum is much larger (40 nmi in
diameter) and is characterized In brown reddish spots,
(uven that the taxoiiomv of all calvptraeids, especiallv
Caliiptracd i,amai-(-k. 1799, and Cnicihulum is on need
ofinajor revision, Bandel probaiilv misidentified the spe-
cies. However if Bandel's obsenations are correct and
111- pidpeilv identified the species, this would lie a
iiiiiinie case of poecilogoiiv among gastropods, which, we
suggest, is imlikelv to liap]ien (see reviews bv Boiichet,
1989, ami Hoagland and l^obertson, 19S8i.

p]inbi"vos of C. auricula di'velop svnchronoiislv within
the capsules of a single female. However they reach
variable sizes (between 680 and 840 fjLin), probablv de-
pending on the iiimiber of nurse eggs and emiinos in-
gested. Hoagland (1986) reported diat. for most Calvp-
tiiUMclae. lanae develop sviK-liidiioiislv, exi-ept in those
siiecies with nurse eggs: and in Cnicihulum .spino.sum.
there is asvnchronv between egg capsules, imjilving tliat
dii- brood does not hatch all at once.

Most Cnicihulum sjiecies herein mentionetl {C. pcr-
.•ionatuin. C. scutcllalum. C. .■ipino.sum. C. umhn-lla)
hatch as free-swimming veligers (Hoagland, 1986: (^oe,
1949). Peiicha.szadeh (1984) reported crawling juveniles
for C. inarcii.se and Bandel (1975; 1976) reported veil-

Paee 44

THE NAUTILUS, \o\. 115. No. 2

foiich.s for C. aiiriciilii in the (-oloiiibian ( 'ai ihlicaii in
tlii.s stucK', we foiiiRl thai ('. (iiuicnlit lialilus as ciawliiig
ju\(Miil('s, witli some velar (ciliar) remains Imt are unable
to swim. Even in tlie absenee of a Iree-swimminw sta<2;e,
this speeies has a wide geographic distribution in tlie
western .Atlantic, liom tlie coast ol South Carolina south
to the Caribbean to tropical Brazil (Abbott, 1974: Dfa/
and Puvaua. 19941.

ACKNCWLEDGMENTS

We wish lo ihank Oliver Contreras tor his research as-
sistance, both in the field and in the laboraton'. We are
also indebted to Claudio Paredes and Ana Karinna Car-
bouini lor their \aluable lielp in the lalioratorv. We sin-
cereK' thank Dr. Diarmaid (J Foii;hi!, Museum of Zo-
oiogv', Univensitv of Michigan, and Dr. Roberto Cipriani,
Unixersidad Simon Bolivar, for their comments on an
earlier version of the manuscript. This work was sup-
ported in- a grant of the Decanato de Imestigacion \-
Desarrollo to the Grupo de Ciencias Marinas (Uni\er-
sidad Simon Bolivar). We also thank l^r [ose H. Leal,
The Bailev-Matthevvs Shell Museum, loi- the photo-
graphs in figure L

LITER.\TURE CITED

.Abbott, R. T. 1974. American seasliclls. 2ii(l ed. \"an Nostraiid

Reinliold. 66.3 pp.
\liliolt, II. T. and S. P. Dance. 1982. (>)nip<-ii(liuni (il seashells.

Odvssev l^ublishing, Hoiiii Kong, 41 1 pp. ( 199S printing)
Baiidel, K. 1975. Das Elinlinonalgchansc kaiiiiischer .Meso-

und N'eogastropoden (Mollusca). Akadeinie der W'issenss-

chaltcn inul der Literatur, Mainz, 1: 1-133.
liandcl K 1976, Ohseivations on spawn, enihnonic develop-

uiinl ,uid ccolog\' of some Caribbean lower mesogastro-

poda. The Veliger IS: 249-271.
liouchcl, I' 1989. \ review of poecilogonv in <j;astropnds. |oin-

nal ol Molluscan Studies, 55: 67-78.
(:li,.parro, (). H.. H. V Ovar/uii \ \1 Xcr^ara anti H. J.

Thompson. 1999. Energv inxcstment in nurse eggs and
egg capsules in Crepicluhi dilatata Lamarck (Gastropoda,
(^ahptraeidae) and its influence on the hatching size of
the juvenile. Journal of Experimental Marine BioloijA and
Ecology 232: 261-274.

i'.ov. \\. K. 1938. Se.xual phases in the gastropod C.nicihiiliiin
sjiiiHtsiim. |()unia! of .Moiphologv 63: 345-361.

( !oe. W. K. 1949. Divergent methods of development in inor-
]iliol()gicallv similar species of prosobranch gastropods,
loiinial of Moiphologv 84: .383-.399.

Di.iA. J. .M. and .M. Pu\ana. 1994. Moluscos del C;anhe Col-
omhiano. Un catalogo ilustrado. 1 era ed.. Colciencias-
Fundacion Natura-INN'EM.XH, 291pp.

Hoagland. K. E. 1986. Patterns of encapsulation and hmodnig
in the Cillvptraeidae (Prosobranchia: Mesogastn)p()da'.
American Malacological Bulli'tin 4: 173-183.

Hoagland, K. E. and R. Rofiertson. 1988. .\n assessment ol
poecilogoin in marine invertebrates: phenomenon or faii-
ta.sv':' Biological Bnllctin. 174: 109-125.

Keen, .\1. 1971. Sea-shells oi tropical west .America (.Marine
mollusks from Baja California to Peru). 2nd. Edition.
Stanford Universitv Press, Stanford, 1094 pp.

.\lilosla\icli. P. and L. Dnfresne. 1994. Development and effect
of femafe size on egg and juvenile production in the neo-
gastropod Biicciuinn ri/tiiifiiin from the Saguenav fjord,
(lauadian jouni:il of Fisheries and .■V(jnatic Science 51:
2866-2872.

Miloslasicli, P. and P. Penchaszadeh. 1997. Spawni and devel-
opment of Ftisiiins flostcr Philippi, 1850 (Gastropoda:
Prosobranchia) ln)ni the X'enezuelan Carihliean. The \ e-
liger 40: 93-100.
Penchaszadeh, P. E. 1984. Direct development in Cnicibti-
liiDi //irtrc/i.sf Weishord. 1962. (Clastropoda: Calvptraeidae)
from Golfo Tiiste. Wnezuela. Journal of Molfnscan Stud-
ies 50: 237-238.

Spight, T M.. C. Birkeland and A. Lvons, 1974. Life histon
of large and small nmrexes (Prosohrancliia: Mniicidae).
Marine Biologv' 24: 229-242.

Robertson, R. 1983. Ohsenatioiis nl tlic lile histonol thevvcn-
tietrap Epitoniiim alhUhnn in the West Indies, .\niencaii
Malacological Bulletin. 11-12.

Tliorson, G. 1940. Studies on tin' egg masses and lanal de-
velopment of gastropods from tlie Iranian Gulf Danish
Scientiric Invfstigations in Iniii. part II: 159-238.

THE NAUTILUS 115(2):45-49, 2001

Page 45

The family Mitridae (Gastropoda) in the Lower Miocene
Chipola Formation of northern Florida

C.iiv\ \\. Schinelz

5575 I2tli A\L^niie Si)iitli\M"st
Naples, FL 34116 USA

AHSTHACT

A brief R-\ie\\ cil tin- Mitridae trom tlic Limcr Mnnciic (^lii-
poia Foiiiiatidii ol iioitiiern Fliirida is pa-st-iiteil. willi tlii' di-
scriptioii ol t\Mi new species and the ideiitifieatiDii dI adult
specimens ot Mitrii dctro^li/phti Gardner, 1937, 'I'lic new spe-
cies are Mitra ciilli(nincnsis. a common, lars;e liisilonii species
tliat bears a resemblance to Mitra fn.sifdnnis (Brocclii, 1814)
from the Miocene ol Europe, and Scahricohi clajxiliiiiii. a rare,
MiKiolli-sliell, species that is limilcil Id the anncnl ni-l cnxi-
Kiiiiiieiit aloni^ Tenmile Creek and the ( 'liipiii,! liiMi

.\ilililu>ii(il kill nunls: Neogene, Milrn. Sfiilii-iculii '/Jim

ixth()]:)U(;tion

Accdrdint;; to C'cni(iliinsk\ (197fi), the faniiK Mitiidae is
characterizetl 1)\ "fusitorni, eloritiate-cAate or t\lin(lrieal
shells which liaxc conwx or annulate whorls. iisualK a
ii.uidw apeitiiic, Iroin 3 to 1 1 lolds on llic columella
and a distinct siplional notch. Members ol this hnniK
are found in tropical to temperate waters throughout the
wdild where tlicN thrive' in intertidal areas down to
ilepths not e.xceeilini; 1465 m (CA'rnohorsks, 19761. The
first lossil exiclence of this faniiK' appeared in Uppei'
Cretaceous deposits and numerous other fossil specie's
have lieen collected and deserilied lioni I'ertian depos-
its throu'^^liont ihe woild (( 'ernohoiskx, 1976).

Gardnei" ( 1937) reportetl 16 species ol .Mitridae from
the Alum Blnll Ciroup of Ploriila, "with probabK as
nian\ nioie repre.sented bv material too imperleit to de-
scribe.' ( )1 the 16 Alum Bluff species, ( Gardnei' listed 1 1
Ironi the (diipola Formation. Ol these, ei<i;ht are now
classified in Costellariidae and one, Milni i Plciopti/iii/-
ina) prodroiiia Ciarduer, 1937. has been placed in the
PlcioptNginatidae iQuinn, 19S9). CnrrtnlK. onl\ '/Jlxi il-
hicklata (Wbodring, 192S) ( = Mitrii iiiitnulilii ( iardnei;
1937) and Mitra (ictvoijl\jph(i (Gardner, 193)7 remain in
the Mitridae.

The objectixes ol tliis paper ai'e to: (1) describe two
new species of Mitiidae from the ('liipola i'"ormation ol
northern Florida aiul, (2) update the taxmioniie status ol
the t\v() remainimj; species from the (Inpohi l''oriii,ilioii.

Tvpe and fi<j;ui"ed specimens referred to in this paper
are deposited in the paleontological collections at tlie

United States .National Miisiami, Sniitlisonian Institu-
tion (USX.M) and the Florida ,Muscami ol Natural Ilis-
ton-, llni\ersit\ of I'lorida lUl-").

SYSiKMATK S

FaniiK Mitridae Swaiiisoii. IS.31
(Melius .U/7/V/ i„iniarek. 1 79S
Sul)ii;emis .)/;//■(/ i,amarck, 1 79S.
Mitra {Mitra) tiftin^liiplia (iardiier, 1937
(Figures 1-3)

Description: Shell lusilorni-o\ate. inoderati'K heaw.
i^rotoconch iisiialK eroded. Sutures deepK impres,sed.
Spire with 5 i()ii\e\ wlioils sculpliir<'d with punctated,
e\enl\' spaced sjiiral gidoves. Fast whorl with 20-2S spi-
ral grooxes. penultimate whorl with 7-10. and remaining
whorls with 6-7. .\pertnre narrow, more than half shell
k-ngth. Outer lip thin, not lirate witiiin. Parietal wall \t']\
thin. C'ohiniellar lolds 4-5, sluupK elexateil, obli(]ue.
e(|iial]\ spaced, inereasiii',^ in prominence in post(>rior di-
rection Anterior canal modeialcK long, not shaipK' de-
hued. broailK emarginate al the e\ti'eiiiit\; anterior las-
ciole slightK swollen

Nfaterial examined: FF 95999. length 41.3 nun.
width 15.1 mm; UF 96000. lengtii 43,9 mm. width 15.1
mm.

Type locality: USNM locality 2213, 1 mile below Bai-
le\s I-'ern. ( Inpola liixcr. (.'alhoiin Ooimtx. I'lorida.

Distribution: Milrn (irlro^h/jilia is a inoderateK rare
spc'cies that has been loimd at two (,'liipola Formation
sites along 'i'emnile ( a'eek in (!alhoun (.'ounty Floriila.
and at a single site along i''aile\ ( a'eek in ("alhoun (3oini-
t\. .According to N'okes ■ I9S9 . the sites along Tenmile
( a'eek probabK represent an .incieiit reel en\"ironnient
while those al Farlex (,'reek suggest a back-reel habitat.

Discussion: ( iardner ( 1937) described M. actco<Jiipha
Iroiii a single, broken, jux'enile specimen (figure 1 ),
Since the ju\eiiile specimens ol all three sjiecies ol mi-
trids examined lor this paper had spiral grncnes similar
to M iii-lroij^liijilia. it IxHaine iux'essar\' to obtain the
t\pe speeimeu and compare it to material from the F"lor-
ida .Vlnsenm of Natural Histon and the imestigators

Page 46

THE NAUTILUS, \oI. 115. No. 2

collt-ctioii ill orilcr to (Iclcniiinc wliicli iil tlic adults liorc
till' greatest siiiiilarit\ to tlic juscnile de.scribecl by Uard-
ner (1937). .\f'ter earefiil coniparison of the t^pe with
rccentlv collected adult and jiiNciiilc spcciiiiciis, it was
apparent that the iiiitiid illustrated in (i<j;nres 2-3 was
thi> adnit form ol \l (irlc(>^li/i)liii This decision was
based upon the Fact that both had similar shell sliapc-s.
the whorls ot both bear the punctated spiral sculpturing
described In (iardner ioji. cit.l and both had a similar
number oi Coluinellar folds. The specimen illustrated in
figures 2 is most similar to the holotxpe. Like the ho-
lot\pe. it has 5 columellar folds and nearly the same
number ot punctated spiral grooxcs on the last whorl
(26 \ersus 2S in the liolot\pe). The specimen in figure
3 has onK four columellai- folds and about 20 punctated
spiral grooves on the last whorl. Further examination ol
3 partiallv broken adult specimens iu this investigators
collection revealed that all had 4 columellar folds.

.\mong New World fossil niitrids onlv the Miocene
MHia (ili)ia^rensi.s (•oraUiophila Olsson. 1922, from
northern Peru, bears some resi'uiblance to M. (ictvonhj-
pha. Olsson (1922) described M. ciirdllidjiltild as a slu'll
seulptin'ed with slightlv elevated spiral cords separatetl
b\ finek sculptured gr(.)o\'es. In M. (■onilli(>j)liil(i there is
a Icjtal of five spiral cords on each ol the whorls except
for ihe last, and the columella beais 3 [)roiuinent folds.
Milrii iirlcc)^liij)h(i differs trom ,)/. coralliophila h\ its
larger size {41.3 nnn and 43.9 nun for the adults versus
17 mm), larger nmnber of spiral cords ( 10 \ersus 5), and
its larger number of columellar folds (4-5 \s. 3).

Because of strong similarities between these two spe-
cies. Cernohorskv (1976) believed that .\/. actcoiiUji>1t<i
mi<jlit be related to the recent Milni Ixirlxiilcnsis (Gme-
liu. 1791 i. llowi'xcr, M. haihiidciisis entireK' lacks the
punctated spiral grooves that characterize M. (irtco^hi-
plia. .^niong extant species, .\/. arfro^li/pha seems to
bear a stronger resemblance to the Indo-Facific Mifra
(imhioua .Svvainson, 1829. A compaiisou ol these two
species shows similar shell uiorphologies, with shells of
the two species showing punctated spiial groo\<'s. How-
ever, M. acteo'^hjpha is a smaller spei-ies (43.9 nnn
lengtli vs. 53.4 nun) with fewer columellar folds (4-5 \s.
5-6) and a fewer number of piuielaled spiial grooves
(20-28 vs. 25-35).

Subgenus l-itsinutnt Conrad. 1855

Milra {Ftisiiuilra) callKuniciisis new species

(Figiu-es 4—5)

Description: Shell slender, large, moderatek heaw.
fusiform-ovate. Protoconch usnalk eroded. Suture deep-
ly impressed. Spire with 8-9 weakk convex v\liorls I'irst
six teleconcli whorls sculptured with 7-8, eveulv spaced,
pitted spiral grooves. Pitted spiral scnljitnring on re-
maining teleconcli whorls lainl. disappearing near the
aperture. .*\pcrture slightlv shorter tliau s|)ire. narrow
and elongate, sinootli within. Clohnuella with ihm ( .illus
ill adult specimens and 5-6 strong, obli(|ne. e(|ualk

spac-ed folds. Siphoual laseiole straight, siphonal notch
distinct.

Ilololype: UF 9599S, length 92.1 nnn. width 24.3

Paral\pe.s: Paratvpe A. UF 87784, incomplete speci-
men, length 60.5 nnn, width 18.3 mm, (Ihipola Forma-
tion, north bank of Tenmile Creek at powerlinc crossing
about one mile west-northwi'st of month of creek at
"Bailevs Feny" (SE 1/4 See. 12, 4 1\, KIOWi. Calhoini
(;ount\\ Florida (Tulane Universitv localilv TU 830),
I'aralvpe B, UF 89579, incomplete specinien, length
62.0 mm, width 20.8 iniii. From tvpe localitv.

Type localilv: (^liipola Formation, Tenmile Creek,
about 1.75 miles west of Chipola River (NE 1/4 Sec. 12,
TIN, KlOW), Calhoun Countv, Florida (Tulane Univer-
sit\ loealitx TU 546: = USCiS 2212, "one mile west of
Bailevs Ferrx").

Distrihiilion: Onlv one complete specimen of .V. cal-
luiiiiicnsis is known. However, nunierous fragments of
this shell art' present in the Tulane University Collection.
now housed at the Florida Museum of Natural Historx;
(Jainesville. In addition, the investigator has obtained
unmerous shell fragments over the last ten vears from a
broad range of C.hipola Formation collecting sites along
Tenmile and Farley creeks and both banks of the Chi-
pola Hivcr.

Et>molotf\-: Named after Calhoun CoimtA'. Florida.

Discussion: l^'rom the examination of unmerous shell
fragments it is evident that M. ciillutiiiiriisis attained a
very large size with manv specimens reaching total
lengths iu excess of 110 mm. In general shell shape and
size of this species bears a strong resemblance to the
living .Mediterranean Mitm fiisifonnis zoiialii Marrvat,
1818, and its fossil coimteipart Milra fiisifonnis fusijor-
mis (Brocclii, 1814) from the Miocene and Pliocene de-
posits of Portugal, France, Italv, Austria. Ilimgai-v, Po-
land, England, and the Island ol Hhocles, ll is verv likely
that .\/. flisifiiniii.s and M. zoiuilii share a coiimion an-
cestor with M. aillHuincnsis.

Milni fusifiiniiis can be distinguished from .\/. caUiou-
iiciisis bv the more slender shell and ]ireseiice of well-
developed [litled spiral grooves on the first five teleo-
eoncli whorls ol the latter species, .\iiiong New World
Neogeue s|ieeies. .\/ calliouiiciisis bears some resem-
blance to Milrii tiltiii i(;abb, 1873) from the Pliocene
deposits ol the Dominican Hepnblic as well as the Late-
Pliocene species, Milni (CDoc/n/ig; Olsson, 1964, from
the I'lsiiieraldas l'"ormatiou of Ecpiador. and Milrii
siniiiisdiiii ilniihiiri Olsson. 19.32, from the I'lioeeiie
Tunibe/ Formation of iiorthc'iii Pi-rii. 15olh .\/ lildii and
M ciillioiniciisis are lusilorm-elongate shells reaching
lengths in excess of I 10 mm with the carlv teleconcli
whorls having disliiui spiral sciilptniing, llowever. .)/,
nilhoiiiiriisis has ,i thinner lip and has 5 (i columellar
folds. Ill addition, the spiral sculpture on the eaiiv
v\li(irls of .\/ liliin li.is line spiral grooves with evei'x

Can W. Schmclz, 2(){)1

Page 4'

(ourtli one deeper (Pilslin-, 1922). while in .\/ nillii)ii-
(ic/i\)s tlie spiral grooves are all tlic same deplli on tlu'
earl\ wliorls.

Mifrii swaiiisoiiii iliiulxiri ami M ini()(/r/;ig; are \\n\
similar and nia\' represent the same speeies. This simi-
lai"it\ was noted li\ Om()horsk\' ( 197fS). who eonsidi ic-d
M uoddriiii^i to lie eonspeeifie with M \niiiit\oiii iliiii-
hiiri. Ahhott (1974) also listed M itdddriii^j^i as .i s\no-
iiMii nl' M/tra swaiiisouii swaiiisiiiiii (Broderip, hS.'5(i), an
imeonnnon Reeent "snhspeeies" that li\es in deepwaler
Irom Sonth (Jaiolina to the Wist Indies, Milm cdllKUi-
iirnsis is similai' to M. uuodriii'^i and M mi iiiii\i>iii diiii-
lifiii in thai it is a lusiforni-elongate shell widi dislinet
spiral senlptnring on the earK wlioils. llo\\e\er. unlike
the latter, M cdllKnntoisis lias a less stont and twisted
anterior eanal, it has 5-fi eolnmellar lolds. and has fewer
spiral groo\es on the apieal wliorls.

Among extant taxa. M. cdllioiincnsis is also similar to
Mitni swaiiisonii antillciisis Dali, 1S(S9. Cernoiiorsia
( 197fi) listed this as a deep-water snhspeeies that oeenrs
Irorn Nortli t'arolina to Yucatan and the .\ntilles. IJke
M. calhoiDwiisis. M. swainsonii (Jiitillcnsis li.is 5~(i eol-
nmellar lolils and has well-de\ eloped spiial senlpture.
However, M. calluniiicnsis has a more slender shell
shape, more dt'i'pK' impressed sutures, tewer spiral
grooves, and the spiial seulptnring disappears near the
aperture. It is possible that A/. (y;///ii/(/ic//,s7s is the lossil
aneestor to the present da\ M swcdnsoiiii eomplex Irom
the Carihhean pro\inee.

New World ancestors to the fusiforni-shajied M I'lil-
liiiniiiii\i\ date liaek to tlie Cretaceous. Tlu'se imliide
['alcufnsiiiiitrfi cloi><s,ata Sold, 1963, from the C^retaceons
Hiple\ Formation of Mississippi (Sold. I9fi4), Fiisiinitra
inilli}iiit(>ni (Conrad in W'ailes, 1S54) from tlu' l'"ocene
MoocK's ]5rancli Formation in Mississi[ipi (Doeki-n,
1977) and Mih-ii coiujiiisita Conrad. 1SS.5 li-om the Low-
er Oligocene of the X'icksliuig (Iroup in Mississippi
(MacNeil and Docken', 19S4).

SiilifamiK Imliricaiiinae Troscliel. 1 Sfi7
Cienns VaIhi II. and A. Adams. 1S5.3
Ziha illacidatii (Woodring, 192S)
(F'ignres fi — 7)

Mitra (Tiara) hcnckcni iUacidata Woodring. 192.S. pp. 243. pi,

14, fig. 13 ("Miocene" [Late Pliocciicl Howdcn Foriiia-

lion. Jamaica):
Mitra I Tiara) luilmdita (;ui(lncr 19)57. pp. 4(IS 4(IV), pi, 4S.

figs. il)-ll ("t iiiife lielow Bailc\'s FcriA. (Iiipnl.i Hiwv.

( )allHiuTi ( )ninit\. Fla, ',

Description: Shell fusiform. iiiodeiateK slender. Pro-
toeonch gloss\, witli 3 whorls. Sutures deepK mipressed.
Teleconch whorls 7, con\ex. sculptured with strong spi-
ral cortls and axial threads. Four spiral cords on the sjiire
whorls. .Aperture narrowK lolnite. alioni li.ill shell
length. Outer lip thm, the margin creiiale in h.iiinonx
with the external sculpture. Parietal wall think gla/<'d,
Colnmellar Folds 3, ohlicjne, e(|uall\ spaced, jiosterior
fold more higldv elevated. Siphonal notch narrow, deep.

Hol<>l\pe:

mm.

T\pc loc'alil\:

maica.

ISWi :5(i9i:i:i, length 10 mm. width fi.7
■ate Pliocene Bowden Formation. |a-

Distriliiilioii: liowdeii h'orniation in |ainaica and Clii-
pola Information sites along lenmile ( aei-k. l'"arli'\ Oreek.
and the ()hipola Hivcr in northern Florida.

Remarks: 'Ailni ilhiiiddlii helongs to a grou[) of iiiod-
erateK small mitrids that are scnljitnri'il with a presu-
tural carina, spiral grooxcs or I'oi'ds and axial lirae in the
inttaspaces. Cernoliorsk\ i 1991 ' tentafiwK jilaced nii-
tritls with these characteristics in the separat(> genus
/Jhti until the radnla of fixing spi'cies could he examined
and compared with those lielonging to the genus ('an-
rilla.

Ziha illaciilala was descrilied li\ Woodring < 192S)
from the Late-Pliocene Bowden I*"ormation of Jamaica.
Later Gardner ( 1937) found a similar species in the early
Miocene Chipola Formation tleposits of northern P'lor-
ida and named it Milm iiiilrodila. Ceriiohorskx (f99f ).
alter examining liotli Milm illacidala and Milm iiiilrod-
itti stati'd that hoth shells are identical and, irrespeeti\'e
of the age difference, listed .\/. milrodiUi in the sxnon-
\'im ol the geologicalK N'ounger .\/. iUacidata. This ar-
rangement is followed herein and. for comjiarison jiur-
poses, the holot\]ies of .\/ iUticiilatn and .\/, initrndiln
are illnstrati'd in figures H-7.

Ck'uus Scahriidl/i Swainson. I SKI
Subgenus Sicaiiisoiiia II. and -\. .\danis. 1S33
Scabricola (SuaiiisDiiia ) chipolaiia new sjieeies
(Figure 8)

Description: Shell Insilorm-elongate. moderatcK sol-
iil. Pi'otoconch eiddeil, Sninre distinct, impressed. T(>-
leconeh whorls 7. wcakK coincx. sculptured with 7-S
pitted spiral grooves on the earlv whorls. Spiral grooxes
ven' faint or ahsiiil on ihe pcnnltimate ami last whorl.
.\perture hall the leiiglh ol ihe spire, elongate, fairly
narrow and snioodi widiiii. Oiitei lip linn, outer edge
smooth. Colnmella with thin i-allus, callns more promi-
nent anteriorK. Colnmellar folds 4-5, moderately strong,
oblique, eijiialK spaced and ol Cqnal size. Siphonal fas-
ciole straight or slightK reciiiAed. si]ihonal notch prom-
inent.

Holotv

ype:

LI'" 93997. length 5(1,2 mm, widdi 1 1.

T^pc localil>: Chipola Formation. Tenmile Creek,

about 1,25 miles west of Chipola River (SK 1/4 Sec. 12,
TIN. Blow'). (Calhoun Conntv. Florida Tiilane Unixer-
sit\ localitx TL 951 ).

Distribution: Sinhricola cliiitolinui is a \i'r\ rail' spe-
cies thai has oiiK been collecled liom txxo localities
along remnile Creek and a single location along the Chi-
pola Bixcr. The txvo sites along Tenmile Creek are sep-

Page 48

THE NAUTILUS, Vol. 115, No. 2

Figures 1-8. Mitritliic Inmi tlic l.dwcr Miocene (Jlupola Fiiniuition, Florida, 1-3. Milm i.\/(/rr/i (uicti'Jiijpha Ciidiier. WVi .
llolohpe USNM 114;32S, Ient;tli U) mm, widtli 7.0 mm (Photograph of illustration in Cardner. UJ:,?: pi. XLX'lil. Kg. .5>. 2. Seleeted
specimen, UF 95999; length 41..3 mm, width 1.5.1 nun. 3. Selected specimen, I'F ^)(S()()(): lengtli 4.3.9 nnn. witlth 1.5.1 mm. 4-5.
Mitra (Fiisiinitra) callunincDsis new species. 4. Ilolohpe, UF 9598S. length 92.1 nnn. width 24.3 mm. 5. Parahpe A, UF .S7TiS4,
length 60.5 nnn (incomplete specimen), width IS. 3 mm. 6-7. Zilui iUaiiddIa (Woodring, 192S). 6. Molotxpe. USXM .3694.33.
lengtli 10 nnn, widdi 6.7 nnn (iiinstration (roni Woodring {192S) pi, 14, hg. 13). 7. USNM .3714.35. li()lot\pe o'i Mitra iititroilitii
(lardner, 1937, length 27 mm, widtli S.S mm (illustration from Clardner. 1937; pi. 4S. fig. 10). 8. Srahricold iSuauisonhi'' clnpcilinid
new species. Holot}pe, UF 95997, length 50.2 mm, width 14.7 mm.

aralcd l)\ ;ip|)r(i\iniatel\ a liall mile, anil locati'il (ni (ip-
posilc sides (il tlie eri'ek.

Et\'m<)lc)g\': Xaiiu'd lor llie ( 'liipdhi Hi\ci' in nDrtlicni
Florida.

Discussion: Scuhhcdld cliijidldiiii is oiiK known Ircnii
one complete and two liadK lirokeii spceinieiis. It is less
hulixms and lacks the pronoinieed spiral sculpture eliar-
acteristie of .\/(7;v; (icIi'H'^h/iilid (;;ii(lner (19.37). and is
MUicli smaller and more slendt'r than Milid idlli(iiiiic)isi\.
Tlie ilistineti\<' siiell sliape ol .S. rhiiHiliiiid rendered ge-
neric allocation diffienlt. ,\n initial rex iew of fossil and
extant New World mitrids h\ tins in\eslig;itor revealed
that S'. chlpolanii was not compatilile with aii\ known
.species. Howewr. a comparison of tins shell wiHi Indo-
Pacific species showed that its shell features wire most
similar to those in the gcmis Srabiicola. Memhers of (his
geinis reach a length of fit) mm, ha\-e o-S eon\c\ or
siil)angulate whorls, are scni|iliired with deep, pimelat-
ed, or striated groo\es, have ,ui aperture thai is e(|u;il in
length or longer than the spre. a collumel!;i with (i-S
obliijue lolds, and ha\e a distinct siiihonal iioleli Sca-
hricold rliijxildtid was assigned to the siil)'.^eiiiis Sniiiii-

.soK/r; because ol its sk'mler and smoolhei' appearance
and the punctated spiral scnlptuii' was conhned to the
earK spire whorls. .Among extant lu(k>-l'aeifie specii'S S.
scdhrirohi hears some rest'inhlauee to tlie sh;ilkiw -water,
sand-dw idling sulispccies Sctihricold {Sicainsoiiia) cd.sta
((imeliu. 1791). liowe\er, S. casia has a stouter shell
shape, fewer punctated spiral groo\es on the earK' spire
whorls (.) \s. 7', le\\er i-olumellar folds {4-5 \s. 5-fi),
and its cohimellar lolds are situated nioic toward tlu"
antei'icH- end. Scdhiirold rliipolaiui is the first and onl\
known example of this genus Ironi the New World. OiiK
<iiie fossil exampk' of this genus. Scdhiicdiii iScdhrirohi)
ilcsiidiii^sii iKieuer, 1S.3S), has been reported Irom the
l'li<icene of the Indo-Pacific (Cernohorsk\, 1991 '. and
Miere AW no known lossil represent;ili\ I's ol the siilige-
iins Sil(li}lsoitid.

\( K\()\\i,i:n(:\ii:NT.s

The .uilhin extends a special iiole ol lh;mks to Roger W.
I'oilell for ;illowiug examiiuition ol (liipdla Formation
speemielis under his e.iic licnn ImHi the ldiind;i -\luse-
iiiii ol \.iliii;il 1 listoiA and d niane IniNeisitx eolleelions.

C,M\ \y Sdiiiicl/. 2001

Paet' 49

as well as, proxidiinj; assistance witli all pliototiiapliic
work and re\ie\\in<:; earlier versions of this niannseiint.
Acklitional notes ol appreeiation are extended to Warren
C. Blow, Unitetl .States National Mnsenin. loi llic Idan
of specimens from the (iardner collecti<in. lo irirx 1 ,011
from the Unixersitv oi Florida lor photoe;ra|ilne work, lo
Dick Petit tor proxidinsj; copies of rare pnl)lieations. to
Gun Rosenlieri; for his helpfnl snggestions. and to ( Veil
,Sexton and Bnrt Haves lor iirantin^ the anlimr permis-
sion to collect on their properfv.

LITERATURE CITED

Abbott, R. T. 1974. American scasliclls 2n(l rd \.iii XosdaiMf

Reinhold, New York. (i(i.3 p., 24 pis.
Cernohorsky. W. O. HJTIS, The .Mitiidae of tin' Woild i'.iit I.

The subtairiiK Mitriiiar lii(li)4'a(.ifit \lolliisca l.> il7):

273-52S,
Cernohorsky W. ( ). IVWl. The .\litndae ol the Wodd. I'art 2

The .subfamilv .Mitrinae iDiicliuled and siibtainillcs Im-

bricariinae and CAlindrdiiiitruiac .MniKi^r.ipiis nt Marine

MoIlusca4: 1-164.
Ddcken.T. D. III. 1077. .Mollu.sca nfthe .MixkKs Branch Kca-

uiatiim, Mississippi. .Mi.>isissippi Ck'Dlegical, Kconoiiiic and

Topographical Snncv, Bulletin 120: 1-212. 28 pis.
Gardner I. A. 1937. The iiKilliisi.in launa nf the Alum Bliill

group of Klorida. Part fi: Pteropoda, Opistolnanehia, and
Oleiiohrauchia iin part). United States (ieological Snncv,
I'rolesMdU.il Paper I 12-F: 251-435, pIs. 37— 4S.

\l.ic\cil. P. S. aud I), r. Dockerv III. I9S4. Power Oligocene
( aistropiida, Sca|ihop<>da. aud (.'eplialopoda of the X'icks-
liiiiH (aiiup ui .Mississippi Mississippi Departuieut of
Xatiii.il lirsouKis Burr.iu ol (a'ologv Bulletin 124: 1-
41.5 pis I -72.

( )lssi)ii \ A, 1022, The Mioicui- ol uorllicru ( ^osl.i Kiel v\ith
uolcs iiu its general stratigraphic relations, i'art I. Bulle-
tins ol ,\ruericau Paleontoloirv 0 i.39': 1-167. pis. 1-15.

Pilslin II \ 1022, H.-visiouofW.M.CahhsTertiaiyMollusca
ol Sanlo Do il;o. Proceedings of the .Vcadeuiv of Xat-
iii.il S.iriKcs ,,! Philadelphia 73: 305^35, pis. 16-47.

<,'i I, I P, |r PJSO. Pleioptvginatidae. a new fuiiilv of uiitri-

lonii gastropods ( Prosohrauchia: Xeogaslropodai, The
Xaiitiliis 103: 1.3-10.

Solil X P 1064. Xeogastropod.i. ( )pistliolii,incliia ,iiid B.isoru-
luatophoni Iroiu ihr liiplev, 0\\\ Creek, aud Prairie Bluff
ioruiatious. I uitnl St.iti-s Geological Snni'V. Professional
Papia .531 B: |,5:5-.344. pis, 10 52,

NokcN. 17 II, lOSO. .\u ovei-vii-w ol the < liipohi I'orui.itiou,
uortliwestian Ploritia, Tuhme Studies iii (icolo^v aud Pa-
lcoiitnl(..„n 22: 1.5-24,

WiHiilnirj; \\ I' I02S, .Midrcuc iinilliisks Iroin Bimden, Ja-
maica: pi, 2, (iaslropods ,uid discussion of results. Publi-
cation 3S5. ( !arucgie Institute. Washington, 546 pp., 40
pis,. 3 figs.

THE NAUTILUS 115(2):5()-.54. 200!

Paee 50

Fisidiiim chkjuitanuni new species from Santa Cruz de la Sierra,
Bolivia (Bi\'a]\ia: Sphaeriidae)

Crisliiin F. Iluarte

Drpartiinieiito /.Doloiiui liiNcilchiados

Museo df La I'lata

1900 La Plata, Buenos Aires.

ARGENTLXA

cituartc@niusL'o.k-ii\ iiiiiiilpi'iliiar

ABSTRACT

A new species nf Pifikliiim C. Pfeiffer. 182L from Santa Crn/
de La Sierra, Boli\ia, is liere described, Pisidiuiii chUjintinuiiu
new species is defined In the striking o\ate slid! outline. l5ac!<-
ward displaced bealcs. enclosed ligament pit, and In the pres-
ence of two siphona! apertures and t\\o dcinihnmchs on eaeli
side,

Additunml kii/ (r('/(/,s, Soutli ,\nierica. freslwater, bnaKi',
clam.

INTRODUCTION

Little is kiiowni ahnut the species ol the ij;emis Pisitliiiiii
C. Pieifler, IS21. in Bolixia. At present, oiiK two species
ha\e been reported troiii tbat eonntn: Pisidhim incicr-
hrooki Kiiiper and Hinz, 1984, cle.serihed from a pond
at 4100 ni altitude at Cordillera de lo.s Frillies (19°34' S.
65°45' W, ea.stem Potosi, BolKia), also reported from
Nubi Lake (15° S. 69° \V) at 4fi00 ni altitude, near Ulla
Ulla, and Pisidiiiiu rile Filsbn, 1897. reeentK reporti'd
from Estaei(')n Parapetf, Sauta Cm/, de la Sierra (Itnarte.
1995).

The species originalK describi'd as Pi\idiuiii (Pisi-
diitm) tUiracciisc Pilsbn; 1924. and Pisidliim huliiiciisf
(Sturanv, 1900) actually belong to the genus Sjiliafriinii
Scopoli, 1777 (Kuiper and Hinz, 1984). .AddilinnalK, ( >/-
das forhesi (Phiiippi, 18fi9) was erroueonsK cited b\
Pilsbiy ( 191 1 ) as a Pisidiniii species. Pisidhim cliiiiuildii-
uni new species i'roin Santa Cruz de la Sierra, Boli\ia, is
described in the present work.

MATERIALS AND METHODS

The source of materia! is described in the svsteuiatics
section. The specimens were fi.\ed innnediatek alter col-
lection in 80° etlianol. Specimens lor scaiming election
microscopy were cleaned by repeated rinsing in distilled
water followed by a short treatment (2-.3 seconds* in a
10% sodium hvpochlorite solution. For periostracum re-
moval scNcral specimens were treated with a concen-

trated solution ol commercial sodium. Linear measure-
ments (shell length (SL), shell height (SH), shell witltli
(SW) and pre-siphonal suture (PSS)), morphometric in-
dices and ratios ((height index (HI = SH/SL), comexit\-
index (Ci = SW'/SH), ratio hinge length (HiL):sheil
length (IliL/SD), were calculated according to the cri-
teria followed b\' Ituarte (1996). For each calculation (n
= 21), mean and standard deviation values are given.
P\)r comparatixe puiposes, specimens of Pisidiitm )iia-
'^I'llaniniin (Dall, 1908) from the Princeton UniwrsitA'
Expedition to Patagonia lodged at the .\cadem\ ol Nat-
ural Sciences ol Philadelphia (ANSP) were useil.

SYSTEM.ATICS

Pisidiinu chiqiiilimum new species
Figures 1-14

Diagnosis: l^istinguishable In somewhat low and
markedh (nal shell shape, with low and liackAxard locat-
ed beaks, position of ligament, internal but extenialK'
visible, the presence ol branchial and anal openings, two
demibranclis on each side and nephridia ol closed tvpe.

Description: Shell: Tliiii, transluci'iit. of small to me-
dium size (mean SL = 3.9 ± 0.25, maximum obsened
size: 4.2 iimii. iu)t high (mean III = 80 ± 1 ). moderatelv
eoii\t'\ (mean (^i = 61 ± 4). sh(>ll outline iiiarkeclK oval,
elongate, anterior end produced in a shaip cunt', pos-
terior end short. \vid(4v rounded, sometimes slightK'
Inmcatcd and straight (figures 2. 3). Beaks knv, slightlv
projecti'd above dorsal margin, narrow, displaced back-
ward, located al about (i2'» ol SL. Shell surlace finely
and soinevvhal iircgiilarlv striated, dull glossv, straw -vel-
knvish.

Hinge plate solid, liiuge line rather long (IliL/SL =
56 ± 2). Hinge: Right valve f figures 4, 5. 8': cardinal
tooth (C;) strongly euncil m llie middle. (|iiite narnnv
ill .iiileiioi- half. gr( allv cnlai'.ii'd in a vvcaklv groovi'd.
rounded iii]'). Lateral Icclli robust, mner anterior lateral
(.\l) long, vvidelv curved, cusp little displaced lonvard;
outer .Ulterior lateral tooth i.MlI) i|iiite sliorl, ciisji (lis-

Cristian F. Itiiartc. 200

Paec 51

Dib, CRTremoullles

Figure 1. Location map. ♦: T\pf localih.

tal; liiiu'i- postcrioi- lateral (PI) nearK straiij;lit, rather
sliort. cusp sul)-central: outer posterior lateral (Fill) re-
duced in size with distal cusp. Lett \al\e (figures fi, 7):
cartliual teeth short, the inner (C\) triangular, lieiit up-
waid, slio;htIv oblique in relation to antero-posterior ;L\is.
the outer (C4) a narrow, uniforTulv cuned blade, (|uite
oblique, overlapping C, at posterior half; anterior lateral
tooth (AID strong, straight, cusp sub-central; posterioi-
later;il tooth (FID relatively short and weak, cusp distal.
Ligament pit enclosed, deep, inner margin straight to
slightK' sinuous, concave at posterior entl (figiu'e 8). Es-
cutcheon long, lanceolate, well marked b\ a delicate line
(figure 9). Ligament long, strong, internal, even though
\isible, from outside, in anterior halt through a\en nar-
row, and in some cases rather long, gap between \aKt's,
ne\er protnided (figures 9, 10). Ligament length is 23
± 1% of shell length.

Anatotnij: Anal siphon and branchial inhalant opening
present. Pre-siphonal suture rather long, lepresenting 1 1
± 2 % of SL (figure 13). The branchial inhal.inl opening
is determined bv a weak fusion of the imier mantle
lobes, without muscular .sphincter; a weak ventral re-

Irattoi' muscle is preseul. Anal siphon well-developed, a
p;iir ol powerful siplioual retractors present (figure 13).
Muscle scars (S scars), corresponding to the inner radial
iiumtle muscles, well marked. King apart from the pallial
line, those corresponding to anal siphon retractors coa-
iescent wilh |)osterioi ;i(ldiKlor muscle scars (figure 11).
limei' and oiilci' drmibranchs |iresent. Outer denii-
branch siiuillii-. lornied li\ 11-1.5 \er\ sliort descending
filaments, reaching back to the 14-16"' filannMit of inner
demibranch (figin'e 12). Brood ])ouches dexclop some-
what upwai'd and poslcriorK ol inner deiiiilnanchs
(howi'xei' in contact wilh. or slightK coveretl b\. the as-
cending lamella, which is well dexeloiied) (figiu'e 14).
Each brood pouch contains 3 — \ embrvos ((.'mbn'onic
shell length: 1.25-1.40 nnu). Xephridia of closed l\pe.
dorsal lobe \ariable in shape. coimnouK sub(]nachate
(figure 12i, lateral loop not \isible in dorsal \iew.

T>pe localih': Unnamed pontl at "La Siberia", a site
west of C.'omarapa, Manuel Caballero Frcnince, Santa
Cmz de la Sierra Department. Holixia (figure 1); M. C;.
Cuezzo coll.: 09/0S/199T.

Et^Tnologj': The name refers tt) (Jliic^uitos. afioriginal
inhabitants of the lands presenth' kiiown as "Llanos de
Chiquitos" (central-eastern Rolixian plains), near die
tvpe locality

Type material: Holot\pe and parat\pes in the mala-
iiilogical collection. Museo de Historia Natural "Xoel
Kempft Mercado", Santa Cniz de la Sierra, Bolivia
(MHXB 34734). Other p:uat\pes: Department of In-
\ertebrates. Museo ile La Plata (MLP 5362); Depart-
ment of Inxertebrates. Museo .\rgentino de Ciencias
Xaturales (.M.VCX 34734); Fundacion Instituto Miguel
Lillo, Tucuman. .Argentina iF"IML 14235 and 14236);
and Museum national dllistoiri- Xatnrelle. Paris
(MXTIX mmumbered).

DLSCUSSIOX

Pisidinin rhujinliiitinn new species can be easiK identi-
fied among South .'Vmerican Fisidium species b\' its strik-
ing oval shell outline with low and narrow beaks and
internal (howe\'er \isible Irom exterior' ligament. The
new species is also characterized b\ two. inner and outer
demibranehs on each side, two siphonal openings anil
nephridia of closed t\pe.

Pisuliinii cliiiiuitinnnii new species resembles I'isi-
(liiiiii iiu'iirhniiiki Kui])er ;uid llinz, 19S4 from Pern and
Holixia. which is the onK known species Irom tropicid
Soutli .Americ;i with both, branchial and anal siphonal
openings (Itnarte. 1995'. /' lueierbrooki differs from R
chkjuitaninii in li.ixing a more convex shell (Ci =
77 — 80). fullci' and more backward displaced beaks.

Pisiduim chicjuUdnuin new species is mcwt similar to
those specimens from Ecuador and Peru figured and
reported In Kui])cr iind Ilinz (1984) as PisidUnn cnscr-
liniuni 'Poli. 1791'. :i F.urasian species, extremelv vari-
able in shell shajie and currentK regarded ;ls cosmopol-

Page 52

THE NAUTILUS, \(il. 115. No. 2

Cristuiii F. Ituaiie, 2001

Paw 53

Figures 2-1 I. I'i^iiliiiin ,lii,iiiil,iiiiiiii new spccicv 2. Ilololvpr .\III\H> :{-Il. I'arahpis \ll,r5:>(S2. 3. I'lisli-nor xicw ol a
specimen. 4. lliiii;e ul a rii;lil \aKe. 5. linjit \aKe. detail ol eaidiiial lodtli li'j;aiiieiil and anUTicir lateral teeth. (>. llniiie ola left
valve. 7. Left valve, detail of ligament, cardinal teeth, and pcislcnor Literal tixith S. Detail olcanhnal tooth and ligament ola ri>Jit
vaKe. 9, 10. Dorsal \iews of two spicimens showiiit; \aiialiility in shape and leii<;th ol tin' i^aj) hetween \aKes. 11. Inner \ie\\ ol'
a ri<.;hl \,il\r margin, showing seais ..I mner radial mantle nmscles and anal si])iion retractor larrowhead). .Scale hars: Kigures 1-7.
11 = 1 imu; Figures = ()..5 mm; l'"igures !). 11) = 2(K)|j.in.

itaii (Biireli. l97o: Kiiiper. U)(i(i, U)S,x Kiiiper ami Ilin/,,
19S4; Holopainen and Kiiiper, 19S2). H(we\er, these
.specimen.s are larger than /' cliiiiiiitiiinini new species
(mean SL from measinenieiits gi\eii In the aiitluns was:
4.51 ± 1.31 and maximum shell length: 7 mm', haxiiig
more ci'iitral beaks ami less produced aiiteiinr end. I''iii-
theriiKiie, /' rJiUjUitiimiiii new species diHeis Irdiii /'
Cdsciidiium in heing less eon\c\, with Iowit and iiaiKiw-
er beaks, decidedk' loeatetl baekAxard. Moreo\er. /' clii-
ijuitaiuun has a comparativeK long pre-siphonal sntnre,
being about 1 l-lo'/f of tlie shell length, whili' /' cdscr-
Idiiiiin has a shorter one eomprising ea. lOOi ol shell
length (Holopainen and Kiiiper. I9S2: Korninshin.
1996).

Pisiiliiiiii (■hujiiitdiiinii new species shares with V'lsi-
(liiini dKhu-llddiiinn (Dall. 190S) tile sanu- si|ih(iiial ai-
laugemeiit. the t\"pe (il nephndia and tlii' imniliei' ul
tlemibranchs. Howexer, /'. iiui^clldiiicuni. the oiiK /'(\-
Uliiim species from Argentinean and (diilean Patagonia
known to ha\e two siph(nial openings aiul two deini-
branehs. dilters trom P. chhiiiitdniun b\ the more cen-
tral position ot beaks (mean = 5(i'7f, range 53-5S'7r ol
SL), shorter presiphonal suture, representing about
7.7% ot shell length, and shorter ligament (mean =
19%, range: I<S-20% of .SL) (mi-asurements and indices

Figures 12-14. Pisidiuiii rhiijiiiliiidiin new species. 12.
Gross anatomv 13. Mantle muscles. 14. Inner \iew ol inner
deniihranch. (bp: brood pouch; as; anal siphon; ia; inhal.inl
aperture; id: inner deniihranch; inn: inner nidial mantle nms-
cles; n: neplnidinni; od: outer demihrancli; pss; presiplioiuil
suture; sr: retractor muscles of siplioiial openings). Scale bar:
1 mm.

aie those icpdited li\ Ituaite i 199(ii ;ind e(ni-es|)ond to
speciiiiens ol /' Nid'U'lldniiiini from Win ( :hieo, Saiit.i
(ill/ I'niviiiic. \igeiiliii,i studit'd b\ Pilsbr\ ll91F.
winch ,ire p;irt of lot SSSl 1 .WSI^).

Pisiilinni rhUjiitldnnin new species cliiiiU differs
Ircnii the giiiiip III spi'cies distributed .iloiig the drain-
ages ol siiulliweslerii Hra/il. Fiugii,i\. ;iiid nortlieaslern
.Argentina (Itiiaiie. 2l)()()) in liaxing (wii demibranehs
and two siphonal openings, .\niong those. Pisitliiiiii
sIcrkiduniH I'ilsbiA. 1.S97. a species wideK distributed
in central and iKirtheiii South \meriea. also differs
linm /' iliiijdilddiDn new species b\ an e\ideiitl\ ex-
ternal and pnitrnded liganieiit. ha\ing a more inlfated
shell and nepliridi;i ol open t\ pe (Korninshin, 199S):
Pisiilnun (/(iihi^in/i (.'lessin, 1S79. hum Friigii;i\ and
Pisiiliinn vile PilsbiT,, 1S97. deseiibed from Frugnax
and also (list li I II I ted in l^o|]\ ia and \igenliiKi. ;ire much
smaller specie's: Pisiilium pipocii^c Ituarte, 20011, is
larger than /' ihhjuilinuiDi new sjiecies. having a
marked snbi|uadrangul.ii shell dutline. while Pi^idiiun
Idrd^iii/ciisc Ituaite. 2000. chlleis in lia\ ing a larger and
ijiiite globose shell, with tiiller beaks. /' cliUjuUdiunn
new species dillers from Pisidiuiii forriisc Meier-Brook,
1967. from Mm. is (ierais, Brazil. b\ its smaller si/e,
rounded shell milliue. lowci' and less eoiucx shell, and
b\ liaxing A iiKii'e iiiterual ligaineiit.

ACKXOW Ff.nCMIATS

The aiithoi' wishes to ;icknowleclge l)i'. M. C.. (aie/./o,
(Institnto .Miguel Lillo, Tueuman, .Argentina), who col-
lected and kiiiclK sent specimens for this stuck. Her fielcl
liip was partK sup[)orted b\" tlie .Museo de Historia Nat-
ural ".Noel Kempfl Mereado", Santa ("rii/ de La Sierra,
Bolivia. Di". I\ui|iei kiiidlv assisted with the bibliographv
and Dr. A. \. korniiishin (fisenssed and commented on
tlie relevance of soft part auatonn features as diagnostic
characters. The .lullmr is Researcher ol tlie Coirsejo N'a-
cioual de Inxestig.iciones (aentfficas \ Tc''C'nicas ((X.).\-
ICET). .Argentina.

LITER.'XTFHF CITLD

Biiich. |. 197.'i I'icsliwiilei splKieriaccan cfniis ' Molliisca: I'e-
lec\pod;i' ol Xuitli .Viiieriea. 2'"' lulition. Amerie;m M;il-
;icologic.il liililic.itions. Hamhurg. 9fi pp.

Holopainen, I |, .uhI |. (i. |. Knipcr. 19S2. \otes on the iiior-
pliometiA and aiiatoiiiv ol some Fisiilimii and Siiliairiiiiii
species (Bivakia, S]ilKieriicl;ie L .\iiiiales /oologici l'"eniiici
19; 93-107.

Iln;irte C. F. 1995. Nuevos registros de I'i.sUliiim Bleifler. IS21

Page 54

TUE NAUTILUS. Vol. 115. \(

V Sphaehum Scopoli, 1777 iHi\al\ia: Spliaeiiidae) en

Chile, Bolivia V Norocstc an^eiitiiio. Xcotropica 41 (105-

106): 31-11.
Ituarte C. F. 1996. .Vri^eiitiiii' species of I'isuliuin I'li-ifier. 1S21.

aiid Mu.sailiiiin Link. 1S()7 ( Bi\aKia: Sjihaeriidaei. Tlie

\'eliger 39: 189-203.
Ituarte, C. F. 2000. Pisklium laraj^ni/cn.w and Piskliiiiti pi-

poertse, new species from northeastern .Argentina ( Bi\ al-

via: Sphaeriidae). The N'eliiier 43: 51-57.
Koniiushin. A. V 1996. .Moiphonietrieal and anatomical chai-

acteristics of Pi.'iiiliiiiii aiscii/iiiutii (Poll) from the Lake

Biwa (Japan) (BivaKia: Enlaniellihrancliiata: Pisidiidae).

Malakologische .\bhandlungen IS: 53-57.

Korniushin, .\. \'. 199S. .A comparative iuxcstigation oi nephrid-
ia in hngeniail and pill clams. .Malaeological Hexiew.
Suppl. 7: 5.3-63.

Kuiper. j. C;. |. 1966. Critical revision ol the New Zealand
sphai riid elanis in the Dominion .Museum. Records of the
Dominion Museum 5: 147-162.

Kuiper, J. C,. J. 19S3. The Sphaeriidae of .\ustralia. Basteria
47: .3-52.

Kuiper |. C. |. and \\. lliiiz. 1984. Zur Fauna der Kleinmus-
clieln in dvn .'Xndeu (BivaKia: Spliaeriidaei. .\rclii\ liir
Molhiskenkunde 114 [1983]: 137-156.

PilshiA 11. A. 1911. Non-marine MoUusca of Patagonia. Re-
ports of the Princeton Universitv Expedition to Patagonia
(1896-1899) 3: 51:3-633.

THE NAUTIIAIS 1 15(2):55-61. 2001

Paw 55

Favoriiuis clenalexiae, a new species (Opistliohraiicliia:
Aeolidiidae) from the eastern Pacific Ocean

Francisco J. Garcia

Depurtaint'iito tie Fisi()l()ii;ia \ Bioloiji'a

Animal, Facultad dc Biologia
Awnida Rcina Mcn-ctk-s, 6
41(IS() Sc\illa

sr\i\

l|t;arcia(2'cica.L's

Jcsiis S. TroiK'oso

Arc.i (Ic ISii)li)i;ia Animal: Faiiillad

( ai-iieias dt'l Ma?
Uiiiversidad di' \ it;n
La^das-Marcosciulc. \ mi)
Sl>\l\
tri)iici),so(i''ii\i''i).r.s

ABSTRACT

A new speeies ot Xndibranehia. belonging tii the genus lui-
vnriniis Cra\, 1S50. is deseribed. It ditlers Ironi ntlier speelis
ol that genus inaiiiK b\ its rhinophoial uniaiiieutatinii: l-'iiiii-
linu.s cleiHilcxiaf new speeies has tliree large. lAeilappiii'j;
flanges v\ith the upper margins free, like three eups. in alninsl
all the speeies the rliiiiophures are papillated or laniellatrd.
The external anatoiii\, radiila. and reprodnctise system are Ar-
seribed and eompared with those of other speeies ol the genus
\ table eompariiig summarizing the ditterenees amongst the
speeies ot the genus Fincuiiiiis is ineluded.

Atlditiiiiiiil krij ncnls: Xndibranehia. taxononn.

IXTHODUCTION

The genus Fiiiiniiius includes fourteen speeies charae-
teiized b\ the presence- of a cleioproctic position of the
anus. The I'crata are arranged in areheil clusters, lacking
enidosac, the niasticatoA edge of the jaws have sex era!
row'S of denticles and the radular teeth aie inodideil to
feed on the eggs of other gastropods. The teeth Iuim-
\eiv elongate cusp and conipleteK' lack or have \en re-
duced denticles. In thi.s paper, a new species ol this ge-
nus from tile eastern Pacific is described and a eonipar-
ison with other Favoriitus species is presentetl.

SYSTEM. \TI(:S

Genus Faioriini^ (;ra\. 1S5()
Favoriniis clciialcixa new species
Figures I -7. Table 1

Description: Flxtcnuil (indtoiiuj: The IukU Is elon-
gate (figure 1). The largest specimen examined was 15
mm long in chstended state. The bocK has up to 7
eeratal groups ini each side. The three anterior groups
are horseshoe-shaped, while the rest are oblicpu' rows
slightlv ciir\'ed at their distal end (figure 2b). The
number of cerata in the clusters was Hi. 12. 11. 5. 4.
2, and 2 in a specimen 14.5 mm length The eerata
have a smooth surface. The genital papi

fi. 12.

The eerata
I lies on the

right side at the lex el ol the first eeratal group. The
amis lies on the same side at tlie lexel ol the seconil
eeratal group. The liiiuophores are bhint-tiiiiii'd and
liaxe three large oxerlappiiig llaiiges with the iijiper
margins frt'c in the niaiiner ol three tups ifigure 2a).
(.)n the aiiteiiiii" ami posterior surface of tiie rhinop-
liores there is .i thin crest jiimiug the Hanges. The" oral
tentacles are long and cxliiidrical antl the propoilial
tentacles are long and (lossess a xential. longitudinal
grooxf. The loot is relatixelx broad and the tail is long
antl piiiutctl.

The backgroiiml color is translucent white. On the
lieatl antl back there are fine opaipie white dots, which
can come together as spots of different sizes, except
at the base of the rhinophores and base of the oral
tentacles. The oral tentacles are translucent white at
the base ami opaipu- xxhite along their upper middle.
The rliinoiihiiics are dark broxxii except for their base,
xxhicli is tiansliieeiit xvliite and their a[)exes. which are
opaipie white. The eerata liaxe xxhite spt)ts tlistribiitetl
on their surface; at the lex'cl of the cnidt)saes the sptits
join as an irregiilai' ring around the cerata. Internallv
to the cerata. the exteiisioiis ol the tligesti\e gland are
pinkish (figure 2c). All the digesti\e branches of a
cluster of cerata are joinetl at the liase of the cluster.
The tligestixf blanches are pink in color. The loot is
traiisluceut xxhite.

liitiiiiiil (iiKilKiiiii: Txxo specimens xxcre ilissectetl,
with 14.5 and 15 mm length. Salixarx glanils xx'ere nt)t
found. The radnlae liaxe H) anil 20 teeth, respectix'cly.
.Ml the teeth consist ol an archetl base with an elon-
gate cusp. Both sitles oi the cusp are smooth and the
apex is sh.irplx pointed (figures 3a; 4). There art" no
basal denticles. The twt) jaxvs are large and amber in
cohn-. The masticatory edge is long and eunetl ttnvard
the conxt-x lace of the jaxx' at the outt>r mitldle (figure
.■5b). .\lt)ng the eilge are four or fixt' roxxs of small et)n-
ical tlenticles and a marginal roxx of long conical ilen-
ticles (figiu'es 3c. 5'.
The reproductixe sxstein is ilhistr;iled in figure fi. Tlie

Pase 56

THE NAUTILUS. Vol. 115, NO. 2

Fimiro I. I'lnithnusclciiiilrxiac. F,\lcni,il \ii'\v.

goiiail empties into a liennapliniditie diut. wliieli en-
larges in anterior tlirection as an ampnlla. Tins is lon^j;
and coiled, l)itnreatin<4 at its anterior end into the def-
erent dnct and the inner o\idnet. The deferent dnct
is short; it I'nlarges into a short prostate, which leatls
to the penial papilla. |nst anterior to the penial papilla,
the jirostate gix-es \\a\ to a straight post-prostatic dnct
that connects to the penial papilla. The jienis is con-
ical, nnarnied, and pointed apiealK.

The inner o\idnct connects to a small and spherical
seminal receptacle. Tlie inner oxidiict coiniects to the
onter oviduct, which penetrates the li'male gland, '{"his
latter is large and envelopes the seminal receptaile.
From the female gland, the vaginal dnct proceeds to
the genit;il orifice.

Type material: Ilolot\pe: .VINCN 15.05/3205 1 12
mm length, leg. Francisco J. (Jarci'a, 12 Feb. 1997; I'ar-
at\pe: MNCN I5.0.5/.322.30. 2 parat\pes, 7 and 5 mm
length. Isla (Joiha, Pacific coa.st of Pananui. intertidal
zone. le<T. Francisco J. Ciarcia. 8 l'"eh. 1997.

Type locaiitv: Isla Coihita, Pacific coast of Panama, 3
m depth.

Other material exaniinecl: Chievitas. North of Pnnta
la Ciringa. Bahi'a de los Angeles. Baja California. Mex-
ico, 6 in depth, 1 specimen fi mm total length, leg.
Il;nis Bertseh, 1 No\. 1993; Sonth side Isla ( ;erral\().

Figure 2. F<nniiiiii\ cluuiUxiac. a. Rhinopliore. a. Diagram-
matic right profile sluminii the insertion of the cerata and po-
sition of the anus and lepiixfutive apertnrcs. c. ('crata cluster

Baja (California .Sur, .Mexico. 2 spt'cimens. with egg
masses. 5 m depth, 7 jnne 198.5 (one of these used for
SEM); Punta de Mita, Naxarit. Mexico. I specimen. 9
mm length, fi ni depth, leg. Hans Bertseh. 17 JuK
1991; Gnanacaste, west Costa Rica, 4 specimens, 6 m
depth. Leg. .\ntonio J. Ferreira, 13 Feb. 1972: CASIZ
88218. Islas l.adrones. Pacific Coast of Panama, 1
specimen, 12.8 m dejith. leg. Mart\ Beals. 1.3 .\pr
1993; CASIZ S8217. Isla Montuosa.' Pacific Coast of
P;m;iin;i. 12.2 m depth. 1 specimen, leg. Terrence M.
(iosliner, 15 Apr 1993; CASIZ 881.35. fsla Jicarita. Pa-
cific Co;ist of Panama. 8.8.3 m depth. 1 spc^cimen. leg.
Terrence M Cosliner. 17 Apr. 1993; CASIZ 88216.
Islas Contici;is. Pacific Coast of Panama, fi m depth,
19 Apr. I99.>. 3 specimens, leg. Terrence .M. Gosliner;
CASIZ S8222, Islas Secas. Pacific Coast of Panama.
19.2 m depth. 5 specimens. Ie<i. Terrence M. (iosliner.
21 \pi i99.v (:\SIZ 975;51. Isla i);uwin. (Galapagos
Isl.mds. l-An,idor. IS. 28 m depth, 1 spi'cimen. leg.
Terrence M. (iosliner. 13 Ma\ 1994; CASIZ 97510.
Isla Isahela. (Galapagos Islands. Fcnador, 31.4 m
depth. 1 specimen, leg. Terrence M. (iosliner. 14 .Ma\
1994; CASIZ 97522. Isla Rabid;i. (;ahipagos Ishmds.
Ecuador. 1 specimen. 27.12 m dejith. leg. Terrence .\1.
Cosliner. Ifi Mav 1994; Isla Coiba. l';icific coast of
l'an;ini;i, intertidal /one. 4 s|)eciniens 1.5. 14.5. 5.5.

Fraiifisfo |. (Garcia and [e'si'is S. Trniicoso. 2001

aiif o /

Figure 3. Finoriiiiis cUnalcxiuc a. Ividiilar tt-clli. 1). |.i\\ c.
detail oi tilt' ina.sticatDn edse.

and 3.5. mm iii k'ngtli, leg. F'l'aneisco |. (iaii-fa, S VAi.
1997.

Di.strihiition (Figure 7): Fiom Balii'a dc Fos .Xngclcs,
Baja Calilornia. MiAied. lo the Pacific coast ol l^mama.

EUillolog^k': Tile name nl tlii.s species, ficiiillcxiac. is a
juxtaposition of the names ot tile xoungest daughters ol
F. J. Garei'a and |, S. Troncoso. respectixeK. Flena and
Alexia.

DISCUSSION

The most similar species ot Faioriiuis cicntilcixa is R
tsuntaaniis Baha and Abe, 1964. This species, de-
scribed from )apan, was referred to originally as ha\-
ing rhinophores with two bulbs (Baba and Abe, 19641.
Later on, howe\er, tliese authors described (he liiin-
ophores as ha\'ing three cup-shaped flanges i Baha and
Abe. 1975), a feature also mentioned in the descrip-
tion of specimens tVom Eastern .\ustralia In \\ illan
(1983). Willan [op. cit.) stated that this character of
the rhinophores was known onl\- in F. tsuru^diius. giv-

en th.il nther Faroriiiiis species then known lia\e tile
rlnrH)|)lioies with bulbs along their siialt or are lamel-
late, iheielore, /'. clciialcxiac constitutes the second
specii's ol the genus bt'aring tliat character.

Faicriinis ilciinlcxiiic dillers Ironi F. Isiiniiitniiis.
Iiowexer. m.unK li\ ils eiiliirat inn. -\lthough both sjie-
eies Ikivc a t laiislueent while groinul color and the
ihinopliores are d.u'k. /' Isiinii^iiniis has a golden \cl-
low or lii'ow nish-xellow head, while that of E clciuil-
cxidc is opatpie white. The cerata ha\e a deep black
cnidosac aica and ihe di<4esti\e glands are orange or
orangt.'-\ell(i\\ ni /' /•^iinc^/imis. In /' ilciialixi/ic. the
cerata lia\e snperlieial white spots, and a wliite ring
around the li.ise ol the cnidosac; and the digestive
gland is pinkish, fhe number oi cerata per group is
slightK higlu-r in the anterior clusters in our species,
while the sjiecinu'us described b\ Willan i I9S3) ha\e
more cerata in the posterior elustei's. In our specimens
the three anterior clusteis ai'e arched as a horsesiioe.
while the n^st appear as ,i more or less cuncd row. In
F t•<llnl'J,lllln•^ almiist .ill .ii"e horseshoe-shaped (Baba
and .\be, 19751.

InternalK', Willan I 19S3i described the presence oi
lobe-like sali\ar\ glands with a broad duct and a clus-
ter of spherical acini. We did not see an\ sign oi th(>se
glands in o\u' specimens. The unisticaton' edg(> oi the
jaws has the outer middk' cuned toward the cou\ex
surface oi the jaw in /•' clciialcxiac. while in F. fsiini-
i^aiiiis it is straight. The radnlar teeth are similar in
both species. The penis ol /•' Isiiniiianiis is u-shap(>d
with the two limbs closeK pi'essed (Willan. 19S3i. F.
chiialcxiac has a conical penis and it is straigiit. The
internal organs ol the reprodnctixe sxstem ot F. tsu-
ni'^anii\ ha\e not been described.

F cIciKili'xiiic has a similar color pattern to F inir-
(ihilis Baba. 1955. In both species the rhinophores are
dark brown, with the apex whitish, the surface oi the
notnui and cerata are ])ro\ided In small white spots
and the cerata ha\i' a geneial pinkish color. HowexiT,
other external features distinguish both species. F.
iiiirahilis has the rhinophores pertoliated, the cerata
set in 12 groups on eitlur side, the general ground-
color is whitish inclined to brown on the back and
dorsal surface of the In-, id. llie oral tentacles and an-
terior edge of the toot are \cllow. and eacli cera is
marked with a purple spot down the ti]). F. clenalrxiac
has the rhinophores with 3 cup-shaped llang(>s, the
l)od\ onK" has up to 7 groups ol cerata on eitiier side,
the general grouTid color is whitish with opaque wliite
dots more or less concentrated on the surface of the
bod\-, the oral tentacles and foot are wliite, and the
cerata lack a purjile spot and haxc white spots round-
ing the base ol the cnidosac.

In a eom[iarison between /-'. clcnalcx'uic and the rest
of the Favoriiiiis species, the clearest feature permit-
ting a differentiation is the ornamentation ot the rhin-
ophores. Features of tlii' anatonn and coloration of
Faioriiiiis species are listed and compared in Taiile 1,
making it possible to differentiate F. clciialcxiac from

Pasie 58

THE NAUTILUS. Vol.

15. \()

lanlo I. (Iianu'i

tcrs <)t spt'cics

()t / innnniis

Body

Rhinophores

Cerata

Species

Relcrcnces

Distriliutioii

Hlimuplinrt's

Crrata

coloration

coloration

c(tioration

Kadiila

F. I'U'uaU'xUif

Present article

(Galapagos Is-

3 cup-siiaped

Sllionlii

'IV.uisliu.-i.l

Black brown

\\'hite spots on

SiMOolh

land. EiLstcrti

flanges

wintf uith

witli the a[)e\

the surface

l\Kific coast

opaquL- uintc-

white

and euitlc)sac-

of Mexico.
Costa Hica
and Panama

spots

Digesti\e
gland pink

F. hranchialUi (Rath-
ke in .Vliiller.

Ednmnds &
Marcus

Eastern Atlantit
and M.-ditn-

l-2Imlhs

Sllinolll

Translucent
white with a

Brown, tip white

Upper part white
oi>a(]ue. Di-

Smooth

1S()51

(1977). Hn.-
vnt-Fol
(1954). Catta-
ne()-\'ietti et
al (1990)

laiiraii Sea

\'ariahlc pat-
trni of
opa(jne wliite
blotches on
the back

ges-ti\ f gland
\xiriable.
brown, \ello\i-.
red. green

F. jouini (Risbec.

Risbec (1953)

New CiiKdoma

Lauiallate

NA

Translucent

Brown with the

\A

S denticles

1928)

white

ape.v red

F ^ouaroi (Risbec.

192.S)

Risbec (19.531

New Caledonia

2 nnt^s

NA

Vellc.w with
lar^e opa(|Ue
white spots

Opaque white

Digestive gland
yellowish

ft denticles

F. iit>lnci'u~s (Ri.shec,

Risbec (19.53)

New ('alfduni.i

Lamrllate

NA

Head xellow

Pnikish widi the

White \u.let

7 denticles

1928)

lirecmsli,
bodv white
with \'io!et
lilints

ape\ \ioiet

E pacifiats Baba.

Baba (1937)

Japan

Indistinct con-

S,nn,.|l,

Translneriit

Chocolate-tinted

Dark vellow.

Smooth

1937

strictions
;Uong the
whole lent^li

white

except at the
distal third
which is col-
orless

with a white
cap and a
pmple mark-
ing below

/'" Japan icus Baba.
1949

Baba (1949).
Baba & Ha-
inatani

(1964), Gosli-
ner (1980)

lapaii, Hawaiian

Islands

2 or 3 hulhs

Papillalrd

Yellowish with
dense opat|ni'
wliite except
the cerata

Translucent
white

Digestive gland
red, \ellow or
yellowish
brown. wiUi
opaque white
apices

Smooth

F. perfoliatus Baba.

Bal)a (1949)

Japan

Perfbliated

Snioolh

Pale rose

Brown

Digesti\e gland

5 dentic-les

1949

\eined with
rose- red

F mirabilis Baba.

Baba (1955).

Japan

Perioliatt^d

Smr.i.tli

Wiiitish. inclined

Dark brown

Brownish with a

Smooth

19.5.T

VVillan (1983)

lo litowTi on

ihe hack

puiple spot
down the tip

F auritulu~\ Marcus,

Marcus (19.55).

Trnpua! West

2..I 3 l.nll.s

Sm(...th

Trausjucent

Browii, tip white

W hitc epitlennai

Smootli or

1955

Marcus fie
Marcus
(1963). Ed-
munds
(1964), Ed-
niunds ix
Marcus
(1977).
Thompson
(1980)

Atlantic

white or gra\.
with white
spots; the
area behinil
the rhiuo-
phores had
an orange
ci'Uter

pigment; di-
gesti\e gland
;i]most c«k>r-
Icss or pink.
gra\ish or
dark browii;
cuidosac white

tinv later-
al denti-
cles

F- pannucfu-, Biini.
19fi2

Bum. 1962, Kd-
nuiiuls. l9(iS

Austraha

Papillate

Smotith

White

White

Kawn with white
tips

NA

F. ts-untntinu.-i Baba
& .•Mw. 1964

Baba & .'Vbe

(19fM. 19751.
Uilluii 1983

Japan, eiustem
Australia

2 l.nll.s n, 3
(.viilappinti
iLiniics with
tlirir nppi^r
iiiariiins free

S.n..ntl.

Tiansiiuenl
head golden
\ellow or
hrownish yel-
low; hod\'
deuseK' cov-
ered witli
opaque white
spots; anteri-
or margin of
loot \ellowish

Black

White dotN

down the tip.
Cuidosac
black. Diges-
tive gland or-
ange

Smooth

F ^hatu-tisis Ed-
munds. 1968

Ednnuuls
11968), E<1-
tiMirids iSc
Marcus
(1977). Ortea
(1982)

West Africa
l(;lianai

3 small swellings

Smooth

Translueenl
gra\isli-while.
A few white
dots on head,
oral tentacles,
back and tiiil

Tip clear gra\i-sh
with wliite
dots, the rest
pniple-hrown
or maroon

A few white dots
on surface,
and a few
white glands
at tip. Digcs-
ti\c gland
cream or
brown wth

|)iiiple-bliiw-U
hlol.hes

7-S tlcnti-
cles

FraiK'ist'i) |. (larcia and [esus S. Troncoso, 2001

Page 59

Table 1. (j)ntiriiu'il

Specio

R.-rrn-iKr

nislniiiilHi

Hlin,..n],<

HckK

iloiMtinii

Rliinopliorfs
toloration

Ci-rata
^■olorutini

Radula

F hihinus I.niKli.' \ L.-tiKlH- \-
Tlioiiipsnii 1^)74 Tlioiupson

..19741, Kd
iiinruK (S:
Marcus
(1977)

/■ M/fVi/s Ortra I9S2 Olira ^UJSili

\urtK.

Caiian Klaml^

LMlaiitu
Ocean I

- Ilii^-sli.iprc
swcllmi's

: small Iniil.-

Sninotli

K.iint stra\v-\cl-
Inwisli and
sciin li\alnic,
Sliintnji wiiilc
pijiTiicnt on
tlic dorsiil
sulcnl ilic
oral tc-ntaclcs,
conlinuinti
hackwards
Ix-twccn tlic
ilunnplioK-s

Tiansliiccnt
uiiitc with
npaiplr white

niitrkiri'^ on
head, dorsum

and oral Irn-
tacK's

VcIIowisli-hrown Faint vclltmish- Smootli
l>rt)\\ii witli a
niirrow al-
most black
strip on tlie
posterior side

White

W'iiite opa(jne 4—5 deiiti-

clfs

XA = Iiiliiniiatioii iKit a\;Lilalilc

the otlif 1" species b\ the presence of sinootli radiilar
teeth (F. joiibini. F. i^ouaroi. F. violaccus. F. jiciinlid-
fus. F :j_luiinii\i\. and F. iitrnis lia\c dcntiinlatc
teeth). The radnht ut F. iKiuiuiccu.'i Burn, 19(i2, lias not
been descrihetl. However, tliis species dillcrs Ironi F
flciiiih'xidc 1)\ tlie rhinopliort's and coloration. 'Hie
cerata are smooth in F clenalcxidc (in F iiip<>iilcii\
tlie\- are papillated) anil tlie iinier o\iduct does not
have a swelling for a fertilization chamlier as was de-
scribed in F juponicus Baba, 1949 and F gluinctisis

Ednmnds. 19fi8 (Baba ami Ilaniatani. 1964: Ethnunds.
196S). F iijidiivHsis has a small penial st\let.

ACKNOWLEDCMENTS

We wish to express our ifi-atitude to Terreiice M. Gos-
liner and Hans Bcrtsch lor snppKing ilata on th(< dis-
tribution of till' specii's, SEM photographs, and for
their comments about the manuscript. Michael Miller
for assistance in digital imaging and transmissions of

Figure 4. i-aionnus clciialcxmc I.clt. SEM iiiirro^iapli ol a lailulai tootli. Hit;iit. SEM iiiicrograpli ola l.itrra! view of the radiila.

Pase 60

THE NAUTILUS. X'ol. 115. No. 2

I'l'mire 5. I'dioiiiiiis cli-ndlcxidc. SEM iiikTo^rapli iif the
iiia.stiL'iiton cilijc <>l till' |u\\.

tlic SKMs, and Richard (l. Willaii for snppl\in5 us
with some hihlioij;raph\' and lor his foniini'iits rctiard-
ing thi.s |)a])ci-.

We also tliank .Anemia I'.spafiola Ar (^oopcracicni
International and Institnio Xaeional de Heciir.sos Na-

sr-^-

PP

Figure 6. FaiDriiiiis clciiiilcxiar. Heproductixe sv.strin Ah
hrc-xlations: a, aiiipniia; dii. (lol'ertMlt duct; fg, fciiialc gland: lid
iicrniaphroditic duel; io. inner o\iduct; oo, outer o\i(liirl: |)p.
peuial papilla; sr seminal reeeplacle; \d. \agiiial duel.

Figure 7. nistrihiitidii ol F<U(iriitiis clinnlixiiH'.

tiirales iienoxahles Ircnn Panama for tlieir a.s.si.stance
during tlie e\pt'(htion in tlie National Park ol Coiba
(Panama) ;ind Io f'ngineer Lnis Carlos |imenez Cer-
rud and Narciso Bastida (.\IaH Mali) for their constant
help. This research was included in the project In\en-
l;u"io de I;i l'';iiiiui \ flora del Parque Nacional de Coi-
li;i. fiii;iiu-ed In ,\<j;enei;i Espanola de Cooperaci(')n lu-
leiiKieiiinal ,nid Ikis lieen partialh' supported !)\' the
project CICVT PB9S-I 121.

IJTl'iiVn HK CITED

li.tlia. K, lil'iT. ( )pisdii)lii,iiu lii.i III |apaii 11 . |iiiirii;il iil llic

L^epartnieiil nl \'4i iciilliiic. KmisIhi linpiii.il I nncrsitx

5(7): 289-.1I I, '

H;ili;i. K. 1919. ( )pistlii)l)r;iriclii;i ol .Siiganii H;i\ follected In liis

\lajcst\ the KuipiTor ol |;ipaii. Iwanauii Siioten. Tok\().
B:il)a K 19.55. Opisthohrauehia ol .Sagaiui Hav .Siipplrnuiit

ciillccled h\' his Majcsl\ llic I'jnperor ol |;ip;iii. I\\aii;iiiii

Shoten, Tok\ci.
H;il)a. K .111(1']" \lir l!)(il til -i I lids III I'/niiriiiii.-.lsiinii'iniiis

"raiK'isco |. CJarci'a anil It'si'is S. Troiicoso. 2001

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new species, from Tsunisja Ba\. |apaii (\u<lilnaiicliia-l''(il-
idoidea). Publications ot the Seto Marine Bioloijical l.ali-
oraton- 12: 47-4S.

Baba, K. and T. Abe. 1975. ( Jirniiients ol hnllier spctimiiis
ol i'lnoriinis tsiinii^tiniis B. ix: A. Iroiii Kilii/.en-( 'Im nc.ii
Tsurui;a Ba\. Japan (Nndibrancliia; lM)lidiiiilea: l'",i\onni-
dae). Publications ot the Seto \lanni- l-Sidlo^ual i.abora-
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|-i;ib:i, K and I ibiniatani, 19(-i-l The aMaloiiix ol I'm <niiiii\
j(iji(iniiii\ Biibii I XudibrancliKi-I'lohdoideai. Piibhe:itioMs
of the Sell) \l.iniie Biological I „iboi;itoi-\ 12l2): 41—47.

Biiiii 11 U)(i2. Descriptions ot X'ictoriau Nudibianehiate .\lol-
hiM.i. with a comprehensive re\ie\\ ol the l''ohd;icea.
\leiiions ol the National Mnsenni Mrlboiniie. 25: 95-
I2S,

Cattaneo-X'ietti. R., R. Chemello and li (;iamni/./i-Sa\elh
1990. Atlas ot Mediterranean Xndibnmchs. F.ditrice L:i
Conchiglia, Roma.

Edmunds. M. 1964. Eoliil Molhisca Irom him:iic:i, with de-
scriptions of t\\(i new t;ent'ra iiiid three new spct us Bul-
letin ol Marine Science ol the (aill ;md ('arililnMii 14: I-
32.

Edmunds. M. 196S. Opisthobranchiatt' .Mollusca Irom (ilian:i.
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Edmunds, M. and E. du B.-R. Marcus 1977. On Finnriiiiis
(luritiilus Marcus and Favcriiius hiniiiliiiili.s (.Miiller).
|onni:il ol Molhiscari Studies 43: 200-201

(iosliner T M 19S0. Tlic sxstematics ol the .Veolidacea Nu-
ililirauthia \lollus(:i' ol the Hawaiian Islands, with ile-
scriplions ol two new species. Pacific .Science 33(11:37—

l.ciiuhr il a)id T, I'. Thompson 197 1, Three opistliobraiicli

gastropods new lo the British laiin:r Proceedings .\lala-

coloi;i(:il Socictv London 11: 1 S.5 19.3.
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cnldade dc l''ilosofia, (aencias e l-elras, Universidade de

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b thr I .rssci \iilillcs. Studies on the Fauna ol ( airavao

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lliompson. r, 1'". 19S0. |aiiiaican opisdiobi.iiicli miillnscs II.

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71-79.

THE NAUTILUS 11.5(2):62-67. 2001

Paee 62

Population cKiianiics and growth rate of the turbinid gastropod
Lithopoma americanum (Molkisca) m Bisca)iie Bay,
Florida, USA

Silvia Macia'

Center for Marine and F,n\ininni<'ntal

AnaK'ses
Rosenstiel School of Marine and

Atmosplieric Scienee
Universitv of Miami
4600 Rickenbacker Caiise\\a\
Miami, FL 33149 USA

ABSTRACT

The i;astropod Lithopoma americanum is a common, albeit lit-
tle-studied, inhabitant of benthic communities from Florida to
the West Indies. \ survey of L. americanum populations was
conducted throughout Biscaviie Bay, Florida, USA. Abundance
of the species was highest in the northeastern portion of the
Ba\' and decreased to zero in the southeastern portion. This
pattern may be the result of h\drod\iiamic and salinit\' regimes
that could prevent the recruitiuent and sunival of lanae in the
inner portions of the Bav. Abundance of a population of L.
(imericanum at a permanent site within tiie Bav was monitored
for a three-vear period, during which gastropod densitv de-
clined from 26 gastropods. nr' to zero. The reasons for this
population crash are not cleai", but ina\ have been caused b\
high rates of predation and/or iiitraspecific competition cou-
pled with a lack of recruitment. Ciniwtli rates of L. amcricamim
were higher under laboraton conditions (0.46 mm -week'')
than in the field (0.25 nun -week '), suggesting that field pop-
ulations may be food-limited. Crovrth rate of individuals was
also negali\cK correlated with size.

Additional kei/u'oirLs:
Turbinidae. Aslraea-

Seagrass, ecoliigv, Mollnsca, Clastropoda,

IXT1U)UUCTI()\

The marine gastropod Lithopoma (iiiieriraiiiiin (Gmelin,
1791), formerly known as Astracti (iiiicrinniinit and often
confused with lAthopouw tectum (Ligiitfool, ITSfi), is
found from central Florida, US.\, through the West In-
dies, with occasional, non-reprodueing individuals re-
ported a.s far north a.s North Carolina, USA (Morris.
1975; Emerson and Jaeohson, 1976). IJllioponiri (imeri-
camim resides in both soft- and hard-hotlom coniuni-
nities, including seagrass beds, roekv shores, and gor-

' Current address; Hofstra Univcrsitv .Marine Laboraton.
Box 90, St. Ann's Bay, Jamaica, hunil(?(w j;unaica.com

()

goinan/algae-doniinated hard-bottom habitats ilhizlett.
19S4: McC^^lanahan, 1992). Individuals of this species are
most connnon, however, on seagrasses (ThaJassia testii-
(liiiuDi and Sip'iiiiiodinm filiforme), vv^here thev feed pri-
luariK on t'piplivtic algae (Emerson and [acobson. 1976;
X'anMontlrans et al.. 19S2; X'irnstein, 19S7: Klumpp et
al.. 1992; McClanaium. 1992; MeClanalum and .Muthiga,
1992).

Given the feeding habit ot L. anwricanuni as a gr;izer
of epiplivtes, its feeding aetivitv can have important eon-
sequences for the seagrass community Epiphvtes com-
pete with seagrasses for available light, anil can poten-
tiallv reduce the photosviithetic eapacih of the seagrass
(Howard, 1982; Orth and vanMontfrans, 1984). In such
a case, reduction of epiphvte cover bv gastropods and
other grazers can increase the productivitv of the sea-
grass population, as well as the seagrass connnuuitv as a
whole (Howard, 1982; Howard and Short, 19S6;'()rth
and vanMontfrans, 1984).

Lithopoiiui iniu-ricduuui is also an important member
of the seagrass eonnuunitv as it represents a trophic link
between the prim;in producers and higher-level preda-
tors such as crabs, lobsters, and fishes i H;mdall. 1967:
Targett, 1978; .McClaualian. I992V Nerv little work has
been done on /. aiiwiicaituiiL thus the research pre-
seuteil here proposes to provide basic information on
growth i;it(.'s, population dviiamies, and distribution ot
/,, iimoicdiium [lopulations within Biseavne Bav, soutli-
erti i'lorida. USA.

M.VfKHIAl.S AND MliTHODS

Bis(:a\\i; lU'i Sin\ i:-!

The popuLiliDU densilv of /.. iniliiiciiiiuiii tlniiu',^li()ut
Hisc;ivne i5av. Floiid;i was quantilied ;it v;uious sites in
the ba\ ( ligiu'e 1 i. Most suiAevs vvxM'e conducted during
the summer of I99S, with ;idditiou;il sites sunevcd in
the sunuuci" ol U)99. At each site the number ol /..

S. Mafia, 2001

Faw' 63

Figure 1. Population densih of the ijastropod IJthopotiui
ainciiiiniiiin at \arious sites tiinmij;iioiit Bisca\ne Ba\, Florida.
WP = West Point field site. N'uniliers indicate the mean nuni-
lier ol (gastropods -ni"- at each site. Sites marked 0 had no L
iiiiifriciniuiu, eithei" withm or ontsKJe of the cjuadrats. At sites
with an asterisk (°), L iiiiirriiiiiiiiiii were present in low niiiii-
liers lint ohsened onl\ outside ol the quadrats. LIndeiliiied
.sites were suncxed in snmiiu'r 1999; all otliers sniAoxed m
summer 1998.

(iiiiciiciiiiiiin was (juaiitihcd in ■^ilii witli \isiial (.■ensiiscs
ol 15 haphazard liir (|ua(lrats. \\\ seagras.s hhidcs and
drift algal mats within cac'h i|iiadrat were eareliilK iii-
■spected, but this tccliiii(|iie nia\ haw created a liias
against veiy small individuals. Because the same tech-
nique was used for each sune\. li()we\er. this bias sliouid
not affect comparisons among tlie diilerent sites.

Water depdi at each site was measured, and all sites
were classified into three di'ptli categories: shallow (0.7-
1.5 m), moderate (1.5-2.5 ml, and deep (2.5-.3.7 iiii.
Sites were also qualitatixeK classified b\ substratum into
t\\o categories: sparse' Thnl/is.sui tvstiuUiunn seagrass lap-
proximateK- < 250 slioots m -) and dense 7' tcsfiiiliniiin
(> 250 shoots ni"-). Sites representati\e of these cate-
gories were selected in all regions ol the ba\ (uortlun-ir
central, and southern), so that there was no (i\eiri(linii
geographic pattern to their distribution.

W'lsj- PoiXI Pui'l 1 \ll(i\ D'lWMlC s

i,ung-term i three Ne.irs). moMitoriug oi a L. amcricaimm
po|)ulatioii was conducted at a single site (West l^)int)
in northern Bisca\iie Ba\' (figtu'e i i. 'i'his site is a dense
(o\in- 500 shoots in -) seagrass {T. t('stii(liit>iiii> bed with
deptlis of 0(1 ISO em I \iaeia, 2000). Montlily smveys
were conilucted Iroiii September 1995 to Septemi)er
199S. .\t each siuaia lixc parallel 50 m transects were
laid out 5 III apart. ( iaslriipods were \ isiialK t-ensnsed //(
situ within a 0.25 in- (juadrat at fi\c randomk' selected
points alouij; each transect (except for the first tx\'o sur-
\"e\s, in which oiiK 15 and 10 total quadrats I.'5 and 2
transects], respectixcK. wire coiintedi. .\s with the
abo\i' stuiK, there iiia\ ha\e been a bias against ww
small individuals, but this bias shoukl not allect com-
pai-isoiis among the different months. Shell iieight of
gastropoils (maximum distance between apex and base
ol the shelli within the quadrats was also measured with
calipers.

Growth R\tk Exit ■.him i:\ts

Twent\-fi\e L. fniicricaniini were individnalK' marked
with a distinct s\inbol painted on the shell and main-
tained ill an eiiipl\ oiitdonr tank (80 X 50 X 40 cm)
with a How -lliniiigli siipph ol seawater Previous at-
tempts at niaiiilainiii'j; L. (imcricduiun in other larger
tanks in which seagrass had been planted inihcated that
the gastropotls remained on the sides ol the tanks, leed-
ing on the algae growing there, and did not spend nmcli
time on the seagrass itself Thus, lor ease of retrie\al of
the gastropods, the smaller, emptx' tank was used lor the
growth rate experiments.

Shell height of eacli snail was measured weekK from
September .jO-Deeember 3, 199S. Snails were classified
into 4 size classes: 10.1-15 mm. 15.1-20 mm, 25.1-.3()
mm, and 30.1-35 mm (no snails in the 20-25 nun size
class coidd be found in the fitdd at the time of this ex-
periment). Some marked gastropods died or were lost
duiing the coiiise ol the stiuK. OuK those indi\iduals
lor which at least 4 consecaiti\e weekly measurements
were available were used in the anaKses. .^s a result of
these losses, and because \er\ few individuals of tlie
larger size classes could be found at tht' initiation of the
experiment, the following are the final number ol snails
measured in each size class, rt\spectively: 13. 5. 4. 2. .All
available vveeklv growth increments were averaged for
each indiviilnal, and these values were used to calculate
a mean growth rate i imu-week"') for each size categon'.

HKSl'LTS

Bis( viM-: Bvi SiHVF.v

Individuals of Lilliojioiiiii amcricamnu were most com-
mon in the northern jiart of Biscavne Bay (figure 1).
Sites marked with a 0 in fis^nre 1 represent sites in which
no snails were observed either in the quadrats or in the
surrounding area while swiminiiig about the site. Two

Paije 64

THE XAUTILU.S. Nol. 115. Xo. 2

Table 1. .\iial\sis ot l.ithopoma amcricniinm ;il)un(l;iiicc :i(
various sites tiiioiiglioiit liiscaxne Haw Florida. Assmnjitious ol
noniialih' and lioinosccdasticih were mil met, thus the data
were analv/ed with the nonparainetric Seheirer-Ka\-Hare e.\-
tensiou of the Kniskal-W'allis test.

S( illici ■

dl

\1S

I'

P

water (l<'ptli

.•)

.3.39.44

3.9S

0.029

substratum

1

.3.50.94

4.12

0.0,50

depth °sul)stratum

2

200.66

2..36

Oils

error

34

85.11

site.s (marked 0°) had L. aincncaiiiiiii present but in such
low density- that the\' did not occur witliin the quach'ats
(figure 1). No snails were ohsened in the extreme south-
ern extensions of the !)a\-. .\lthon^h the northernmost
site sampled had a densit\ ol .5.3 ijastropods- m -. most
sites had less than 1 gastropod -ni -.

When testing for the effects of water depth and suh-
stratuni on snail densits'. neither the assumptions of nor-
malit\- nor honioscedasticit\ could he met. Thus, the
non-parametric Scheircr-Ha\ -Hare extension of the
Kniskal-Wallis test was used in lieu ol a parametric t^\()-
wav.\NO\'A (Sokal and I'mliH. 1995). The Scheirer-Ra\-
Ilare e.xtension ranks .ill \alnes (with the lowest value
given a rank of 1 i and uses the ranks to perform an
ANOV'A. Litliopoiitd aiiwriraniiiii was significantK more
abundant at dense seagrass sites (0.5 ± 0.3 (SE)
snails- nr-) than at sparse seagrass sites (0.1 ± O.I
snails- m-) (table 1).

There was also a significant elfect of water depth on
snail densitA' (table ll. but this result is equivocal. When
using actual abundance tiata moderate depth sites had
the highest mean abuntlance (0.42 ± (J, 41 snails-m"-).
lollowed b\' shallow (0.33 ± 0.17 snails-ni -) then deep
(0.24 ± 0.09 snails-m -) sites. When using the rank data.
howe\er, the pattern was rexersed, with deep sites hav-
ing the highest mean lauk (25.3 ± 2.7). followed b\
shallow i2().9 ± 3.0) and moder.ile sites (15.3 ± 2.3).
Finthermore. a separate linc.ir regression aualvsis found
no significant correlation belweeii water depth aiul
abimdance of I, aiiifric/iiuiin (p = O.Sl; r- = 0.0()2i.
Thus, no clear relationship between depth and |iopula-
tion densitv of gastropods can be delerniiiii-d lidiii these
auaKses.

Wi-;si- Point P()Pii.,\ti<in Dinwik s

The highest al)undance observed lor /, iiiiicnciiiiinu ,il
West Point, 26.0 ± 1.2 (SE) gastropods m -, was (huiii'4
the first survey in September 1995 ( figure 2). Ciastropod
abundance declined steadilv during the subsetiucnl sev-
en months, then remained relativelv low (less th.ui 10
gastropods .m-) for the next hvo years. On the final Iwd
surveys density of L. (imcricaiiuin within the qnadrals
was zero, although a few snails were observed dnongli
out the site.

The high<>st abmrdance of gastrojiods coincided wilh

I

I I I I I I I I I I I I I I I I I I I I I I I

S N J M M J S K J Nr \1 J S

J MM J S

I I 'JOS

Figure 2. Aliuud.iuce iuieau ± SE) ol a LilhiijhniKi timcri-
cinuiiii population at the West Point field site in Biseavue Bav,
Florida, from September 1995 to September 199S,

the smallest mean shell size. S.O ± 0,2 nun. Excluding
the two snncvs when onlv one gastropotl was found
(Mav and |nni' 199S). there was a significant negative
correlation between L. amciicaiuun population densitv
and mean shell height (df = 22; F = 23. .SO; p < O.OOOlj,
with an r- value ot 0.53.

Mean size of L. ainciiraiium at the field site increased
at a rate ol 0.23 mm .week ' from September to No-
vember 1995 (increase in mean shell height from 8.0
nun to 9.6 nun in 7 weeks) and 0.27 nun -week ' from
November 1995 to March 1996 (increase in nu'an size
Iroiii 9,6 imn to 1.3.9 nun in 16 weeksV During the three
vears ol the studv a single recruitment event was ob-
servi'd. occurring between |ulv .iiid ( )et(ilier I996(figure
3).

(;ia)v\rii H vn-: Fapkhime.xts

Cu'owth rate ol /, iniiiTicdiniin maiutaini'd in llie l.ibo-
ratoi^v was iuverselv correlated with initial shell height
[di' = 23; F = 11, '.3S; p = 0,003). with ;iii r value' of
0,34, Snails in the smalli'st si/e categoiT grew at the
laslcst rate (0,46 ± 0,06 unn-week ' \ while snails in
tin' largest size category grew at the slowi-sl r.ile lO, 14
± 0,04 mnivveek ') (figure 4).

niscissiox

The .ibnnd.uiee ol l.ilhopomii (niiiiicdiiiiiii in Biseavue
15;iv decreased Irom northeast to southwest, with the
tiieatesl abundance ol I, (iincricaiiuiii in the northeast-
ern corner ol the liav, Xo gastropods were louud in (lie
sondieiii portion ol the bay or along most ol its western
b<H(ler, I'lieie is no clear exphuKitiou lor this pattern.
Seagrass densKv and water depth d(i .illi-el the abim-
dance ol snails, but those areas devoid ol /„ (UiiirUinunn
ineliided sites with both dense and sparse sc;igrass. as
well as sites in all three depth categories, 'i'hus it ;ippears
that these lactors alone ai'e not responsible lor (he ob-
served b.iv-w ide pattern.

\ltliiiii'^li the western side ol Biscaviu' j-iav is snbjei-t

S. Macui. 2001

Fasie 65

O

Oh

o

Oh
Vh

O

d
u

0)

a,

n .l-

9A;5

n:-

n-T 1

||^l -

, A

i 1 'l^

II -1-
0 2-

J

7t

II 4-

n-

■>;■')(.

II 1-

4,4(1

'<:-

ritlfk

ii -1-

S/yi,

II :-

n rf [kVn

II 4-

(),yi>

0 2-

n l-T t{1

114-

7/')(.

■■:-

U4-

S/4t)

u::-^

rTL n, nJTI 1 T-,-,

na-

9/V(,

if-

-rrfl-l

^-"hrlirhn

joHItti-.

-rwfl

lO/'X.

H^TUffl^,

1 :«>(>

^m-^K

m-n-Rn.

ib_

4;'i7

Ml

jm

(,/V7

^^iIktu

n nrm

7;47

-CLL

Hm

? S 11 14 17 20

I I I I I n !( [f III I I I I I I M
? S 11 14 17 211 2?

shell height (mm)

Figure 3. Si/.v rn.'(]iK'iic\ distrihiitioiis Im a |ii)|)iilaliiiii dI
Utliiipiiiiui (iiiiciicinuiin in Bisca\iii.' Ba\. I'ldiuhi, Inim Scp-
Iciiilicr 1995 to August 1997. Population size after Anij;nst 1997
was tiKi small lor ineanin<:;lnl size tre(|ueiK'\ distrlKutions.

to larj^e ciian<2;es in salinitx n\siiltin^ Irdin licsliwatei" ca-
nal disci larjffs troni the urban and aLi;rii'iiltural aieas ol
Miami (Fatt, 19Sfi; W'anij; and Cajfer-Sliahica. 19SS). sncli
changes arc nut a i-ansc ot ini)rtalit\ lui' .idiill /. niiii-i-
icanum (Irlamli ct al.. 1997, identified therein as L tec-
tum). Freshwater discliargcs to the Ba\' nia\ not alleet
adult gastropods, hut the lanal stages of gastropods and
other in\crtelirates are often nuich more sensitixc to sa-
linitx changes (Kinne. 1971). .Sni-Mxal and settlement ol
lanal L. aiiirricaiinin ma\ thus he nt'gati\el\ alfect<'d in
these western areas of Biscavne 13a\^ where salinitv lluc-
tnatioiis are most scM-re (Fatt, 19'SfS; Wang and ('iiler-
Shahica, 19SS).

Recruitment to the imier portions ol llie Ha\ iua\ also
he limited h\ the pre\ailing water circulation patterns.
ll\drod\iiamics in Bisca)ne Bay are dominatetl h\ wind

0 6-

-1

f

<U

01

kw

•5

^"^^ .

^ 1)4-

5d

B

^'~~~--.>,„_^ 4

•-' I1 1-

<u

cS

S.

V-

\.

« "-■■

\.

J— 1

^.

O II 1-

. ,

u

bO

0-

1 1 1 1 1

10 11^ 1< I 2(1 2" 12'^ 25 I .'(I 'n I .■S5

size class (nun shell hciulit)

Figuri- 4. ( aciwljj rale ' iiicasiind as iiii reuse in sliell lieiijit i
(if LillHtjhimii {iiiiriiiiiiiiiiii iiiainlaiiieil iti llie lahoraton lor at
least 3 weeks. XiiimIhis almxe error liars iiulieale ii lor that
size cateiion.

and tide exeiits. which are semidiurn.il ,ui<l eh.uige di-
rection into and on I ol llie I5a\ e\i'i'\ six lioui'S l I ,ee and
Hooth. 1971; Swakoii and Wang. 1977*. When there are
no \\iiRl-dri\cn ciicnlalioii ellects. direct exchange with
oceanic waters is low in the interior and southern parts
ol the f-iax (Lee and liooth. 1971 '. II /., aiiicricaiiinii
populations within the l^a\ depend on lanal input Inini
populations (iiitsidc ol the lia\ lo mainl.iin tliemscKcs.
siicli h\clro(Knaiiiic palleiiis iiia\ prc\enl lanae Irom
reaching the inner lia\ Tins hvpothesis is supported In
the fact that the site Willi llie highest ahnndauce ol L.
(niitiicinniin is also the one most exposi'il to oceanic wa-
ters.

The oilK pi'exioiisK piililislied reseaicli on the popn-
latioii dMiamies ol I .illii>ii<>iiin iiiiirrictniiiiii was con-
diictetl ill the I'lorida l\e\s. .\lc( .lanahan and Muthiga
1 1992) round /- iiiiii'iicaiiiiiii to he the dominant gastro-
pod species in scagrass areas ol Florida Ba\, with a don-
sit\' (if 3.8 gastropods-m -'. (Gastropod ahimdance was
also i|iiaiitilie(l through a sei-oiid iiielhod lliat measui'cd
densitx as the mimlier ol gastropods seen per hour of
seai'cli time, giving a \alue of fi9.3 gastropods lujur '
(McClanalian and Muthiga. 1992'. .\ pre\ ions studx cov-
ering the same area l McC'lanahan. 1992i ri'ported a
liiiiher deiisiU of 200 gasliiipoils ■ hour '. .Assuming a lin-
ear relationship hetweeii data collected with the t\vo
techni(jues. the higher value ol gastrojiods liour ' rep-
resents approxim.itciv II gastropods- m ■ for the sea-
grass habitat

Maxiiiiimi (leiislt\ III /.. iniici'icdinini in Biseavne Bav.
2fi gastropods- III -. was much higher than that found in
Florida 15av. This population densitv was not sustained
loi- long, however. (lr(ippiii'.i lo 13 gastropods-m - in just
twd months, and a^aiii lo 9 gastropods-m - within 5
months. Tw(i ve.us lalei'. llie ahundance of 'gastropods
had declined to ne.iilv zero, and remained so for the
duration of the sliidv. .V concinrent stiidv at tile same

Page 66

THE NAUTILUS, Vol. 115, No. 2

site f'ouiul HI) siii-li dcc-liin' in (lie local popiilatiDii ol sea
urchins, Li/lccliinus varic<^titii.s. nor was there e\idence
of an\ cataslrophic ciiantics in water temperature or sa-
linity'(Nhicia, 2()()()).

No emptv />. autcricnimin sliclls were ohsencd during
the studw sugge.sting (hal niigiafion oiil ot the site and/
or consumption b\' a predator that di'strins the shell
were the reasons tor the sutlden population collapse. In
comparison to se\fral oilier (Caribbean gastropod spe-
cies, ho\ve\er, L iiinriicaiuiDi has the slowest rates oi
mo\ement (nieasnreil as displacement from an initial lo-
cation), averaging ouK 11 cinila\ ' (Hazlett, 19S4).
Tlius, mo\ement ol the species is relati\el\' slow, and
migration out of the site doi's not a|ipcai- to be a likel\
explanation lor tlie rapid didp in abundance of L. iinicr-
icanmn.

Spin\- lobster, crabs. I'asciolarid gastropods, and vari-
ous fishes are potential predators oi gastropods such as
/,. (imericamim (Rand;ill, 1967; Targett, 1978; Mc-
Clauahan. 1992). The I'asciohiriid gastropod Ffiscioltiria
lulipa, the majid crab Lihiiiia diihia. ;iiid the pufierfish
Spliocroidcs testiidinciis were commouK obseived at the
Bisca\ne Bay site, and mav have been at least partK-
responsible for the decline in abundance of L (luivr'i-
cantun. A similar predator-meiliated crash in gastropod
abundance was reported for a population ot Anachis
avara, another seagrass bed resident found in Bisca\iie
Bav, Florida (Hatfield, I9S0). This species demonstrated
large seasonal fluctuations in population densit\' as a re-
sult of high periodic predation rates as well as seasonal
recruitment events (Hatfield, 1980).

Whatever the reason for the initial decline in popu-
lation si/.e of L. ainchcaiium. the occurrence ot onl\ one
significant recruitment event in .3 vears is likelv to have
plaved an iinpoi1;iut I'ole in the !ai-l\ oi recown' of the
population. .^ recruitment pulse began in |ulv 1996,
reaching its peak in October of that vear. This recruit-
ment event was not able to sust;Lin ihc population, how-
ever, as gastropod abundance l)cg;iu to decline steadily
after December 1996.

The growth rate of Z^. aiiicnctniiiin maintained in out-
door tanks was behveeu 0.14 and 0.46 nun -week ', with
smaller snails growing at a laster rate than larger indi-
viduals. The snails at the field site in Biscaxne Ba\, with
a mean size less than l.i unn. grew at appro\iruateK' 0.25
mm -week '. nuich more slowK than the 0.46
mm -week ' obseived for similarK sized snails in the lab.
This discrepancy suggests that /, iiiiuriciiiuiin in the
field are food-limited. T\\f lal)orator\-maiutaiui.'d gastro-
poils were feeding on the epiphytic algae growing on the
sides ol the tanks. These algae ma\ be more ;ibnudant
or mitritious than those found on the seagiass blailes in
the Bav, thus providing the snails with a superior lood
source.

As in the {'"lorida [)opnlation ol /,, (imcncduiun stnilied
here, the congener L. iinilosd (Wood, IS28) from (Cali-
fornia, USA, also e.vliibited an inverse relationship be-
tween population densitv and individual gastropod size
(Alfaro and (iaqienter 19991 This pattern icsulted from

intraspecific competition among the snails i.\llaro anil
(Caipenter, 1999). If the tjuantitv or (jnalitv of food avail-
able to L. (iinciicdinim in Biscavne Bav seagrass beds is
poor, as suggested bv the growth rate data, the inverse
relationship between snail size and abiuidance mav be
caused bv intraspecific competition. Alteniativelv, sucli
;i iclationship could be the result of an umisuallv suc-
cessful recruitment event, followed bv high levels of
mortality (possiblv as a result ol predation pressure),
with tlie remaining individuals simpiv continuing on
tlicii- normal growth trajectory.

Although it appears that intraspecific competition and
pretlation previ'ut Litluiponui (imcrictinuin from main-
taining hi'j;li population densities. (.)Ccasionallv large but
short-lived [)opulations sucli as the one in Biscavne Bav
mav temjiorarilv increase the productivity of the seagrass
commnuities in which they reside. Tlu-ougliout the Bav,
L. tiincriciiiiiiiii provides a temporallv and geographically
heterogeueons sornx'e of (rrazinsr that mav release sea-
grasses from epiphvtic competition lor light. Snail pop-
ulations also proviile a heterogeneous food resource for
higher-lev i-l consumers within the seagrass eonminnitv
food web. Overall abundance ot L rniiciicditiiiii
ihionghoni Biscavne Bay is low. but given the abnu-
daiKc of suitable seagrass habitat in the Bay, there exists
tiie potential for successful recruitment events that
could affect the populations of lioth priinan producers
anil higher consumers in the seagrass couimnuitv.

ACKNOWT.EDGMENTS

.\ianv people helped in the fieldwork reijuired for the
completion of this research, autl I am indebted to them
all: B. Orlando, P. Biber, T Jones, L. Kaubuan. A. Mo-
rales, M. Brown, D. Lirman, [. Montague. .\1. Blanco,
M, liobinson, and K. Irlandi. I thank Mike Robinson and
lv\o auon\ mons reviewers tor pi'oviding ven helpful
comiMculs on eaiiiei" draffs of this m;iiiiisciipt. l-'iiiaiicial
support lor this project, for which 1 am evtremely grate-
ful, came from the NO.\,\ Coastal Ocean Program
.\vvard #N,\37RJ0149. The Sauibel-( :;ii)tiva Shell Club,
.mil the RSM,'\S .Vrionvniniis l^onoi- Awaid.

l.nKKVll RE CITED

All. IK). .\. (.'. ami K. C. CJaipcnter I'^m). liivsical .iiul hiiiloi^ical
piDccsses iiiffiiencing zonation patterns of a siihtidal pop-
uhitioii of the niaiine snail, Astraca (LitJiopoma) tiiidosa
Wood, 1S2S. |()unial of E.\periineiital Marine Hiologv' ;uui
l':ri)logv 24():2.59-2S.l

iMiicrsiiii, W. K. ami .\l. K, j.iiolisdii l!)7(i. i'lic Xiiiciii an Mu-
se ii I Natural I liston ( liiulc In SliclK \ II nil \ KiMipl,

\(\\ \(irk. 1S2 pp.

l''all. I (^ li)S(i. (;aiial Impact on l^isiavne liav Salinities. M..S.
I'lu'sis. tiii\crsil\ ol Miami. Coral (iiihles, 229 pp.

Il.illiiiif Iv H. U)SO, Xalmal I iisloiv and population lliictuution
of the gaslropoil Aiitirliis ai ma (Sav> in a tropical seasjiass
iiahitat. Miami. I''loriila. Bulletin of Marine Seience ■)();
HOI (il2.

S. Mafia. 2()()1

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Hazlett. B. A. 19.S4. Dailv iiiowiiuMit ol some tnipital marine
ijastropods. Marine Belia\ii)rai PliNsiolo^x 11:35— IS.

Ildwaid. R. K. 1982. Impact of teecliiig attixitio.s of epiluiilliic
aiiipiiipods on surface-fouling of eels;ra-S.s leaxes. .Aipialic
Hotan\ 14;91-97,

Howard. !•!. K. ami K. T Sliort. U)S(i. Seai^rass i^nmlli aiul
suni\(irsliip under tin- inllnenee ol i'pipli\tc ma/crs
Aquatic Botany 24:2.S7-:3()2.

Irlandi. E.. S. Macia and J. .Serat\. UJ97. Sallnit\ rednelmn
from freshwater canal discharge: effects on niortalil\ and
feeding of an nrcliin [Liitccltiiuis Vdrii't^iitiis) and gastro-
pod {LithofiDiiKi ti'tiiiin). 15nlletin ol Marine Science (il:
869-879.

Kimie. (). 1971. Salinit\: animals: nixertelaales. hi Kiinie. ( ).
(ed.) Marine Ecologv. Wjlnnie 1. Part 2, \\ ile\ lnteisi.-j-
ence. London, pp. 821-995.

Klumpp. D. W,. J. S. Salita-Espinosa. and M. D. Fortes. 1992.
The role of epiplntic peripiix'ton and macroinxertehratc
grazers in tile trophic fln.x of a tropical seugrass cimnnu-
nit\-. Aquatic Botan\- 43:327-349.

Lee. T.'\ :ind C. Booth. 1971. Circulation. In: Bader R. C.
and M. A. Boessler (eds.l .■Xn Ecologic:il Stuck of Sontli
Bisca\iie Ba\ and Card Sound. LIui\ersil\ ol .Miami. Cnral
Gables, pp. 111-1146.

.\l;icia. S. 2000. The effects of sea urchin grazing and (hill algal
hlooms on a sulitropical seagrass bed counnunit\. Journal
1)1 E\perimental Marine Biologx and Ecologx 246:53-67.

.McClanahan, T. R. 1992. Epibentliic gastropods of the Middle
Florida Kevs: the role of habitat and cinironmental strt'ss
on assemblage composition. |ournal ot Fxpcritnental Ma-
rine Biolo2> and Ecolog\ 60:169-190.

McClanahan. T B. .uul \, .\. Muthiga. 1992. Comparative
sampling methods lor subtidal epibentliic gastropods.
|onrnal el I'Aperinient.il Marine Biolog\ and EcologN' 164:
87 nil.

Morns, 1' .\. 1975. A Field Cnul,- to Shells of the Atlantic and
( inll ( Coasts and the West Indies I liMi^hlmi .Mifflin Coiii-
p.iiix Boston, .3.30 pp

' 'rth. 11 I .mil ] \,in-\h)ntlr,iiis. I9S4. Epiplnte-scagrass re-
LitKinships with .111 emphasis on tlie role of micrognizing:
a re\ie\\. .'\i|ii.iti( Bo!.iii\ 18:4.3-69.

Handall. J. F. 1967 Fixid habits of reef fisiies of the West
Indies. Studies in Tnipieal Oceanographs' 5:665-847.

Sokal H. H an. I |- \ liolilf, 1995. Biometn.' \V. 11. Freeman
and ( .(Jiiip.iin. \i\\ ^ork. 887 pp.

Swakoii. v.. A and |. I). Wang. 1977. Modeling of tide and
wind induced How in south Biscavne Ba\ and Card Sound.
L'ni\crsit\ ol Miami Se:i ( iraiit 'fechnical Bulletin .'57. Cor-
al C;ables. 14 4 pp

Targett. T. E, 1978 l-'oml rescmrce partitioiiiiig b\ the puller-
fishes S)>licicr(Hcl(-< splin^cri and Splini-rDules Icsludineus
from liiscaMie Baw Florida. Marine Biolog\ 49( I !:83-9I.

\aiiMontfrans. j.. R. |. Ortli :iiid S. A. \'a\. 1982. Preliminan
studies of grazing b\ Hillnim i/iiiiiiu on celgrass peiiph-
\ton. Atjuatic Botaus 14:75-89.

\ irnstein. R. W. 1987. Scagrass-associaled iincrtcbrale com-
munities of the southeastern U. S. A.: a resicw. Florida
Marine Research Pnbheations 42:89-115.

Wang, J I), and S \; Corer-Sh:ibica. 1988. The effects of f res! i-
water canal discharges on salinities in Biscaviie National
Park. Report to the National Park Seniee, Miami. 17 pp.

THE NAUTILUS 1 lo(2):6,S-75, 2001

Fa«e 6S

Anatomy of Boonca jcidisi (Olsson and McGinty, 195(S^
(Heterobranchia: Pyramidellidae) from the western Atlantic,
with comparisons to other species in the genus

John B. Wise

Houston Miisciiin o( Natural Sciciuc
One Hi'iiiiaiiii (jixlc l)ii\c
Houston, Texas TT:«(I-I7!)H USA
j\\ise@lnnns.ort;

ABSTRACT

Booneajiiilisi lias a tliirk, t-onical shell witli spnal eouls crossed
by perpendicular axial ribs, wliii'h are absent on the lower half
of the bodv whorl. The smooth, heterostrophie protoconch is
oriented 120° to the teleoconch axis and submerged 30-35%
within the first adult wiiorl. Its head-foot, like those of other
members of the genus, is translucent antl lentiginous, with an
unnotched mentum. The tentacles, with tentacular pads, are
subtriangular, connate, and \entro-laterall\' folded. Tlie buccal
pump is divided into anterior (bpl) and posterior sections
(bp2), with bp2 three times longer than bpl, Boitiicd jddisi is
a simultaneous hermaphrodite and piddiK'es a noii-entieulai-
ized speruiatophore, (Characteristic Icaturcs (e.g., alimentan
tract configuration) allocate tins species, previously included in
the genus Odostomia . in the 'j;enus Koonea. A description of
the genus and a discussion ol the Included .species are pre-
sented.

Additional key unrds functional morphologv, s\steinatics,
new combination. ()d(>\li)iiiiii. P\ ramidelloldea, Heterostropha.

ixTiionrcTioN

In 195'S, Olsson and Mi(iiul\ deseiilicd tlic specie.s
Odostomia {Chn/sairula) jadisi. This taxnn, as well as 5.'3.3
other species and suhspccies iucludeil In llieir work,
were obtained "niostk (roui l-ioeas del Torn island. Pan-
ama. (Caribbean Sea. It is clear IVotii llieir records that
O. jadisi was colleetcd on the island; liowe\er. it remains
unclear exactlv where. 'I'lie inatcrial obtained (lining
tiieir 1953 expedition was from "shore collecting and bv
the picking of bcacb drift, earebillv selected In the field."
Later, Rios (1970) listed this species from noitliern Bra-
zil, while van Rcgtcren Aitena (1975) collected it In two
locations in Suriname. More rccenllv. (y jddisi v\as col-
lected alive at Sebastian Inlet. I'dorlda. US.\, which al-
lowed lor anatomical stiuK and its alloealloii In the ge-
nus Boonca. .\ dcserijitioii ol the genus and spec ies and
a discussion ol species eiiireiilK assigned lo the genus
are iirovided.

MATERIALS AND METHODS

Specimens ol Booiic/i jadisi were collected bv scrubbing
the undersides of inter- to subtidal rocks. Debris were
separated through a series of sieves, the smallest with a
mesh size of 0.25 nun. Mud and sand was rinsed oil
bdiii the inateiial with seawater, and specimens sorted
miller a dissecting microscope. Living snails were main-
tained in bowls of aerated seawater. Snails were photo-
graphed with a Pentax 35 mm camera mounted on a
Zeiss Tessovar dissecting microscope. Sliidls were
cracked with a vise and removed, and snails were dis-
sected whole. Whole animals and alimentan and repro-
ductive svstems, betore and alter excision. v\cre stained
with toliii(hne blue to enhance moiphological detail.
Line drawings were produced under a Leica MZS tlis-
secting microscope with drawing tube. Shells and oper-
enla were cleaned bv sonicatiou. air-dried, coated with
gold-palladium, and examined under a |EOL [SL 6000
scanning electron microscope set at 5-10 K\'. Institu-
tional abbreviations are: .^NSP. Academv ol Natural Sci-
ences ol I'lilladelphia; LISNM, National .Museum ol Nat-
ural Histoi'v. Smitlisoniau Institution: IIMNS, Houston
Museum ol Natural Science.

sY,sri;M viics

b'aniilv Pv r.miidelhdae ( u'av. I S40
Siiblaniilv Odostomiinae Pelseener, 192S
(a'lius Hooiica Robertson, I97S

Botincd belieilsiiii, 1;)7S: .jlil. I'vpe sjiecies: jdiniiiid sciiuillidd
(.',B. .\cl.uiis. IS;>9, bv original desi'j;iiatl<Hi.

Description: Shell thick, eli:ilkv white eoiiie:il. .i-f-i
mm ill leiigtli. v\itli 3~fi adult v\lioils, Whoiis with or
without spir:il c'ords. :ixial ribs, or combin;iti()ii ol both,
lioclv whorl 5()-(i5'/( of shell length. Iniblliens iniimte
cH' absent. Smooth, heterostrophie protoconch oriented
12(1 I 10 to teleoeoiieh, paitiallv snbiiieiged In lii'st
;i(liilt wlioil Apeiinre aiirilorm. with single .leiite eolii-
iiiellar lold. ( )|iei(iiliim I, ill or brown, aiiiilorm, p;iii(is-
piial, with siibceiitiie micleiis.

|. B. Wise, 2(101

Pa<'(.' m

Hi-ad-foot white, translucent, and oiten IcTitiiiinoiis.
lM>()t naiTowinii posterior to propodiuni, widening and
naniiwing again posteriorK to a hlunt tip. Posterior pi'd-
al '^land producing attachment thread. Tentaeles sul)-
triangular, connate ( = conjoinetl at tentacular bases, see
Wise 199fi; p, 455, figs. 2rA-C), ventrodaterall\ folded:
tentacular pails present. Eves suhepitlielial. on median
sides of tentacle bases, ,\nterior iiientum edge imiiotch-
ed and mentuni not bifurcate ( = not biseiled longitu-
dinallv). Intnnert-proboscis aperture on \entral side ol
head, dorsal to meiituiii base.

Dorsal and xcntral ciliateil strips |ommg on mantle
I'ool at posterior cut! of mantle ca\it\. \eiitial ciliated
strip gland composed of uiostK white cells, with a lew
scatteretl \ellow to orange cells, underking 2()-25'/( of
most anterior portion ot \entral ciliated strip. Small, ob-
long pigmented mantle organ (PMO) composed primar-
il\- of bright \ellow cells, although generalK' some num-
ber of clear, orange, green, browii, or ri'd cells present
depending on species. Intro\ert joining buccal sac.
which is composed of sucker, mouth, sheathed st\let
with separate opening, oral tube, and stxiet bulb. Buccal
sac joining buccal pump, which is di\"ided into anterior
(bpl) and posterior sections (bp2).

E.sophagus originating on \entral siirlace ol bpl-b[i2
junction, Salivan' gland ducts entering gut and extending
parallel to one another within walls of bpl and entering
stvlet bulb without exiting alimentan' tract. Salixan
glands not attached distallv to esophagus. Penis globose
or small, with rounded anterior and liiilboiis posterior,
Peni,s located outside and \entral to nene ring, Non-
cnticularized spermatophores transterreil ilnring mating
and usnalK (although see below) attached to posterior
portion of snail's neck (e.g., in Boom/i sciiiiiintld ' or deep
within mantle ca\ibi' on dorsal mantle wall to lell ol e\-
cni-rent siphon (e.g., in B. iinprcs'ia and /■> luMititi filial

Bintiicd jiidisi (Olsson and McGint\, 195S) new

combination

(Figures 1-13)

OildstKiiiid iClinisdllidii) jadisi Olsson and McCIiiits', 1958: 43,

pi. 1, (igs. 11a, 11: Abbott, 1974: 294. no. 3497.
Mcncstlio jmlisi (Olsson and McGintw 19.5S): Ode and Speers,

1972: 7,
Chnjsdlluld j(i(li\i (Olsson and McOintx. 195Si: \aii Hegteren

Altena, 1975: 78, figs, 33a. b. Hi.)v 19V)t: IS7, pi, (il. W'l-

874.

-Material examined: Holot\pe ot Oilostuniid \C'lin/-
Millhla) jailisi Olsson and McGintx 195-8, AN.SP 21 191b.
T\pe localit^: Bocas del Toro L, NE Panama, March
1953. T\pe lot of ()(/()s/(i/;i;(; t('Iin/\(illi(l(i ' iiinhd Hall
and Bart.sch, f911, U.S\M 2232-S4, 7 specimens (a dis-
tinct species examined for comparatixe puiposes), T\pe
localitx': Bernnula. All specimens fortius stnd\' were col-
lected' at Sebastian Inlet. Florida, USA (27°51.fi' \.
S0°26.9' W), b\ author uruler rocks just west ol the
bridge, on the' north side of the inlet ( 199.S-2()0()),
Voucher specimens ol' B. jadisi IIMNS C:at. No. 49()fi9,

Ta.vonomie remarks: Dejong and Coomans (1988^
suggested tli:it the species is closeK related to C/i ;"i/.sr;/-
lidii iiitihii ii):tll and Bartseh, 1911). E.xamiuation of tin-
txpes ol this species ii'sciiled that this .species is con-
chologicalK simil.ii to /> jiidi\i llowe\er, as noted l.)\'
Dall and Bartseh (1911: 2S(ii in their original descriji-
tioii, ,ind b\ ne|oiig and ('oomaiis i 1988), the proto-
concli. unlike those ol most |)\ ramidellids, is scored b\'
spiral grooNcs. The t\pe lot ol C. itioba (7 specimens.
L'SNM 223284) contains a siiecimen, originalK desig-
nated as the figured t\pe . llowc\cr, this t\pe was not
figured in the original description (Dall and Bartseh,
1911:286).

Description: Shell dnd iijicrcnhiin I'iiiiii'cs 7— S '; Shell
white, thick, conical, at least 3.5 mm in length, with 5-
b adult wiiorls (figurt's 1. 2, 8). Teleoconch whorls with
4-5 spiral cords parallel to sutures. ( a)rds of upper
wliorls ci'ossed b\ perpendicular :i\ial ribs, giving a can-
cellate appe:irance (figures l-3i. hitersection of ribs and
cords (forming \v\\ slight nodes) delineating nunierous.
fairK deep, generalK rectangular depressions. Bod\'
whorl appioximatcK 5iV'/t of shell length, witli ouK iiji-
per one-half of spiral cords crossed In' axial ribs, bower
lialf with onlv spiral c-ords, separated by grooves c-on-
stnicted of linear series of irregnlarK' shaped depressions
(figures 1, 2). Oxate, auricular aperture with scalloped
thick outer lip and llnted base. Recessed, single, acute
colnmellar fi>ld on upper half of columella peipendicular
to coluniellar axis. Smooth, heterostrophic, protoconcii
oriented 120° to teleocoiR-h ;ixis. submerged 30-35%
within first adult whorl, with earliest portion of proto-
concii partialK exposed (figures 4, 5), Operculum brown,
auricular, paucispiral, with su!)centric nucleus, lacking
notcli to accommodate colnnu'llar fold, and completeK
closing aperture i figures (i. 7),

Hcdd-fnot and iisiri/d udiw (Fif^un's U-IOJ: Head-foot
transluct'Ut. lentiginous, with scattered wiiile c-ells. Pro-
podium with \eiT .slight medial indentation antl rounded
antero-lateral edges (figures 9, 10). Foot narrowing pos-
terior to propodiuni (pd>. then widening to gradnall)' ta-
per to blunt apex. Pechil gland i producing attachment
thread) within posterior end of (k'c'p and long medial
groove (pmg) extending from miiliUe to posterior end of
\entral surface of foot ifignre 10). Tentacles subtrian-
gular, connate, vcntro-lateralK fiilded. Tentacular pads
present. Exes widelx sp:ucd. bhick. with lenses beneadi
epithelium on median sides of tentai-les ite'. .Anterior
mentuni (me) edge mmotchetl and not longitudinailx bi-
sected (figure 9). Digestixe tissue of xisceral mass pale
orange, xelloxx- broxxn or light brown, witli red flecks.

Miindiitan/ tract (Fiffirr 11 <: Hetnicted introxert-pro-
bo.scis (p) extending posteriorlx from its aperture on xeii-
tral side of head, dorsal to nieiitum base, to enter ce-
phalic hemocoel. Proboscis joining biiccd sac (hs) that
is connected to buccal pump. Buccal pump dixided into
anterior (bpl) and posterior sections (bp2), xxith bp2
three times loiima tli:m bpl: bji 1 round in cro.ss-.section.

Page 70

THE NAUTILUS. \ol. 115. No. 2

Table 1. Species a.ssigned to tlie neiius Booncti Holiert-soii. 197S. .-\ngle of protoeoncli ineliiuiHoii (= deviation scnsu Knii;ht et al.
1960i. "r siiliinerijeiice witliin teli'oconch and ratio.s of hjil and hp2 lor se\('n species ol B(n)iira. Tlie uut anatfimies of/? hiMilurnli'i.
/>' khijiiiiiii and /i' sainiii .iic unknown. NA = Inlorniatioii luit a\ailai)li"

Species

.Angle {")

Snhniergence (%)

hpl vs. I)p2

Boonea l>i.sutiiralis (Sav, 1822)

B. cincta ((."aipenter. 1864)

B. iinprcssa (Sav, 1822)

B. jndisi (Olsson and McGinty, 1958)

B. kinpana Hon and Nakannn'a, 1999

B ok/iiininii Hon and Okntani, 1996

B sciHinudn (C.B. .Vdanis. 18.39)

B. soincri (Aerrill and Bnsli. 190(1)

B siuunui Hon and Nakaniura, 1999

B. iiiitliitnuHitlii Ilori and ()kntani. 1995

120
130
130
120
140
135
120
130
130
135

40-45

30-35
40-45
30-35
50-55

33.3
40-45
40-45
30-35

50

NA

1,0:1.0
3.0:1.0
1.0:3.0

NA

1.0:1.7

51.5:1.0

NA
1.0:1.4
2.0:1.0

thickened along last one-third of its length; hp2 wider,
somewhat lateralK' flattened, distalK roinuleil (figure
11), Esophagius (es) elongated, witli tnherculate surface,
originating on \entral surface of alirnentai-v tract at junc-
tion of l)pl-l)p2. Esophagus extending into \isceral mass
to join stomach. SalivaiT gland ducts (sd) entering ali-
mentaiy tract immediately anterior to hpl-hp2 junction,
e.xtending parallel to one another within walls of bpl,
and entering st\let bulb (sb) without exiting alimentary
tract. Paired sali\ai"\' glands (sgl) not attached distallv to
esophagus. Well-developed vesicles (sglt\) forming tei-
minus of each gland,

I'allidl cavitij: A]iterior mantle eilgc finch plicate, with
small, single dark ceils at suiface and larger, aggregates
of white cells just posterior to mantle edge. Dorsal and
veTitra! ciliated strips joining on mantle roof at posterior
end of mantle ca\itv. \'entral ciliated strip gland com-
posed of mostK white cells, with a few scattered \ellow
or orange cells, underKing 20-25% of most anterior por-
tion of ventral ciliated strip. Small, oblong [pigmented
mantle organ (P.VIO) consisting of ven' dark brown cells
suspended in a matrix ol niaiuK vcllow cells. PMO re-
leasing thick, bright \ellow exudate when snail is dis-
turbed. Kidnev suspended from mantle roof inimediate-
Iv jiosterior to P.VIO. Peiicardi:i! ea\it\ with heart com-
posed of single auricle and \cnliiele, immediatelv pos-
terior to kidney at jmiction of maulle ea\il\ and \iseeial
mass,

Reprofliictivc st/stcDi (Figure 12): Simultaneous her-
niaphrf)dite, with ovaiy and testis within loiiules of single
gonad ( = ovotestis). Gonad filling concave side of vis-
ceral mass, Ovotestis connecled to seminal vesicle bv
narrow hermaphroditic duct. Short duet joins seminal
vesicle to coelomic gouoduct. This |)ortiou of gonoduet

inchides fertilization chamber and convergence points of
seminal receptacle and albumen, mucous and pallial
glands. Reproductive .svstem monaulic. Common pallial
gonoduet extending anteriorlv beneath mantle floor to
open on right side of head, anterior to right tentacular
base above dorsum of foot. When retracted, penis lies
inside cavit>', outside and ventral to nene ring. Penis
with rounded (retracted) to slightly attenuated (protract-
ed) anterior end (atp), widening to ciliated (c), bulbous
posterior (bbp) (figure 12). Posterior muscle fibers (nif")
attach penis to cavitv. Penis extending to outside via me-
dial opening, immediatelv' ventral to nientum, \on-cu-
ticularized spermatophores extruded through gonoduet
opening. Spermatophore placement not obsened, how-
ever, in one individual, it was attached to an opercuhun
(figure rS).

NeiTous si/stem: Epiathroid. Nene ring, including vis-
ceral ganglion (but minus o.sphrachal ganglion and buccal
ganglia), is highlv concentrated and encircles alimentan
tract. The outl\ing osphradial ganglion is coimecteil to
supraesophageal by long nerve extending across nen'e
ring immediatelv anterior to proboscis. Buccal ganglia
attached to surface of prinian' retractor muscle (figure
11, prni' at distal portion ol inverted proboscis,

EeologA' and Distribution: hi silii feechug was not
obseived, althougli on several occasions in the labora-
torv, B. jadisi fed ([uite readilv on Ccriihidco costata
(pers. obs.) Boouca jadisi occiu's from Florida, into Texas
and south to Uruguav (Ode and Speers, 1972: .\bbott,
1974: Rios, 1994).' Rios (1994) reported it on San^assum
in Brazil.

DISCUSSION

Robertson i 197S, U)9(i) h:is aigued that slu'll eli;uacter-
istics provide poor clues loi' developing liv jiotlK'Ses of

Figures 1-8. Scanning electron niieropliotograplis ol sliell and oi)ere\ilnni ol Booiu-a jadisi 1. Apertnral vit'w. 2. Dorsal vii'w.
Scale bar = 600 |xin. 3. Enlargement of upper portion of teleoconeh to reveal detail ol sliell sculpture. Scale bar = 100 |jLin. 4.
Apical view of protoconch. Scale bar = 100 nin. 5. UatiTal viev\ of protoeoncli. Scale bar = 100 |j.ui 0. 7. Opcrcninni. 6. .Mtachment
side. 7. Unattached side. Scale bar = 100 |j.iii 8. Shell with speiinatophore attached to opeiciiliiiu. Scale bar = 500 jj-in. spin =
sperniatopliore.

B. Wise. 2001

Pa«e 71

ra-'e 72

THE XAUTIIA'S. \ol. 115. No. 2

pd-

Figure 9. Livins^ Boonca jinlisi. Sc;ile bar = 1 iiiiii. iiu'
uu'iituui, [xl = propodiiiiii, te = tentacle.

pn)i)iiu|iiit\ witliiii the F\ raniick'lliilac. 'I'iic p\ laiiiiiU'llid
genus Boouea contain.s stneral .species with dilteriini; in-
terspecific shell nu)iph()l()<i;ics (t'.i;., sc\)un>i(hi \s. iiii-
pressa or himiumUs). inchidint:; protoconi'h c()iifiy;nia-
tions. as well as interspecific variations in angle ol incli-
nation anil percent oi protoconcli sul)iner<j;ecl witliin first
teleoconch whorl (Table 1). Moreowr. it lias vet to be
snfficientiv determined it protoconcli nii)ipholog\- is a
reflection of plnlogenv or developnieiital mode (for con-
trasting opinions see Robertson, 1978, anil Wise, 1996).
Hegardless. anatomical st\iilies clearK nnite these taxa
(Robertson. 197S; Wi.se, 1993, 1996; Ilori and Okutani.
1995, 1996, Hori and Nakamura, 1999). Cnrrentiy, the
anatomies oi 7 of the 10 species assigned to the genus
are knowii (Tabli' 1).

Members of the genus Bouiicd possess apomoiphies
(e.g., mentum configuration, leiilaeiikii' pads, and open-
ing of the extroveit/proboscis at the base of the dorsal
surface of the mentum) of the subfamiK Odostomiinae',
while a unique gut moiphologv is an autaponioiphv of
the genus and its configuration (figure 13) distinguishes
this taxon. .Xs in manv pvramidellid taxa. the buccal

' Hi'Cfiitly. Sclianilcr ct ai. (1999) placed lioiiiiiii in llic siil)-
fainily Clirxsallidiiiae, family Odostoiiiiidae iwliciviii lln\ di-
\ide(l tlic P\Taiiiidelloidfa into fi families and 1 1 snlitaniilies).
Tlie value ol this new classification remains luiUsUd. particn-
larly because the authors offered no satisfacton explanation lor
their enlarged taxononiic scheme. A scheme wliicli lollows
names proposed by Sanrin, 1958. 1959. iJronn. ISIS, .mil I'cf
seener, 1928 and not milike dicsc earlic-r workers, have made
this leap uithont an imderpinnlng ol livpollieses of honiologv
based on elearlv defined eharaeters and their states. Tliei-ef()r<\
lor the time being, I choose to work uilliin the admitteilK
narrower interpretation of the snperfamiK willi onK 3 families.
one ol which, the P\ramidellii!ar, includes all p\ raniidcllids.

pmnp is divided into 2 sections, separated bv their oxer-
all sha|ie and the junction of the esophagus. However
with llie exception ol the undivided I'sophagus (e.g.. also
present in certain species of the genus Odostomia^ and
the iuti'rspi'cific variation in l)pl/bp2 ratio (Table 1' the
above-described configuration of the buccal pump is di-
agnostic of the genus (e.g., figures 11 and 13: Hori and
Okutani, 1995: fig. 9, 252).

As ili'scribed for other pv ramidi'llid taxa i Hori anil
Okutani, 1995: Hori and Nakannna. f999: Wise 1996:
2000), Bootica species have a repioduetivc tiacl consist-
ing ol a conunon mouaulic, pallial gouodnct. that ex-
tends anteriorlv beneath the mantle lloor to open ou the
right side of the head anterior to the right tentacular
base above the dorsum of the foot. This is contrarv to
Robertson (1978) who described the "female pore" in
Boonea as bi'ing on the neck of the snail. The penis, as
in the pvramidelliti genera Sai/clld. Iloiihricka. and Pi/r-
innidcUii. is located in a cavitv beneath the mentum. an-
terior and ventral to the uei"ve ring. Hori and Nakamura
(1999) suggested that because the penial complex was
beneath the mentum in B. siioana this species nnist be
assigned to the gen\is Booiii'ti. However, as stated above,
that arrangemenl is not uiii<]ne to this geinis. In Baoiicd
spei'ies, thi' pi'uis is globose or tvpicallv small, with a
rounded anterior end that narrows, then vvidcTis. becom-
ing large and bulbous posteriorlv (Robertson. 1978; Hori
and Okutani, 1995, Hori and 'Nakamm-a, 1999; Wise.
1996; figure 12). The penis is attached anteriorly and/or
posteriorlv to the floor of the cavitv In luuscle fibers ;ind
opens to the outside via a medial aperture beneath the
mi'ntum.

Ill Booncu scmiitiula. B. iinprvsxa. ;uul B. bisittumlis.
non-cuticularized spernuitophores are attached to the
ilorsal mantle wall to the left ol the excurrent siphon,
deep ill the nuuitle liivitv. hall-wav betwei^n the siphon
and luiterior end ol the iiumtle i';iv itv <ir ;it the neck
(Robertson, 1978). In a previous ;inalvsis (Wise. 1996:
488), I iiicorri'ctiv combined the speriuatopliore attach-
ment preferenies in the genera Far^oa and Booiicd.
Species of FiiriiDii ;itt;ich their speruKitojihores to the
shell, tvpicallv :it the basal disc on the apcrliir;il side ol
the last whorl, posterior and to thi' right ol the posterior
end ol the aperture (Robertson. I97S). .\i'cording to
Robertson (1978) sperm;itopliores produced bv Bootiea
species arc small (e.g., in B. sciiiinuild ;iiid B liisiiliirali'i)
to vciv small (e.g., in B. imprc.'^sa) relative to shell size.
.Ml ;ire iioii-eiiticulari/ed. without basal discs. :ind bulbs
lie;!!' or ;it (he unatt;ii'lied ends. In B scitiiitudd. ;iiid oiilv
in this specii's, a l;iter,il spout is present in the sper-
nuitophores, while ill B. hi.siiliii'dli.s ;uul B. iiiij)rrs.sa it is
;il)seiit. The spernuitophores ol B. jddisi resemble those
ol /)' lii\iiliiifili\ I Robertson, 1978: (ig. 42, 370). Sper-
iiKildpliiires luive not been repiiitcd lor the other Sjiecies
ol Biioiicd

The nenoiis svstein in Booncd spei'ies (minus the os-
phradial ganglion and bui'cal gangli;i. but including sub-
and siipiMcsophageal g;inglia ;iiid \isccr;il g:ingli(iii) as in
other pv I'.imiilcllids is comprised ol a hi'j;hK i-oneeiil i;it-

B. Wise. 2001

10

Pmg

me

Figures lO-l:}. Hooiira jmlisi 10. Niiilral siirl.uc n\ IcidI Scale liur = 25(1 p.iii II. Mininilarv tracL Scair liar ^ odd |a.ni, 12.
Diagram <il penis. Scale liar = 100 |iiii 13. IIcmI ami aliiiieiilan trael n\ B Miiiiiiuihi Seale liar = 500 |xiii. atp = anterior end
of penis, hlip = Imlbous liase ui penis, lipl - liiueal |)iniip I, lip2 = Imeeal pump 2, l.s = linecal sae. e = eilia. es = fsopluiiius.
me = mentiim, niF = muscle fibers; p = pmlidseis, pd = pKipdiliiim, pint; = pusleiicji medial ^kkuc, |)rni = priinan retractor
mii.scle (attaches at coliimellar muscle), sli = st\lil Imlli. sd = sali\aiA gland duel, sgl = sali\an gland, sgltv = salivaiy gland

terminal vesicle

ed ring witiiin the lieail. Tlie ring t'licircles tlic aliineii-
tan tract ( Ilnher. 1993: Wise, 1996)). Tliis .s\st,in is de-
scribetl as epiathrnitl because the pleural ganglia lie ad-
jacent to the cerehral ganglia. Tlie presence ol the (is-
phradinni and its garLglion on the snail's Icll side suggests
it euth\neni"iius condition ( = nntwistt'd condition)
(Fretter and (irahani. 1949; Ihis/priniar. UJ-SS).

All Boiiiica species are tlie ectoparasites ol a niunlier
of inxcrtelirate (niostK' niollnscan) hosts (T:ililc 2). The
genus includes species lidiii the Pacific ()ee;ui (5) mid
the Atlantic Ocean/Cnll oF Mexico (5) (Talile 2), Vw-
douhtedk the innnher ol recognizalilc species ol Booiiiti

will increase as our iniderst.uidnig ol tins hiniilv im-
proves.

AC:KX()\\ Id'.DCMI'ATS

1 tlianls Drs. .\1.(;, ll.nase\\\(li ;ind V. Honclu't tor pro-
viding specimens oT p\ r.niiidellids Ironi Sebastian Inlet,
Florida. I also thank .\lr. J. liarrish and Dr. J. Hicks of
the Texas Children's Hospital. Houston, Texas for mi-
croscope lime ;ind iuxaln.Llile .issistance. The critical
comrniails of M. Wise. S, ilvan, ant! two reviewers im-
pi(i\ed this endiMvor. 1 ;nii particularly gratelnl (as a!-

Page

THE NAUTILUS. \ol. 1 15. No. 2

Table 2. I'lilili.slicd iiiloniialion on liosis and i^coiinipliical
ranL;cs tor species ol Hnniu'ii

Boonea impresxa ,Sa\. IS21 ■; ('ra.ssostrca tirfiiiiica. Hillioliim
variiim. Crepkhila amvcxa. l' rosiilpbix ciiicrcii. Tiipliora iii-
f^rocinctd, "polvcliaete" worm. Molinihi sp.. Far'^oa cL nihlmsa,
Cciikciisin (leiidssn (Sclieltcina. 1965; Holicrtson and Maii-Las-
toNifka. 1979; Wise. 1993); Massacliust'tts sontli to Floridii;
Texas. IJS.\ and Qnintana Roo. Mexico,

B. hisitltirdlis iSa\. 1822); l.illdiinii lillnnii. I'niKdlpinx, ci-
iicrcii, Ih//in/is.sti cbsohid. DiiislmiKi iiIIitiiiiIuiii. Ndssariiifi tri-
vittiitns. C'ri'pidiihi fornicdl/i. ('.. phina. (' cuincxd. C mssosl red
ririiiniid. Mi/tihis rdiilis. Mcn-ciuind iiirrciiidiid. Ar<^i>pfcfcn
irrddidits. ('nii'ihulnin stridltiiu. MiHlinlu^ miHliahis. Pldcopcc-
tcii lud^clldiiiciis. Ccukcnsid dciiiissd. C.rcncUd drciuirid (see
Koheitson and .\lan-Lasto\icka, 1979); New Brvni.swick. Can-
ada to New York. USA.

H. seminuda (C.B. Adain.s, 1839); Did.sloiud dllcnidtiim. Civ-
piduld fornicdtd. C. pldiid. Cnissostrcd i:ir0iucd. MciTciuirid
mcrcendria. Arfitijxctc ii irnuhdiis. (.' 'jjhhus. Cnicil>uliiiii stri-
dliim. Modiolus inodiohis. Vldcopciicn iiid'^cUdiiicns, Mi/tilii.s
cdiilis. Urosalpinx liiicrcd. Ili/diidssd oh.soletd. Ndssdriiis trivil-
tdtus (see Robertson and Man-L:isto\ii'ka. 1979); Prince Ed-
ward I., Canada .south to Florida; Texas. USA.

B. cincta (Caipenter, 1864): .\slri-ti iiiidiKd. A. ^^ihhcrosd. Nor-
ri.sid uorrisi. Hdliofis sp. (see l,;il''iillelle, 1977); Santa Barbara,
Calilomia, US.*^, soutli to ('.ulf of ( l;ilil()rnia, Mexico.

B. jadisi (Olsson and McCintx. 1958); Host ;// situ unknown;
in the laboraton-Cc/-;7/i(V/iY/ costdtd (this study); Florida, into
Te.xas, USA, and south to Urni^uav.

B. someri (\errill and Bush. 1900): unknown: Berniiid;i soutli
to Texas. USA (see Ode. 1993).

B. iiiulxtiiiocohi lion ;Ln(l ()kul;nii. 1995: I'liilxniiiiiii iiioiiih-
ffnim. (see Hon ;in(l ()kut;iin. 19951; Tonnoka Ba\. Ixunia-
nioto. Japan.

B. okamurui Hon ;ind ()kut;ini. 1996: ('(dius okamolni. C.
sdZdukd t/oshioi. I'.. Jidiiilds (see Hon and ( )kntani. 1996);
Okataura Beach, Hachijo Iskuid, |.ip.in

B. Huoana Hori and Nakaniura, 1999; Bdiiidlia i iicsci'iis (see
Hori and \ak;nnnra, 19991; Vaniat;uchi Prelectiue. |;ipan.

B. kinpinia Hori ;ind \;ik;unnr;i. 1999: proh;iliK Aliiiiti pcr-
tindtd (see Hori ;in(l \;ik;iiiMua, 1999): '>ania'j;uclii Pi-electure.
Japan,

ways) to Dr. M. Rice and her st;iri at tlic Snutlisdinaii
.Marine Station at Ft. Pierce, Morida. This is Siiiithsoii-
ian .Marine Station Contrihiilion No. .515.

LITERATI) iu<: (:rn<;n

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liulieilsoii U I97S Sperm, ilophoies ol si\ {iislein North
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nal of Molluscan Studies 66: 199-124.

THE NAUTILUS 115(2):76, 2001

Paee 7R

Notices

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FHE NAUTILUS

Volume 115, Number 3
October 31, 2001
ISSN 0028-1344

A (juarterhj devoted
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TH EtoNAUTI LUS

CONTENTS

Wtiiniic 1 15, Xiunhcr 3

October 31, 2001

ISSN 002H-J344

D. L. Geiger The idenliU i>\ Hulit>li\ \li>iitiiliiiiioriiiis Rccxc, ISKi linm

B. Owen the Meditei laueaii Sea l( iasliopmla: Netiiiastropoila:

Halioti(lae) 77

/\ntoni<) Bonfitto .\ii\li'i>(l(ijiliiiillii (/ro/ri/i oxiv new species I Casliopoda:

Mauro Morassi 'Inmdae ' Iroiii Veiiicii l!eil Sea. with ikjIi's on .\ r//rev//s

Bruno Sabelli (MeKill. H)()Hi 84

J. R. Slono A chalistie anaKsis i>l speeies n) Ltnuhis I ( ^astrojioda:

Stromhiilae) 90

Yuri I. Kanlor Redisco\'en n\ CankUa dorri W'atteliled, 1SS6. with

Richard Neil kilburn discussion (it its s\steiiiatic jiositioii (CJastropotla:

Neoi^astnipdda: Xassariidae: Wiwodaiilii^ 99

Jeffrey C. Nekola S\steinatii's and cc()l(i'_:;\ nl Cdstrocoptd [iUislrocojitd*

Brian F. Coles nyf^iTsensis ((iastropoila: Pu|iillidae), a new species oi laud

snail from the Midwest nl the United States of America 105

Erratum 83

Book Re\ iew 114

, '' °3'^°^nic ins:.;j;,Qi

NOV 1 4 2001

UTililA'V]

^f T^kW^^

t/

K 1 w f' ^H

Wt>^ V

Tliis pulilication is sponsored in

part l)v

till' State ot Florida. Department

of State.

Division ol Cnltuial Affairs,

and

the I'lorida Arts ( !onneil

THE NAUTILUS 115(3):77-83, 2001

Page

The identity of Haliotis stomatiaefonnis Reeve, 1846, from the
Mediterranean Sea (CTastropoda: X^etigastropoda: Haliotidae)

D. L. Geigei

Research Associate, Santa Barbara

Museum ot Natural Histon
2559 I'lirsta del Sol Road
Santa Barbara, CA 93]()5 USA
cLLgeigerCg'hot mail. com

B. Owen

V () 1^()\ (id I

Ciialala, ( :A ;r)H5 USA

bu/abiiianln'nicn.on'

ABSTRACT

Haliotis stoiiKitiiirfonnis Ree\e, LS46, is slii)\\n to be a sniior
S)iionym ot H. ncfllcctci Philippi, LS48. A Icctotspc is here di-s-
ignated. Soft parts, radula, habitat, and behavior of the species
are described and compared to those of other oblong species
with which //. \l(iiii(iti(icf('n)us has been conhised in the past.
Size at maturit\' is estimated to be 21.95 mm using shell mor-
phometries. The species is confirmed as a fourth species in the
Mediterranean Sea liased on discrete differences in shell mor-
phologN' and epipodial moi-jiholog)'. The species is restricted to
Malta and SiciK,

Additional kcij aord.s: Gastropoda, Haliottilae, Mediterranean,
taxononn. soft parts, radula. shell, behavior

IXTHOlJUt.TlON

The familv Haliotidae is a vvell-kiiovvn group ol gastio-
pods, paiticularlv due to the economic inipurlance of
larger species in the seafood indnstn'. .Accortlinglv. the
focus of most studies has heen on the conmiercial spe-
cies found in teni[ieiate waters, whereas ihe tropical ali-
alone species havi' received less atti'ntion. Tiie global
treatment of all ahalone tiixa of Geiger (UJOS; 1099;
2000), Geiger and Poppe (2000). and (ieiger and (iioves
(1999) has renewed interest in this familv. As was to he
expected, some of tlie more ti-ntativc conclusions ol
Geiger (1998) have heen confirmed, vvliile othi-rs are in
need of revision. Here we correct the record ol the high-
ly contentious taxou }lidioti\ stoinuruiffonnis Reeve.
184fi, with hearings on a seconil tavon. //. ur<ilccta Phi-
lippi. 1S4S.

The impetus for this reappraisal came from additional
material recentK' obtained In Buzz Owen from lour col-
lectors on Malta. This snhstantiallv increased llie num-
ber of known specimens and included six animals. ( juii-
paiison of this material with the tvpes ol // sldiiinliac-
formis by Buzz Owen confirmed his initial susjiicion thai
it is a senior synonym of H. nc<ilccia^ Here wi' jiresent
detailed arguments and make comparisons wiUi similar
species, as well as sympatric species.

.MATERIALS AND METHODS

.\bbl■e^iati<)ns <)f C^olleolions: B.MN'H: The Natural
Histon Museum, London; N.MW: National Museum ol
Wales,' Cardiff; SBMNH; S;uita BaH)ara .Museum of
Natural Histon; USNM; United States National Muse-
um. Smithsonian Institution; W)V.: Buzz Owen (Collec-
tion. Cnalala. (California; DLG; Daniel L. CCeiger (Col-
lection, in SBMNH: JKC; [oan Koven (Collection, Astro-
labe Inc., Silver Spring. .Vlanland; KAS: Katherine .'\.
Stewart (Collection, (Carm<-I \iillev. (California.

Material Examined: Lots in nniseinn t-ollections are
listed m (Ceiger < 1999; 2000! with adcUtional material in
HOC. Here we onlv ideiitilv specimens by collection
nmiiber if relevant. Iliilintis stoiiuitidffnnnis: >200
shells, fi animals (3 alive in ;i(|n;irinm ol .\. Wright, .Mal-
ta), all from Malta; four shells from (Catania, Sicilv, Italv
(leg. Danilo Scnderi, B()(C, L.UCM 1.52723). Haliotis

lidnnulata Linnaeus. 17.5S. Ir tli<' Mediterranean; >

1.000 shells. > 750 animals. Utiliolis squnnuila Reeve,
bS4fi: > 250 shells, > 50 ;niimals. Haliotis clissoiui (Ire-
dale, 1929): > 100 shells, 2 dried bodies.

Standard scanning electron microscope (SEMI tech-
niques were emploved; for details see Geiger (1996) and
Stevv;irt ;uid (Ceiger (1999i. The radula terminologv' fol-
lows that usi<l by (Ceiger (199fi). Statistical analysis was
carried out willi St;ilisti(;iM;ic I.I iStatsoft. 1994).

SYSTE.M.ATICS

Haliotis sttuiKiliaijonni^ Re(>ve. 184(i
(Figures l-t. 7-fl. IH. 19, 20 >

llaliolis '.loiiialiaiiorinis Ri'cve ^fS46a; 57). — Reeve 1 1 S4(ib;
species 7.3. fii;. 74). — Sovverby (1882: 2(i-27, pi. 3, figs.
22-23, pi. 14, fig. 113).— Kaicher (1981: 2897): copyof
Reeve's figure. — Ubaldi (1993: II, 3-2). Listed ;js unspec-
ified svnonvni. — Geiger (2000; fi.3-64, 68, figs. 14. 147,
150).— Geiger and I'oppe (2()()(); 88-89. fig. 74. pi. 2 figs.
1-3. 5a-b).— Owen et al. (2001 ).

not: Haliotis sloinatiarfonnis. — Wagner and .\bbott (197S:
()()-2()l) 1= H rrtnV/|.— Talmadge (19(i3; 1.34-1.35. fig.
5) [= H. irtri«|.— Ostergaard (1935; 7, 34) [//. sp.].—

Page 7S

THK XAlTILrS \ol. 115. No. 3

Figures 1-4. S\\v\h ot H. sioiiuitiiicfonnis. 1, 2. Bahar-ic-ciiirluui. .Malta, 2-3 m, 6 Nov. 1999, length 24 mm, BOC 15. 3, 4.
Sciutu il-Kl)ir rocks, verv clo.se to the tiiiv islet of F"ilfla, .South of Malta, 2 m taken live, 7 Mav 1995, length 24 mm, BOC 4.
Figures 5-(i. Shell o{ Hiilioti.s (lissiiiui. Site 142. Astrolahe Reef, Fiji, length 17 nmi. |K(;.

Casto de Elera (1896: 424) [= H. sp.].— Pilsbn' (1890:
89, pi. 49, figs. 30-35) [= H. sp.].— Fischer and Fi-
scher-Piette ('l939: 263) [= H. sp.].— llahe and Kosiige
(1964: 7) [= H sp.].
H. ncuJccta Fhilippi (I84S: 16).— Pliilippi (IS47-IS51: 91-92.
pi. 9 figs. 4a. 4b: .see tieiger 11998: note 8] for error in
figure lalieling). — Ubaldi (1987: 285). Reference to spe-
cie.s as v;did, but not figured.— Ubaldi (1993: II, 3-1).
Listed as unspecified sviionym. Geiger (1998: 96-98, figs.
20-21).

Type Material: Three .speciiiien.s of H. stouiatiacfor-
mis are deposited in the I5.\1.\II ( 195().:3. 16.22-24). One
ol the specimens (36 iiim length) corresjionds in size to
tlie one llhi.stnited hv Hee\e (lS46h: pi. 17 Rg. 74) and
bears a pajier label in(htating "T\pe". Thi.s .specimen
(Geiger and Poppe, 2000: pi. 2 fig. 3) is here designated
as lectofspe; tlie otlier t^vo specimens (Geiger and Pop-
pe, 20()(): pi. 2 figs. 1, 2) become paralectot)pes (shell
length 30 mm and 45 nmi). .\11 three specimens have a
similar coloration and correspond more or less to
Heeve's fignn>.

sp^

Discussion of Synonvmv: llaliotis sti>iniitiacfi>n)iis is
a senior .synonvm ol //. nc^lcrln. The t\pe material in
the BMNII consists ofspecimens of the Medileiranean
species known now from Malta and SiciK. The tspes
have several characters in cominon with recentk eol-
lected, Mediterranean specimens. Direct ciiiii]iai isnn
witli tile t\pe of//. tw<ilvcta is not possil)k', because the
vviiercal)onts of tiic tvpe material consisting of four spec-
imens (Philippi. 1847-1851: 92) is imknown. It could not
be located in Bremen, where some of Philippi's material
is deposited (J. Lehmann, Unixersitv of Bremen, pers.
comm). The species is lound at the t\pe localitN off/.
uc<ilccla (Sicily) although it is much mon> common in
.Malta. P'urtliermorc the two original ilescri|iti()ns (see
below) show a number ol close similarities, which are

indicated b\- numbers in curl\- brackets: |1) the o\erall
oval oblong shape of the shell; {2j the conve.\it)' of the
shell; |3[ the spiral sculpture; 14} tiie fine folds seen in
the juxt'uile part ol the shell: |51 the ven' eccentric spire.

Translation of Original Descriptions:

"Species 73. (Fig. 74, Mus. C^nming. )

llaliotis stomaiidcformis. Shell oblong ovate |1|, xeiv
conxi'x 12], spiralK striated |3), radiallv fineK' plicated
(4|, spire nearh' terminal (5|, elevated; five perforations
open: marbleil with oli\e and green.

Lives in New Zealand."

"76. Haliotis iicjilccta Ph. liaUotis with a o\al oblong
shell |1), somewhat convex, with spiral striation jSj,
white, with red maculatioii; with 4-5 open holes; part
lietween the holes and the margin convex (2); spire
small. side\v:ivs |5|. Length 1 1'"; width R-6.5'"; height 3-
3.5'".

Home: Sit ilv. rare.

Spii'al grooves and lidges approximatelv 34-32 be-
tween spiie ;nid holes, approximatcdv 7 between the
holes and the uuu'gin: little conspicuous, radiating tolds
finalK extinguished towartls the spire 14|. Differs Irom

voung //, niiiosfi |i,am;u'ck. 1S22

or

R

eeve.

1846':^

an(

liniicllosd 1' bv (he more iiairow lorni illdl /'(/giw/ 11'"
long but 7.5'" wide' 2' bv the nmeh snuiller spire |5|, 3)
bv the p:ni between the spile :uid the holes much more
convex 121. I' bv the p:irl between the spii'c and the
margin si mi lark em i vex. not at all giooved. and 51 bv the
ledge can"\ing IIk- holes mil distinct: ;ind also 6' bv the
less picniiinenl spire.

Description: Sliidl (figures 1-4) arched, medium
vvciglil oi- ihickness. nuLxinnnn size 45 nun (jiaralecto-
tX'pe). Spiral cords smooth, ol variable stieiigth, 20-35
between suture and row ol holes. One slicing cord usn-

D. L. Ceitrer and B. Owt'ii. 2001

Paa;e 79

Figurt's 7-15. SI-'.Ms ol ladiilac 7-11. llnlinU^ \hiiiiiili,iijiiiini\ ISOC 7. Oxcnicw (il i.uliil.ir ribbon, scale bar = .500 |jliii. 8.
Rac'liidiuii ami hit. ml tdotli I. Scale bar = 1(1(1 jxiil 9. l.alna! Icrtli .5-5, Sc.ilc bar = Kid (iiii. 10. Cusps of inner marginal teeth.
.Scale bar = 5(1 ijini 11. Cusps (if'uiiikllc ruart^iual tcetli. Scale bar = 2(1 |jlim 12-1.5. Kacliidiau and lateral tooth 1 of some oblong
abalone species. 12. Haliotis clc^ans. US.\.\1 .■5(i(iy4(l, 13. Iliilictis tlissoim. JKC, .Astrolabe Beef. Site .No. SS, in Cnjptoplax lar-
VM'fonuis with coralline algae. Februan 1995; note sliaip angle lormed by cutting edge and primary ridge of lateral tooth I. Scale
bar = .50 )j.ur 14. Haliofis ;J(ihni. Note shaip angle lormed b\ cutting edge and jiriman ridge ollaleral tooth 1. Scale l)ar = 200
(xm. 15. Uiiliotis s(punnata. SBMXH. East of Cape I'resfon. Hetinard Haw Western Australia. Scale !)ar = 200 (xnr
Fiijui-es 1()-18. Representati\e epipodial section (A llnliuli^ spp Hi. IhilUHis sloimitiiicfttniiis. B( K 17. Iliilinlis tiihcrciilatii.
SB.MXli e\ DLC. 18. Haliotis sqn<nn<ita. SB.M.Mi.

allv alternating with nnt' to three hner intermedial
ones; in < \Wc of specimens con.spieiious smooth space
present lietwcen two strong cortls; some with .i(l|acenl
strong cortls. Earl\- wliorls often with low, irregular, non-
colahral folds. Cooler \ariable. Inise color tan. mottled
with green, ri'iL hrown. Holes slightK' ele\ated. some-
what oblong. iisnalK .3-(S open (mode = 4: Fignre 20).
First open liole well in anterior lialf of .shell. .Spire ver\'
eccentric. Interior with steel blue nacre, no Timscle scar.

Kadci.a (Figures 7-11): Kachidian trapezoid, cutting
edge neitlier Straight nor folded posteriorly Lateral
loolh 1 triangular, piiinan' ridge more or less straight,
secondan ridge inserting at 2/.'5 of the height. Lateral
teeth .3-.5 with minor denticles near base of cusp. Inner
marginal teeth slightK as\nniietrical, first dentick" on
outer edge at le\el between first and second denticle ol
inner edge.

Sort P,\irrs (l-'igure Hi : Fpipodinm \c-r\ narrow for

Piiire 80

THK NAUTILUS. Vol. 115. Xc

Gozo

Malta

• F
G. H _

Figure 19. Distribution map and loL-alitics of H. stomatinr-
{iiniiis. XunilxTs in parentliesfS indicate the number oi dead
shells/live animals. A. .St. Pauls Bav (10/0). B. SaJina Bav(6/0).
C. (ihallis Rock (T/2). D. Qalet NJarku i5/0l. E. Bahar-ic-Cas-
haq (83/31. F. St. Juliens Ba\- (3/0). G. Manoel Island. C;7.ira
(7/0). H. Fossa \';illetta (7/0). 'l. Xghagra Zabbar (29/0). J. Mar-
saskala Bay (36/0). K. Filfla Island (2/1). Inset: Italy with its
.southern island of Sicil\'. and marked localit\' for H stciniatiiic-
forinis (Catania: 4/0). Arrow: .Malta Island

genus. Dorsal margin frilled with small projections and
bearing large tentacles, at intenals of 2-3 times the
width of the epipodiuni. Face of epipodiuiii with some
stnbb\- projections. Wntral sick' larger than dorsal side,
witli iiKjre numerous, more narrowK' spaced, but smaller
tentacles. Ventral margin bearing manv partially o\erlap-
ping projections: some projections with bilurcate or tri-
furcate tips; some small flaps further below. Hvpobran-
chial gland with simple himellae.

Dislinbution: .Malta and .SiciK' (Pliilippi, 1S4S: 1S47-
1S31: (;cMgcr, 2()()(): this stud\: figure 19'.

Habitat: Si\ spcc'iinciis wcic taken ;ili\e in 1-2 m
dc'])tii irom the underside ol Hat limestone rocks with ;i
diameter oi less than 20 cm on a bottom of graxcl and
small stones at least 10-15 m from the nearest sc:igrass
(Poskhmia oceauicd | Linnaeus, 1S13| ndile). Larger
rocks of up to I ni in diameter did not h:ui)oi' :m\ spec-
imens. Dead shells were most frequcniK found ;it :i
depth of 2—4 m, to a maxiMuuu of 15 m. Shells Ircuii
deeper water are rre(]ueutl\- of reddish ( olor

Biologj': Some bcha\ioral obs(>natiniis were made in
acjuariums. The species is veiy active at night, last mo\-
iug. and \er\' light sensitixe. It moves nuicli faster :iu<l
is more light sensitive than // fuhcnuldln When^is //
tuherculata will initially clamp dcjwn when distuibed, //.
stonuitiaefonitis iuuiiediately crawls awa\ rapidk. Ilali-
cfifi tiihcmilfitfi is regukuK found in the open at night,
whereas H stoxuitiacjonnis is almost alwaws cnptic The
beha\ior ol //. stoiiuiluicjornm is \en' similar to tli:it of
a recentK decribed species ol ll/ilioli\ (Owen et af.

shell length (mm)

Figure 20. Scatteiplot of nmnber of open holes as a function
of size (n = 200). Note the slight trend of more open holes
with increasing shell size. Break])oint regression identified the
change in slope at 21.95 mm; for shells > 21.95 nun (arrow)
the number of open holes does not change with size (p =
0.11). Insert: histogram of number of open holes for shells
larger than 21.95 nun: mean = 4.2 (SD = 0.66), mode = 4,
n = 146.

2001). One specimen o/ H slotudliacfonnis grew from
approximatelv 25 mm to appioximatelv 35 mm in one
vear.

We were able to estimate size at maturit\ irom shell
mni-phological dat;i. nameK length and the number of
open holes. Breakpoint regression anaKsis on data from
H. tuherculata has shown that the breakpoint tor these
tA\'o \ariables corresponds closely with published size at
maturit\' as deteiniiiied from gonad studies (Geiger,
199S). As in H. luhcrculatn. in H stoiiuitiacfonnis there
is an a weak trend to increase the munber of open holes
with size (m = 0.13), although linear regression anaKsis
was onl\ marginalK significant (r- = 0.016, F, .^^ = 3.24,
p = 0.073, n = 200) o\er the a\'ailable size range (13.5-
42.0 nun). Breakpoint regression auiilvsis with Quasi-
New ton loss function (r- = 0.72) determined the break-
point to be at 21.95 nmi (figure 20), which is taken as
the best estimator oi size at nuituritx. Linear regression
anaKsis for specimens larger tluin 21.95 mm is insignil-
icant (r- = 0.017, F, ,_,, = 2.54, p = 0.11, n = 14fS! with
;i mean number of open holes of 4.2 (SD = O.fifi: figure
20). Haliotis stomal iacformis matures at a considerabK'
suKiller size than //. tuherculata (33 nun), but both ma-
ture at appro.ximalcK luill their m:i\inmm size.

Remark.s: D.X.\ se([uence data oi the Uistone 3 gene
ir;igment confirm that H. stoiiiatiacfonni.s is distinct
liiim //, luhcrculntn . the .32S bp lr:igmcnt shows 23 dis-
ci'ele dillcrenccs. Intcispecilic \, illation between other
species pairs ol llahi>li\ sp. i:iilges Irom 12 to 24. No
intiasjiecific \ariation was detected in similar DN.\ se-
i|ui'nces of three specimens ol //, nihiiiinosa Recne,
IS 10. :uid two specimens ol //, varid Limiiiciis, 1 75S,
((ieigcr, iiiipnbl. d;it:i.).

DISCUSSION

C^oiiiparisons: I luliolis dissoud i I'lgures 5-fi, 13): This
Wcstein l':icific species is disliibiilcd Irom the Marian:is.

D. L. CvU'vr aiul H. Owen, 2()()1

'aee SI

Micronesia, New (^aletlonia. to 'lon'j;a. lis s[iir,il cokIs
are nuieli coarser and their numlier l)et^^^■en ihr sulinc

and the row oi treniata is approxiinatcK 12. e pared

to > 20 in H. stoinatidcfiiniiis. The nniiilier n\ open
lioles is approxiniatek tlie sanu' in // ilissmia i4-5i and
H. stonuitiacfonnis (.3-H. mode = 4'. Th(> coloration is
off-white with lar^e, irre(j;ular c<)Io?-ed aieas in lirown.
red, and i^n^en tones. Finlher illustrations of the speeii-s
can lie lonnd m Kaicher (1981), Geiijer ( UWS: 2()(l()i,
and Ceiger ami Poppe (2000).

R.MJl'L.^ (Fignre 13): The radula shows the s(ron'.,;I\
ansadar lateral toolh I sci-n in some ollirr indo I'acilic
species, such as //. iiniii and //, ;J(ihni (imilni. 1791
(cf. Geiger, 1999, figure 141. Tlie epipodinm and hvpoh-
ranchial gland are cnrrentiv unknown; the two dried
specinu-ns ilid not allow their assessment.

liilliotis stjiKiDKild: I^'onnd predomnuintK m Westeiii
Australia, hut additional specnnens hdiii \ ietnani and
Indonesia lia\c come to light in recent \r.iis iCiei^cr.
2000; D. Reid, pers. connn.; .\. 'iliompson. pers.
comm.). It is mueli larger (7-(S cm) than II. sttiiiidlidc-
fonnis (4.5 em). Compared to H. .stoiiialiaii'oniiis. the
spiral cords ol iiido-l-'aeific specimens ol // siiiuiiiKiIri
are somewhat stronger, and \en much stronger in West-
ern Australian specimens of H. sijunuuitii. which e\eii
bear coarse scales. The nmnlier of holes is much greater
(6-9, mode = 8) as compared to f/. stdiiuitiriffoniiis (3-
6, mode = 4). Tlu' first open hole nl // sijiuiiiuild lies
well in the posterior part of the shell, \\here;is in //
stomatiacfonitis it is found in the anterior halt. The coloi"
is usualK uniform ilark reddish brown to si'pia. Iiregular
folds in the eark whorl are onK' known from one spec-
imen Ironi Indonesia (IDLCJ .\.\B 13g). In most sjieci-
mens the concentric sculpture is confined to stronger
growth lines. The species has been illustrati'd in Hinton
(1978), Ei.senbersi (1981), Kaicher (1981), Wells and
Bn-ce (1985), DJiarma (1988), Wilson (1993), Ca-igcr
(2000), and Geiger and I'oppe (2000).

Radula (Figure 15); The central field is similar to that
of H. stoiiuitiaifiiniiis. Lateral teeth 3-5 ilo not be;ir
denticles. The denticles on the nuirgiiial teeth are s\ in-
metrical.

EpIPODRI.VI (Figure 18): Ilnlintis SijUiniKlid ;m<l // v/e-
matiacfonnis both liavf a r;ither narrow epipodinm. In
H. sijuainald the dorsal ti'iitaeles are separ;ited b\ less
than the width ol the distended epipodinm. whereas in
H. stoniiitidcfurinis tlie\ are sep;u';ited b\ 2-3 times the
width of the epipodinm. The clorsal margin bears finel\
digitate processes on a knobbed, broad lold. The lace ol
the epipodinm is strongK knobbetl. The \entr;il hall is
wider than the dors;il h;df. It bears somewluit siii;iller
and nn)re narrowlv spaced \entral tentacles iind die
same digit;ite processes as the dorsal margin.

Haliotis Inhcrcnlnhi (Mediterranean lormi: lliiliolis
stomatiacf(inni\ is more elongated, and much more
arched. The irri'giilai' radial folds are much we;iker tluin
in the lamellate forms of /V, tubcniihitii. The niimbei ol
open holes (3-9, mode = 5) and the phicemeiit ol the

first open hole in the anterior hall ol the shell an- Nen
similar to // sloiiuil'uicfonnis. The ele\ation of tlie spire
is \;niable kT. ( ;eiger. 2000: fi^s. 12. 13). but the spire
IS more or less perpendicular to the [ilaiie ol the shell,
and in ;i more central position on the shell.

Weink;iiill (188.3: 74. tnmshition) noted in his discus-
sion ol // U(il(ni<li<'<i l\ee\e. I S 1(S = // liihtrciilald
Limuieiis. 1758); ■Sowcrln muted with his //. ■■ilomd-
lidcformh ;i sm;ill beach loiiii ol // tiibcrndtttd Irom
\I;ilta with liigliK ele\ated spire, which Pliilippi hatl
c;illetl //. iicijcctd. Specimens with monstroiisK ele\atetl
spire are not ;it ;ill r;ire in the .Mediterranean. ;uid h;i\i'
been known lor a long time." Weinkauff was certainK
correct in pointing to the high spired forms off/, iubcr-
culdtd from the .Mediterranean (cf Geiger, 2000: fig.
I3i. IIowe\'er. //. sldiiidtidcfoiuis is not distinguished b\'
the high spire, but b\ the ;n-ched shell, and the rather
eccentric s|)ire. Utiliolis liihcrculdid has been illustrated
in K;iicln-r ilOSl: as //. Idnu'lliisriK .Alibott and Dance
il9S3>. Ubaldi (1987). I'oppe and C;oto (1991), Geiger
120001. Gt.i.^er and Foppe (2000), and Owen et'al.
i20()l).

R\i)i i.\: Hachidian tooth tnipe/oid. cutting edge con-
\e\ without ihiekening. Lateral toolli 1 primary' ridge
wilhniil eon(.a\e depression, secoudaiA' ridge snuiHer
than piinuiiA rid'^e. inserting ;it 2/3 ol the height of the
lootli, Later;il leetli .'1-5 without denticles. The marginal
teeth show s\ iiiiiK'triciil denticles on cusp.

Fi'ii'onn \i (I''ignre 171; The epipodinm is ol moder-
ate width, i.e.. much wider th;m in // stnnidtidcfonuis.
The ilorsal margin be;irs ichitiveK small tentacles. bareK'
extending beNoiid the mai<4iii ol the leafv small Haps.
The small Haps bear priman and secondan undulations
and ha\e a hirrowed snrhice towards the face of the
("pipodiuin. Till' imdiilatiiig mid-epipodial fold is well
dexelopeil. Tlii' l:n'e bears papillae 1-4 mm long at \'an-
iii'j; densitit's. The \eiiti;il portion ol the epipodinm is
lar^ir ;md minors in struclnre the dorsal iialf. Some
small tentacles are nestetl within the furrowed part ol
the \eiilial poilioii.

Hist(>i"\: file 1 1 inliisioii smroimdinii //. \ti>iiidti<icfor-
iiiis in;i\ be atlril)nt;ible to the illnstnitiou In Reexc
il84(")bi. which does not show enough tletail. and the
mi(loubti-dl\ erroneous tvpe locality "New Zealand" (cl.
TahiKidge. 190.3 ' Three spec-ies are kTiown from well-
studied New Ze;ilaiid - // mislralis (Imelin. 1791; //. iris
(imelin, 1791: //, i in^iitca (hnelin. 1791 1 and are all en-
demic to the island i^ronp ^(ieiger. 20001. and tlie name
// stumdlidcforiNis has ne\'er again been reported Irom
this area. Talmadge ( 1963:1.35) recognizetl the erroneous
l\pe localitN of //. stomal iarformis (New Zealand) and
di'sign;ile<l .i new one in the Indo-Pacific: "Ncjrth coast
of |;i\;i, liidonesi.i. III doiii',; so he inacKerteiltK' added
111 the confusion. For the most part. //. stonuitiacfonnis
h;is been overlooked, or it has been \ariousK' svnony-
iiii/ed. Where it has been used, it referred to a nuniiier
of different s|iecies. We provide a brief svnopsis of the
v;irioiis applic;itioiis of the name, arranged bv species

Page 82

THE NAUTILUS. \'ol. 115. No. 3

that we tliink was most IikcK al lianil: niir ideiitiHt-atioiis
of these n>eor(ls are olteii \cv\ teiitativi'. because neither
voucher material nor illustrations were axailahle:

Udliolls tli.ssoiui (Ircdalc, 1V)29) — Ostergaard (1935; 7 [as sto-
inatifomis]. 34 [as stoiudldclonius]). referring to speei-
iiieiis ironi Tonga, noted (p. 34): "Ree\e remarks that this
species is well distinguished bv its conspicuous convex ob-
long torni." This makes an identification as H dissonii \er\-
likel>.

Haliotis cxiiiiia Dnnker. 1863 — Casto de Elera (1896) made
reference to a Philippine .species and gave H. cxiffid Dnn-
ker, 1863. another controversial taxon (cf. Geiger, 1998).
;ls a .snionvm. Pilsbn' (1890) sxnonvmized H. cxigud as
used bv some anthois undci // .stoiudtidefonnis. and
grouped H. squdinaUi with //. .stoiiuitidcfonins. The ma-
terial Pilsbr\- (1890) based his treatment on came I'nnn
New Caledonia and Tonga, and was in the 3 cm range.
The description, the si/e of the specimen, and Pilsbn's pi.
3. fig. 4. make it IikcK that he was discussing H. chithrdtd
Reeve, 1846. llo\\c\er. pi. 49. figs. 30-35 of Pilsbn
I 1890). show quite a different species.

Haliotis sijudmatd Reese. 1846 — Specimens labeled H stu-
iimtideformi.s in the BMNH supposediv from the SolouKm
Islands and NMW from \'iti Levu, Fiji, actuallv represent
H. squairidtd. Geiger (199S) sviionymized //, stamdlidc-
fnnnis under H. .sciudiudid.

Udliiitis vdiid Linnaeus, 1758 — Some more elongated speci-
mens of the hvpei"variable //. i/iiid were separated 1)\' Tal-
madge (1963: 134-135, fig. 5) as a distinct subspecies, H.
varia .stoinatidcfonnis. an opinion followed bv Lindberg
(1992). Wagner and Abbott ( 1978) considered the Uvo taxa
synonymous.

Unknovvni species — Fisclier and Fischer-Piette (1939) men-
tioned a specimen from Tanna. X'anuatn. It mav either
refer to H. dissond or to H chithrdtd ^ Habe and Kosuge
(1964) synonvmized unil(>r //. stoinalidfionnis three dis-
tinct species: H. hdiih'i/i Ancev. 1881 (= //, jdcnciisis
Reeve, 1846); H. j)hiiiata Sowcrbv 1882; ;inil //. cri'hns-
nilpta Sowerby, 1914. We ;ire iiii;ilile le e\en guess wliat
species was discussed. Ubaldi considered // sUnndtidcjor-
ini.s either a valid species from the Fainipe;in-Mediterra-
ne;m area (1987) or a s\n(in\in of :l t;i\()ii tlL;it \\;is not
.specified (1993).

Sowerby (I882i on llie oilier IkukI. impressed in a now
vindicated renegade opinion, that H. nc^lccta Phili[)iii.
1848, is either a .synonym (]i. 26-27: "The \ariet\' figured
in our la.st plate represents // ncji-cl/i I'hilippi. and is
undoubtedly identical with our present species [H. slo-
iiuiluicfdnnis]."), or a v;iriet\ leaption hir pi. 14: "IIS.
H. stomotUicformis. R\e. var. nc^cdn I'lr See f. 22,
23. — Malta") ol //. stoinaliacfoniiis. Ilnlinlis nc^hTfa is
itself an overlooked taxon. which \\;is lentati\el\ resur-
rected as a valid species by Cieiger ( HW8) ba.sed on seven
shells from Malta. We are now in possession ol addi-
tional material, including animals, wlneli allows iis to
conclude that H. sloDuiiiacfomiis and //. iic^h-cta lep-
re.sent the same biological species from the iskuid ol
.\l;ilt;i and Sicily in the .Mediterranean Sea, and thus to
confirm the existence of a h)iu-tli abalone species in the
Mediterranean {H. tiihcrculata: II. ])usliihil(i Reese.
1846; Haliotis iniikriiDscusis Owen el al.. 2001 I.

ACKNOWLKDCMENTS

W'e reeei\ecl iniicli ni;iteri;il ol the species discussed
from C. Mifsud. R. Shead. H. Thake. A. Wright. ;iud A.
.Xuereb (Malta). F'urther material was provided In' Mare
Curoua (France); K. Stewart (Carmel \alle\', C^alifornia.
USA); the Schonsleben family (Basel, Switzerland); O.
Kailiea and .\. and L. Bowe (Neiafu, Vava'u. Tonga); R.
Kershaw (Narooma, Australia); [. Koxen (.Astrolabe Inc..
Washington, DC, US.A); A. Thompson (Uni\-ersitA- of
California, Santa Barbara, USA); ;md D;uiilo Scnderi
(C>atania, Sicily, Italy). SEM time was paitl b\- grants
from the Hawaiian Malaeological Societx' and the West-
ern Societs' of Malaeologists to DLCJ. DNA sefjuencing
was made possible through a W. M. Keck post-doctoral
fellowship ;it LAC^M to DLti. Two reviewers contributed
coiistriicti\i' criticism f)f the manuscript.

LITERATURE CITED

Abbott, K, 'I', ami S. P. Dance. 1983. Gompendium ol Seasliells,
E. P. Dntton. Inc.. New York. 411 pp.

Gasto de Elera, R. P. Fr. 1896. C!)atalogo Sistematico di- Toda
la Fauna de Filipinas . . . \'ol. 3. Inipnnta del Golegio de
Santo Tomas, Manila, 942 pp. + l.\i\,

DliariiKi. B. 1988. Siput dan Kerang Indonesia (Indonesian
Shells). PT Sarana Graha, Jakarta. Ill pp.

Ei.senberg, |, M. 1981. A Collectors Guide to Seasliells of the
World. McCiraw-IIill. New York, 237 pp.

Fischer. P.-II. and E. Fischer-Piette. 1939. Gasteropodes mar-
ins receuillis aux Non\elles-IIebrides par M. Iv .\ubert de
la Riie. Bulletin du Museum national d'llistoire natnrelle.
di'uxieme serie. 9: 26.3-266.

CJeiger, D. 1996. Haliotids in the Red Sea. with neotvpe des-
ignation for Hdliotis uuihitrridis Lanuirck. 1822 (Gastro-
poda: Prosobranchia'. Re\uc Suisse de Zoologie 103: .339-
354.

Geiger, D. L. 1998. Recent genera and sjiecii's of the laniilv
Haliotidac (Gastropoda: \'etigastropoda'. The Nautilus
111:85-116,

(ieigi'i. D. L. 1999. .\ total exidence clatlistic aiiahsis ol the
Haliotidac (Gastropoda: Vetigastropoda). Ph.D. Thesis.
I'nixcrsitv of Southern California. Los .Xngeles, xix + 423

PP

Geiger, 1). I.. 200(1. Distribution and biogeograpliv of the Re-
cent Ihiliotichie i (;;istro])oda: \etit;astnipoda) world-wide.
Bollettino Malacologico 35: 57-120

tJeiger D. L. and L. T CJrovcs. 1999. Re\ iew ol liissil abalone
(CJastropoda: \'etigastropoda: Ilaliotiilae) with comparison
to Recent s[)ecies. |onrnal ol Paleontologv 73: 872-.885.

Cieiger, D. L. and G. f Poppe, 2000. I'amily Haliotidac. hi:
Pojijie, G. T. and K, (iroli il'ds ' \ ( ,'on(iiolo<jic;il Ico-
uograpliN. ( .'oiK'libnoks, ll;ickeiilieiiii (ieriii,ni\ 1 3.5 pp.,
S.VpIs,

llalie. I', .111(1 S. Kosiige, 1964. List ol llie liidii-P.u llie Mol-
luscs, Goncerniugto the Japanese Molluscan l'"auua I, Sn-
peiiamiK Plemi)lomaiioide;l. \;Uional Science MnsiMUii.
I leiio Park, 'lok\o. S pp.

limliiii \, 197S, (ailde Ici Aiisli.ili.m Shells Hoberl Brown and
Xssoei.iles Pl\, Port Moresin, P.\,(^. 77 pp. |l):ile Iroui
Wilson I 199:5: 18)|

K.ihIht. D, 1981. Card Cal.ilo^iie of Worl.l Shells P.iek 28,
lliihotis K:Lielier SI. Petersbnrii. Florida.

D. L. Ceisjer and B. Owen, 2001

Paee 83

Lindberg. D H, 1V)^)2, l"\(iliiti(>ii. (lislrilmtKui and s\stiiiiatus

of Hiiliotidiic. In. Sliepheicl. S. A., M. |, Tciriicr and S, A,

Guzman dfl Pn'xi (Eds.). Al)ali)nc uf tlir World: BinldirN.

Fisheries and ( 'iilture. Fislnii'^ News Hcioks. ( Jxlord. p. 3-

18.
Ostergaard, |. M. 1935. Recent and lossil marine Mollnsea of

Tongataliu. Bemiee P. Bishop Museum Bulletin 131: 3-

."iii, pi. I.
Owen, B., ,S, llana\an anil S. Hall. 2()((l, \ new speeies ol

abalone (Halioti.s) from Greece. The Winder 44: 301-309.
Philippi, R. A. 1S4T-18.51. Ahhildnngen nnd Besi'hreiliungen

neuer oder weniger hekannter ( ,'(]neli\lien. No!, .> rhee-

dor Fischer, Cassel, 138 pp.
l'ilslir\ II .\ 1890. Manual of Concholcig\ ; Structural and S\s-

tematic with Illustrations ol the Species 12: 1-323, (i5 pis.
Poppe, G. and Y. Goto. 1991. European Seashells. \'o\. 1. \cr-

lag Clirista Henunen, Wiesbaden. 352 pp.
Reeve, L, A. 1846a. Descriptions ol loit\ species ol IhiHcti^.

from the collection ot H. Guming, Esc|, Froceediims dl

the Zoological Societ\ of London 14: 53-59.
Reese, L. .\. 184(ib. .Monograph ol the (a'uus Udlinlis 22 pp ,

IT pis.
Sowerbv, G. B. II. 1882. Thesaurus ( .'onclixliorum | Fliesaurns

of Shells]. 5. parts 37 and 38:1 -.54. pis. 1-14.

Stalsolt, 1991. SIVIISIICA .Mac 4.1. StatSoft. Tulsa.

Oklahoma. S.j 1 pp.
Stewart, K \ and I). I., (iciger 1999. nesignation ol lectotspe

lor lldliolis crchrisciilptd Sowerln, 1914. with a discussion

ol // cliilliraUi Reeve. lS4(Siii())i I.ichteiistein, 1794). The

\elim-r 42: S5 9(1.
I'almadgi'. K H I9(i3. Insular haliolids in the western Pacfic.

The \eli.j,cr 5: 129 1.59. pi I 1
Ulialdi R I9S7 41ie llallotidae ol die Allanti( Medileiianeau

aiv.i. \r',;onaul;i .5: 2(iS-290.
rli.ildr R. 1993. Atlas of the living abalone shells of the world

I Fust deli\er\ with iutnnhicton' notes and four species

treated [. .Vssocia/.ioue .Malacologica Interuazionale. Rome.

1SS\ 1122-0.309. [ Irregular pa<.iination I
Wagnei", li | .md H 1" Abboll, 197S. Stand. ird Gatalo<; ol

SIh'IIs. Thud l-aliliou, American Malacologists, (Jreenxille.
WemkaufI, II, C. ISS.V l)u' ( ;attung y/r;/i<i//\. /;i: Svstematisch-

cs ( Jonch\hen-{ iabniet \on M.ulmi und (diemnit/, 2(fi)B.

p. 1-83. pis. l-:50.
Wells. F F, and W Bivcr 1985. Sea,shells of Western Anstra-

li,, Weslern Anstr.ili.m Museum, Perth. 207 pp.
Wilson. B. 1993. Anslrali.in .\larme Shells. \ol, 1. Od\ sscv

Publishing K.ill.inio HIS pp

Erratum

In the article li\ Wise (2001), please correct llie ratio lipl/lip2 lor Boiiucd ■.riiiiiiinlti iTahle 1, Ttli line). Tlie ratio
should he 1.5:1.0, not 51.5:1.0 as originalK stati'd,

LITERATURE CITED

Wise, J. 2001. Anatomv oi Boonca jadisi (Ols.son and MeOiiilv, 1958) (Heterobrauchia: Pvr.unidellidae' from the western .Ulantic,
with comparisons to other species in the genus. Ilie \aiitilns 1 15: 68-, 5.

THE NAUTILUS 1 15l3)r84-89, 2001

'age 84

AiistrodapluicUd ijcnienensis new species (Gastropoda: Tiinidae)
from Yemen, Red Sea, with notes on A. alcestis (MeKill, 1906)

.\ntonio Bonfilto
Bruno SabcUi

DipartiiiR'iitu di Biulugiu

E\oluzionistica e Speiimentale
Unixersita degli Studi di Bologna
\'ia Selmi, 3
40126 Bologna. ITALY
I ion fittofe'alnia. unit io.it
sali(-'lli{G'alnia.unil)o.it

Mauro Mora.s.si

\'ia dei Miisti. 17
25121 Bi-e.scia. rL\LV
vitinoni.s@tiii.it

.\B.STRACT

The genus AustrodaphniUa La.seron. 1VJ54, previously known
only from Australian waters, is lieie reported from the Arabian
Peninsula. Two speeies oeeur: A. i/ciiwncti.sis new species, from
Yemen, Southeni Red Sea, and A. alcestis (Melvill, 1906) new
eomhinatioii, a deep water species knowni onlv Irom the Gull
ol Oman. SEM photographs ol both spi'cies are provided.

Atldilii'iKil krij iionl\ liido-Pa(ifie. new eoiiiliinatiini

inae), previously inchuletl in the laniiK Tiirridae, is
transferred to the Conidae. However, this proposed clas-
sification has receiith' been criticized hv Rosenberg
(1999), who demonstrated that their results cannot be
ad('(|uatcK replicated. As a consecjueiice. this paper still
lollows tlie tratlitional arrangement. -Abbreviations used
in the text are: a/1 = ratio ol aperture length to total
shell length; b/1 = ratio of shell breadth to total length;
BMNH = The Natural Histon- Museum, London; NIZB
= Museo ili Zoologia dell" Universita di Bologna.

I\4H()DU(:T1()N

This p;iper deals with the genus AiislrndajthiwUu Lase-
nin, 1954. lornierlv reported onlv Iruin .Australian wa-
ters, but here shown also to occur oil the Arabian Pen-
insula. .\t least two species occur also in the Arabian
region, namely /\. i/ciiicni-nsis new speties Irnm Yeinen,
Southern Red Sea, anil A. alcestis (MeKill, 1906) new
combination, a deeper-water species from the Gulf of
Oman.

Aiistnitlaiiliiiclld i/ciiiciiciisis is piesi'ntlv known oiiK
Irom a lew adult spei-imens and fragments which were
dredged on tlie continental shelf of Yemen b\ the
French oceanograpliic ship .VIvhion i:)i'Fin;sM:. within
the frame of the European Conniiniiitv proji'ct RED
SED '92 (Southern Red Sea ;ui(l Ciilldl .\deii).

SEM micrographs of both spei ics here discussed and
a redescription of A. alcestis are given, .\lthongli the
anatomies ol its included species are unknown, the ge-
nus Austnxlaphiielhi is here assigned to the snblaniilv
Raphitoininae in accordance with previous workers. The
protocouch and teleocouch moqihologies of both species
here discussed are consistent with such alloiMtinn.

Under the cladistic classification of the siip<'iiamilv
Conoidea recentK j^roposed bv Tavlor, Kaiitor ;iiul Sv-
soev (1993), the siibfamilv najilmeiliiiae ( = i{;iphil(ini-

SYSTEM.VnCS

Laniilv ruiridae 11. and A. .Adams, 1853
Snblaniilv Rapliitominae Bellardi. L875
Ccnws Aiistni(lrij)liiiella Laseron, 1954

\iislni(l(ijiliiu-lhi Laseron, 1954, p, 45; tvpe species: Austro-
(Idjiliiirllii thithrtltd Laseron, 1954, hv original designa-
tion.

Diagnosi.s: Shell thin, ven' small to small lup to 10
iiiiii in lengthl, Insitorm, superficiallv resembling genus
niijitiiii 11(1. but totally lacking fine interstitial axial riblets.
Sculpture ol a\i;il ribs crossed bv few widely spaced spi-
ral ridges lorining (juadrangular interstices. Anal sinus
snbsutiiral and ven sliglit. Protocouch multispiral to
p;LiRispiral. diagonally cancellate or with rows ot .spirally
:illglied gr;ilinles.

Kangt": Red Sea, (iiill ol ( )in:iii and \iisl r;ili;iii waiters,
liiiin 9 to 25(-) III,

Remarks: l,;iseroii (1954), in his revision ol the New
Siinlli Wales tiiiiid hiima, (.'ii'dcd the gi'iins .\;/s-
lioildpluiclla Icir :i single speeies, iiamelv A clalliriilii
(hedged Irom nil I'ittwater. .According In lis deseriber.
Aiistrinliiiiltiielhi dlllers Iroiii the genus nuplnu lla
lliiids. IS I I III possessing a liisildrtn shell with :i dis-

A. Bonfittoc/ al.. 2(K)1

Wwc 85

(iiirtK pioduc-cil, instead ot (ixatc. hasi' ami a f(iais('l\
clathrate sciilptiirt' lacking fine interstitial a\ial rihii'ts,
Powell (1966: 124) ieiz;ariled AiistriHldpliiirlla as a valid
genns inelnded within the Daphnellinae ( = Kapliitoin-
inae), a)! opinion reeent]\' tollowx^d hv S\soe\ il99.'Vl.

Shuto (1983) described a second specie's, naniek .\;(s-
trdilaplniclla tonvscnsis. haseil on tew daniaij;ed shells
dred<j;ed fr(nn off Mnrra\" Islanil. Torres Strait, (,)neens-
land. Shiilc} ii"j;arded thi' species as distinetixe liecanse
(il its peiipheial an^nlatioii and connnenled lop. cit,:
24): "it is a [irolileni whethei' it is inelndeil m Aiisli'a-
(laphticlhl Laseron, 1954 or it represents a new snl)<j,eniis
or genns". We ha\'e not examined tlu' t\pe seiies oJ.V,
torrcsensis. lint its morphological leatines seem not dis-
tinctive enongh to warrant a snpraspecilie distinition
Irom A cldthrata.

In this paper, we regard tiie taxon Aiistrodapliiulld as
a \alitl genus on the basis of its teleoconch scniptnre.
whicli (.liffers considerabK' from that of Ddplinclld
Hinds, 1844, this latter being a wideK ilistribnted gemrs
well represented in tlie tropics.

Species assigned to DaphncUa bear dense and minute
axial riblets crossed bv spiral lirae. In some species the
.spiral lirae are numerous and nearl\- ei[uallv thin as the
riblets forming a \er\ fine reticulation (e.g., DapUiicUii
bdtimica lledlev', 1918), but lirae mav also be much few-
er and stiduger than axial elements (e.g., Ddpliiiillii siih-
uhi (Heeve, 1845)), rendering a distinctk caiinatt' aspect
to the teleoconch whorls (e.g.. DapJinclla Mihriiui Mel-
x'ill, 1906). Some other Diiplutclla species have distinct
axial ribs on earl\- teleoconch whorls, but this ornami'u-
tation becomes completely obsolete on the last whorl,
where the axial sculpture is represented onK bv rather
thin riblets that produce, at most, a slight beading at the
intersections with the spiral lirae.

The genus Austnidaplutrlld has witIeK spaced axial
ribs crossed b\ few spii'al ritlges, foiniing relativeK'
broad, (juadrangulai' interstices. The axial sculpture is
well developed on all teleoconch whorls, anil tlie mimite
riblets observed in Ddpliiulld are totally lacking. In ad-
tlition to tlie difference in sculpture, specii-s assigned to
Aii.stnxIdpliiK lid have a well-produced and stronglv ex-
cavated base rendering, the last whorl conspiciiosK more
inflated than tlie penultimate one. Tliis feature, in ad-
dition to the thin s\\v\\. the relatixelv few teleoconch
whorls, and the ven' shallow anal sinus, ma\' give the
shell a somewhat jmenile appearance. This presumably
led Laseron (1954). in his description o{' Austnidapliiicl-
Id cldthrdfa, to olisene that "the t\pe max not be ijiiite
mature". Mi'Kill (1906: 78). probabK due to the small
size antl the general appearance ot the shell, also con-
sidered the described specimen of his Daphiwlld iPlcii-
rotomelld) dicestis. a .species here assigm'd to Anslro-
(hiplinclla. as probabK' immatiu-e. Howevfr. relerring to
other specimens of the species, he noted that "the six or
eight examples that occurred were all much of the same
size". The somewhat immature appearance notetl for the
two above-mentioned species is also present in A. ijc-
iiinifiisis. .'Mtlioncrh determination of the adult state max

be somewhat piobleiiialic in spi'cies lacking a terminal
\arix or swelling, it seems unlikcK that all these species
are actiialK based solelv on juvenile specimens.

The genus Ddplinclld differs distinctk in tliis respect
h'om Anslroildphmlld in p(.)ssessing a less produced and
iisuallv more shallow k excavated base, which gives the
last whorl a more elongate-ovate sliapi'. Furthermore, in
iiianv Dapliiulld species tlu' anal sinus, though not nec-
essarik deep, is nevcillieless distinct; ihi' species as-
signed to Auslroildpliiiclld. as stated above, have a nearly
imperceptible sinus. Whether tlu'se differences between
the txvo taxa are of generic or snbgeneric rank is a matter
of opinion. However, the genus Ddplnulld. as presently
constnieil. seems to represent an ovfrcrowded and het-
erogeneous assessment of species ranging from the Eo-
cene to Recent. Fossibk. anatomical studies on some of
these specii's might produce I'videnee allowing separa-
tion of Ddpliiulld into different genera or subgenera.
Kxamples of similar situations in other tiirrid groups are
taxa such as Cmssispird Swainson. 1840. and Encitlidi'd
Fischer, 1SS3, belonging ri'spectivek to the subtamiiies
Crassfspirinae and the Mangeliinae. Both these two gen-
era are currently accepted as composed bv several dif-
ferent groups recognized on the basis of the shell mor-
phology antl/or radular features (see for example Mc-
Lean. 1971; Kilburn, 1992). The tiixon Aiistroddpliiiclla
seems to represent a snuill but distinctive group of spe-
cies possessing simikii' shell projiortions and sculpture,
and is probabk wortliv ol recognition as a genus separate
from Ddplinclld. even if its ;inatoiiiical features are pres-
ently unknown.

.\part from .-\. ilddiidtd Lasi'ion. 1954 and .A. tonv-
scnsis Shuto. 1983, both from .\ustralian waters, there
are at least hvo other speiies lidm the .Arabian Peninsula
cleark' referable to Aiistraddplinclld. One is hereunder
described as Anslroddpliuclld i/cincncnsis new species,
the other. .A diccslis iMekill. 1906) new combination,
was dredged from off the (nilf of Oman and has never
been recorded sinci".

Ansfroddplii\cll(i iicnicncnsis new species
(Figures 1-7)

Dia^no.sis: Shell np to 8.(i mm in length, last whorl
with 21-25 axial nlis decnssiiteil In 4 main spiral ridges.
Subsntural ramp and interstices bc^twecMi ridges with
secondan sciiljiture. Protoconcii pancispiral. with gran-
iilai" spiral threads.

Description: Shell v\illi a slightk cyrtoconoid spire,
teleoconch ixinsisting of 4.2 raimllv I'xpanding whorls
with strongk impressed sntnie. Whorls profile weakly
shouldered. Subsntural ramp narrow and weaklv con-
cave, sculptured bv 6-7 ion last txvo whorls) verx" low
spiral threads and vcrv fine axial incremental lines. Ap-
erture oblaiK-eolate. gradualk tapering to siphonal canal.
Siphonal c;inal moderatek wide, lacking a terminal
notch. Outer lip rather thin, not preceded by a labial
varix and smooth inside. Columella rather long and gent-

Patre Sfi

THE NAUTILUS. \ol. 115. No. .3

A. Bonfitto ct <ii. 2001

Pas£e HI

\\ c()ii\f\. l^arii'tal region slii^litK i(iii\i.'\. l.ahi.il callus a
tliin glaze. SEM exaiiiinatiun shows surtaei- ol inner lip
coxered with niicroscopie, sparse, somewhat s(jnaniifnini
granules (figure 5). Anal sinus rexerscd, I, -shaped,
Stromhoid notch ab.seut. Sculpture consisting ol narrow,
slightlv angular avial ribs crossed bv widek' spaced spiral
ridges to form spiralK elongate interstices. .Axial ribs or-
thocliue. uearK ecjual in width to intenals or about hall
their width, obsolete on subsutural ramp, lading on last
whorl at lexcl ol upper part of columella. There arc IS
axial ribs on pemiltimate whorl, increasing to 21-2.3 on
last whorl. First teleocouch whorl with 3 spii'al liilges,
subseijuent whorl with 3— i m;iin spiral ritlges. The fii'st
ridge, at the lower edge ot the subsutural ramp, is dou-
ble-spaced from the remaining ones. t)n later two teleo-
couch whorls, an additional weak ridge is obseiAcd in
the interstice between tlie first and second ridge, hiter-
stices betx\een spiral ridges with 3-5 (on last hso whoiis)
\er\ h)w spiral threads. Interstices between spiral
threads with a microscopic sculpture oi spiralK aligiR'tl
granules (fisiure 7). Base with 14—15 ridi'es. those on
rostnun long and nearK' vertical; interstices between
ridges with 1-2 \t'r\ low threads on posterior part ol
base, anterior part ilexoid of secondan' elements. Back-
grounil color from light \ell(.)w to tan. with irregular oi-
ange blotches on last two whorls. Protoeonch pa[)illiloi"ni
consisting of l.S whorls with impressed sutiu'c. scnlp-
tiu'ed b\' miimte spiial threads crossed bv finer axial
threads to lorm rows of spiralK' aligned, minute grannies
(approximatcK l(i on last whorl ot the holotxpei. l^rn-
tocouch diauu'ter 0,41-0,43 nun.

Measurements (in mm): Holot\pe: Ijcngth S,(i.
width 3.3, aperture 4.5, b/1 0.38. ;i/l' 0.52; parat\]H' 1;
length 8.5. width 3.7, aperture 4.rs, b/1 0.43. ad 0.'54.

T\pe material: Holotxpe, MZB 17001: l'ar;it\pe 1,
MZB 17002; Paratvpes 2-3 MZB 17003. I\irat\pes 2-3
are fragments lacking tin- bod\ whorl.

Type locality: Oil Ycnu'u. southern lied Sea.
14°46'72" N, 42°32'82" E. 7t-i m depth on iiincid\ s;uid.

Etymolog)': The spi'cific name refers to the coun(r\
where the uew species was dredgetl.

Remarks: This new species is assignetl to genus .A//.v-
trodditlnifUd l^aserou. 1954, on the basis of its teleo-
couch features. AiistrodaphncUa i/einciifiisis is readil\
separable from its congeners bv its short, papillilorm
protocoucli with lows of spiralK' aligned granules instead
of conical and with the tvpical diagonally cancellate
sculpture. The protoeonch moipholog) of the new spe-

cies iua\ resemble ih.il ol die wideK distributetl hulo-
Pacific genus Emijclotoiiui Boettger, 1S95. iIowe\er,
membeis ol (he hitter geims differ from species of Aus-
liddtijihiK llii ill possessing sti'ong spiral keels on the te-
K'ocoiich whoils, i'lirthermore. species assigned to Eii-
cilcloldiiKi \\:i\f A truuc;ited anterior end, a distinct
thongli not exeei'dingK deep anal sinus, and dense axial
interstitial riblets closcK resembling in this character
species assigned lo genus Diiphiiilhi.

Apart Iroiii (lie proloconch. A. ijcmencnsls differs
Irom .A cldllinilii K.iseron. 1954. which is of Comparable
size being oiiK sligthK larger (10 X 4.5 mm according
to Laseron. 1954' ;ind has similar general shell propor-
tions, in IkinIii'j, ihree rather than two spiral ritlges on
carK whorls and. judging from the original figure (op.
cit.; 45. fig. 2281. less beaded spiral ridges.

AustrtnlaphuiUd (ilcfslh i\h']\ill. lOOfii new

couib'uation

! Figures S-10

Dajiliiiclhi rliiirohiinilla ahr^lis MeKill, 19(l(i: 7S. pi. S. (i<i.
2:5.

I'lcinoloiiirlla ,il,;->tis MrKill. 1!)17: 19(r

Diagnosis: Shell ol 5.4 mm in length, last whorl with
27 axial ribs decussated In 5 spiral ridges. Subsutural
ramp witli arched wrinkles, interstices between spiral
ridges lacking secondan sculpture. Protoeonch niiiltis-
piral, diagonalK' cancellate.

Description (BMNH specimen): Shell with sliglitK
cxrtocouoid spire, teleocouch consisting ot about .3 rap-
iilK expanding whorls with impressed suture. WTiorls
conxex. weakK shouldered. Subsutural ramp narrow and
weakK' concaxe. .\pertme obhiuceolate gradnalK taper-
ing to siphonal canal. Si|)lional canal moderateK wiile,
lacking terminal notch. ( )iiler lip \ei\ thin, not preceded
In labial xarix and smooth within. Columella rather long
and exeuK convex. Parietal region convex. Labial callus
;i thin glaze. .\nal sinus \er\ sIkiHow and broadlv U-
shaped. Stromboid noti-ii ;ibsent. Sculpture consisting of
narrow. slightK angular axial ribs crossed by wide-set.
spiral ridges to form disti]ictK (|nadrangular interstices.
.\xiaK oithocline nincli narrower than intenals betxvei'u
them, forming stronuK arched wrinkles on subsutural
nimp. evaucsciug on last whorl at the adapical part ol
columella. There are 17 axial rii)S on ])enultimale whorl;
this iinmlier increases to 27 on last wiiorl. First teleo-
concli whorl with 4 spiral ridties, the first and weaker
just below subsutural r;mip. Penultimate whorl with 4

Figures 1-10. Species oi Amirodaphndla. 1-7. Ilolotvpe of Ausli-odiiphiwHa i/eiiu-nnisis new .species. MZB I7()()l. 1. .Apertural
view. Scale liar = 1 mm. 2-3. Protoeonch; scale bar Kli) (jim. 4. .\pical \iew. Scale l)ar = 1 mm. .5. S(|iiamilbnn granules ol the
inner lip. Scale liar = 10 jjLm. 6. Teleoconcli vvhod. Scale Itar = .5()() (J.m. 7. Microsciilpliire of leleoeoiich. Scale Ijar = 50 ^ni.
S-10. Lectotvpe of Aiistrodaplinclla dlccstis (.\Iel\ill, I9()fit new c<>ml)iii;ition. 8. l,ectot\pe o\' Ihiphnclld i Piciirolomi'lla i alcvstK
MeKill. I9(I(S'. H\1MI lV)(l(-i.ll).2:3.2.S-9. Scale bar = I inin 9. Ti'lcocoiicli. Scale bar = KK) (Jiiii 10. Protocoucli. Scale bar = 100
ixm.

Paee 88

THE NAUTILUS. Xol. 115, Xc

ridges and a fine thread at ahapiial suture. Last whorl
with 5 spiral ridges. Base of last whorl with about 16
ridges (those on rostrum nearK laded i. Color dull white.
Protoconeh eonieal, O.fiT uun diaiueter, with more than
3 whorls (tip missing) and diagonalK' eaneellate sculp-
turi'.

Measurements (in mm): Length 5.4, width 2.3. ap-
erture 3.0, h/1 0.43. :i/l 0.5fi.

T>pe locality: ( ;nir nC ( )man. 24°58' N. 56°54' E, 2S5

III.

Material examined: Two spceimeiis l.ilielled as s\'n-
t\pes stored in BMXH (reg. no. 19()fi.l0.23.2S-9). Of
these, one agrees with the original deseription. the other
is very different, possibly a specimen ol Cliillmrclln (iin-
pliil)lestnim Melvill, 1904. The specimen ot Daphiicllri
I Piviirotoiiu'lla} alci'stis is here selected as lectot\pe. Tlie
SEM micrographs were taki-n with the specimen un-
coated.

Remarks: This species was origiualK mtnjduced as
Dii])hiH']l(i (Pk'urotomi'IIa) alccstis MeKill 1906, and
compared with Dapliiiclla Uicdsi MeKill, 1904. and Cla-
thurella (iinpltihlcstniiii MeKill, 1904, both from the
Gulf of Oman. Howexer, iieithei' of these taxa is actually
closely related to the species here discussed, which
seems properly assigned to the genus AustrodaphncUa.
AustrodaphiteUa alccstis is readiK distinguishable fnjm
A. ijcmcncnsis h\ its t\pical diagoualK' eaneellate proto-
coneh .sculpture instead ol rows ol spiralK' aligned gran-
ules. Furthermore, Aiistnxlaplinclla alccstis (MeKill,
1906) is much smaller (5.4 mm \s. 8.6 nnn in length),
and lacks the secondan' sculpture ol spiral threads in
the interstices behveen main ridges. In addition, A. al-
ccstis has a dull white shell lacking the orange blotches
obser\ed in all the a\ailable specimens of A. i/cinciicnsis.
It is comparable with A. lorrcscjisis Shuto, 1983 in di-
mensions but differs distincth Irom the latter in its con-
vex whorls lacking the peripheral angulation nt the .Aus-
tralian species. AiistnulapluicUa clalknila is iiiiicli larger
than A. alccstis (10 X 4.5 mm \s 5.4 X 2.3 mm) and
has two instead of lour sj)iral ritlges ou early teleoeoneh
whorls.

Biogeographic remarks: 4'lic picsciicc ol a genus
origiualK' described Irom the Southern Hemisphere in
the Arabian Region ma\' probabK' be explained consid-
ering the effect of the .\rabian Sea ujiwclling. During
summer, the coastlines of Yeiiun and Oman are affected
bv the strong Southwest Monsoon, whirli blows warm
surface water offshore causing its replai-ement In deep,
colder oceanic water. Sheppard et al. (1992) discussed
some of the more significant consecjuences dial deiixc
from this event. Among (jtiier effects, strong iipwclliug
in tropical regions inhibits coral reef growth and induces
the de\el(pnient of macroalgal conununities of a tem-
perate rather than tropical natm-e. 'I'lms, the oi'cnrrenc("
in tlu' Arabian Sea of species oi' Eckloiiia. a kel|) genus
otlu'iwise found onK- in the southern li<'iiiispherc i, Aus-

tralia, New' Zealand and South Africa), is recognized as
probabK' resulting Irom the lower water temperatures
and nntrieut input associated with upwelling. The au-
thors also suggested the possibilitx' that the cold up-
welling water, bv inhibiting reef growth, ma\' act as an
ecological barrier restricting the recruitment ol main
tropical groups in the Red Sea and the Persian Cinll. It
seems possible that larvae of species oi AustrodaphncUa
ma\ liavf fountl in the .Arabian Peninsula en\ironmental
conditions faxorable to metamoiphosis. Sheppard et al.
(1992) also noted that the upwelling effects mav al,so be
strong along the SomaU coastline. A possible example
supporting this \iew is the occurrence off Mogailiscio of
Psciidcxoiniltis fiiscoapicatus Morassi, 1997. a species
belonging to a genus known othenvise ouK' Irom South
Africa, soutliern, and eastern Australia.

AustrodaphncUa i/cnunicnsis differs Irom all others
s]ieeies assigned to the genus AustrodaphncUa in pro-
toconeh moipholog\' (paucispiral instead ol multispiral).
This difference is generalK' regarded as indicative ol \^\■o
different t\pes of de\'elopmental strategies, i. e.. uon-
planktotrophic versus planktotrophic dexelopment.
However, it is generally agreed that different txpes of
lanal dispersal ma\' develop in species of the same ge-
nus, so that a species with a paucispiral protoconeh does
not necessitate inclusion in a genus separate Irom that
including species with multispiral protoconchs ( Bouchet,
1990).

A number ol rii\ ii'oniiK'ntal tactors associati'd both to
geographic and climatic iactors (such for example geo-
graphic isolations, temperature changes, eustatic, and sa-
linitx' changes), ma\' act in promoting loss of plauktotro-
ph\'. Planktotrophic species have a prolonged lanal life
and aie therefori- more exposed to pinsical and biolog-
ical factors increasing lanal mortalitx rates (Ruuuill.
1990), Ac(|uisitiou of lecithotrophic or "direct" lanal de-
\elopmeiit nia\ therefore be an ecological adaptation fa-
vorabK selected in ri'lati\el\ hostile environments char-
acterized In climatic fluctuations (O!i\erio. 1996). The
diaiiiatic cliiiiatie changes in the .\rabian Peninsula that
particnlai'K .iflected the Red Sea and the Onlf of Advn
iluriug the Ouaternan. and the continued strong season-
al tiMiiperature Ihictuations related to npwi'lliug condi-
tions are factors that probabK induced, and nia\ still be
inducing, uoii-planktotroplnc lanal (k'xclopment.

.\(:k\()\\ i,i;i)(;mk.\ IS

\Ve wish to th.uik Dr Marco Ta\iani (C.N.R. — Italian
National Hesearcli (^oinicil^ Italian j'jartner ol European
{'ouunnnitv project RED SEP) '92 and Ms Katliic \\a\
(BMNH) for the loan of t)pe .specimens.

LITERATURE CITED

lioiiclii'l, I' !!)!)() Innid ij;ciu'ra and iikkIc of dcsclopiuciit:
llic use ami .iliiisc ol pi'oloconcli iiiorpliolog\. Mahu'ologia
:V2: (W-77.

Kllhiini, W \ l(Ji)2 ■I'linidac I .Molliisci: Casli'opoda' of south-

A. Bonfittd li <il. 2001

Paue S9

em Alricii ;ind MozaTiiliicjiu'. I'art (i, SiihrainiK Maii'^rli-
inae, section 1. Annals ot tlie Natal Mnseinii 33: 4(i 1-575.

Laseron, C. F. 1954. Re\ision of the New Soiitli Wales Tiiiri-
ilae. Avistralian Zoologieal Haiulhook, Renal Zooldniial
SocietA' of the New South Wales, S\(lne\, .56 pp.

McLean, J. H. 11-)71. FaniiK Tin-ridae. /// .\. M. keen. Sea-
shells of Tropical West .Anieriea. 2"' edition. Stanford Uni-
\ersit\' Press, Stanford, pp. (iS(i-71i(i.

MeKill, |. C. 1906. Description ol tliirt\-(iue a;astropc)d,i and
one scaphopod from the Persian (nillaiul (inlf'of ( )nian.
dredt;ed In' .\Ir. F. W. Towiisend, 1902-1904. ProceediiHj;s
of the Malaeological Societ\ of fjindon 7. 69-.S0. pis. 7-.S.

MeKill. |, (' U)17. Revision ol the Turridae (Plenniloniidae '
occurnni; m the Persian (Jiill. (nilf of Oman and the
North Arabian Sea. as e\ndeneril mostK- through the re-
sults of dredgings carried (jiit li\ \lr. F. W. Townscnd.
1893-1914. Proceedincrs ol th.' Malaeologieal Soeiet\ ..I
London, 12: 14()-2(>1. pis S-IO.

Oliverio, M. 1996. Contrastinu; tle\el()piiieiital strategies and

speeialion in N. K. .\tlantic [)rosol)raiichs: a preliininan
analysis. /;/ |. 1). Ta\lor (ed.) ()ri<;in and evolutionan' ra-
diation ol die Molliisea. Oxford Uni\<-rsitv Press, London,
pp. 261-266.

I'owell. .\. W. B. 1966, Hie iiiollnsean lainilies Speiilhliidaeand
Turridae. Rnlletin of the .\iicklaii(l liis(ilnl<' and .Miiseniii
5: I-1S4. pis. 1-23.

liiiscnlieig, C l<)09. Reproducibilit\' of results in plnlogenetic
an:il\sis ol inollusks: a reanalvsis of the Ta\ lor, Kantor, and
S\soe\ I 1993) data set (or conoidean gastropods. .Xnieri-
can .Malaeological Bulletin 14 (199St: 2I9-22S.

liiniirill, S, S. 1990. Natural mortalitv of marine iincrtehrale
lan.ir. Ophelia 32: 163-198.

SluppaKJ ( \. Price and C. Roberts. 1992. Marine ecolog\
ol the .Arabian Region. I'atterns and processes in the ex-
treme tropii-al emiromnents. .Academic Press, l^)iidon.
359 p[i.

Slinto, T 1983. Ne\\ liirnd la\a Iroiii the Anslralian waters.
.Memoirs ol die Facultv ol Science. L'nivcrsitv ol KmisIiii.
series 1.), ( ieoloiIX' 25: 1-26.

THE NAUTILUS 1 15(3):9()-9.S, 2001

I'aiii' 90

A cladistic aiuiKsis ot species of Lamhis (Gastropoda:
Stronibidae)

|. H. Slone

Dcpartnieiit of Animal Rcologv'
Exolutionan' Biologx' Centre
Uppsala Uni\ersih'
NorbvAiigen ISD
SE-7.52 36 Uppsala
SWEDEN
jon.stone@e\iiliiti()ii.iiii,sc

ABSTRACT

A cladistic analysis of the 9 species ciirrentlv classified as the
genus Lainhis (Abbott, 1961) was conducted. Three species in
the genus Strombus (Abbott, 1960) were used as outgroup.s.
Features of the radula, soft anatomx', and shell were coded as
binarv' character states (i.e., in a iiiaiiiK r that allottctl one char-
acter state for everv bit ol inlorination). Each ol two most-
parsimonious cladograms obtained contains a clade that in-
cluded all 9 species ol Liniihis. one including a single species
ol StvomlniH and the other including two. On tlie basis ol this
cladistic anaKsis, Laiuhis is paraplnlctic and Stroinhii.s is poK-
phvletic. Therelore, the traditional classifications ol species
within these genera are untenable. The results obtained are
used to reassess hypotheses ol moi-phological evolution and
adapti\e function ol circuni-apirlnral projections ol strombid
ga.stropods.

AililUidiuil key ui>ril\ .Adaptixc Inuclidii. gastropod, inlorma-
tiori. moqjhological cMilntinu. ph\loi;i'Uctic swsteuiatic auaKsis.

INTHODUCTION

Ijiiiiliis i.\l)l)(itt. 19fil) is a gcinis (il iiuiniu' gastropods
(•ii(!cmic to the Iiulo-Pacific region and a nienihcr ol the
lanilK' Stroniiiidac Iialincs(|n(', 1<S15, which also inchides
the genns Stroiuhiis Linna<'us, 175S. .\i)l)ott (lOfil) di-
\iclcd the 9 species cnrrentiv comprising the genus Lain-
Ins into three snbgenera: I,, hiinhis (Liinie. I7.5S), L.
rrocatn (Link, 1S07). and /., Ininnilti ( I Inmphrex. 17S6)
comprise tlie subgenus Laiuhis s.s. liiiding, 179S; L. iiiil-
Icpcda (Linne. 1758), L. di^itata (Penv, 1811), L. ro-
htistn (Swainson, 1821). L. .scoiiiins (Linne, 1758), and
L. vAolucca (Swainson, 1821 ) comprise tlie subgenus .\///-
lepes Morch. 1852; and /.. chira<irii (Liime, 1758) is llic
lone member of the subgenus Uiirpci^^o Morch, 1852.

The most outstanding moiphologieai fealinc of spe-
cies o( iMmhis is the set of "cirenm-aperlnral projec-
tions" emanating from the edge of the dared apertnral
lip ol the shell. The number of circuni-aperliiral projec-
tions iCxeluding the si]iii<inal canal) varies among spe-

cies, ranging Irom 5 (e.g., L. chira^ni'^ to 11 (e.g., L.
violacca) excluding 7. Circum-apertural projections have
been ,showTi to confer protection against crushing (Palm-
er, 1979), postulated to have evolved as a response to
predation (Vermeij, 1989), and Inpothesized to provide
staliilitv on substrates (via "snowshoeing," or weight dis-
tribution) and a means of circumventing geometric con-
straints chniug ontogenv (Savazzi, 1991). Cireum-aper-
tiu'al projections also might enhance stabilitv via hvclro-
dvnamie jirocesses (i.e., drag forces might counteract lift
forces prochncd bv currents or waves). .Anv eonibination
of these factors might be responsible, through evolution,
for tlu' existence ol circum-apertiu'al proji'ctions. How-
ever, wilhoul ic'lerence to a rigorous cladistic analvsis
from which a hvpothesis of the evolution ol specii'S ol
Lainhis can be fornndated, explanations ol the origin and
utilitv ol circuni-apcrtnial projections picdomin.mllv tU'e
speculative.

Ill this pajiei", a cladistic analvsis ol speiies ol Ijiiiihi.s
is presented. The results obtained were used In reassess
hvpotlieses ol iiioiphologieal evolution and adaptive
liiiK'tion ol ciiciim-apertnral projection.s.

M.\TEH1\I,S \Nn METHODS

.Sample nialerial: Inlormatioii was obi, lined lioiii ex-
amination (I'.g., scanning electron microscopv. dissi'c-
tion. obsenationl ol specimens (table 1) and literature
(,\libiill, 19(i(l: I9(S1 1 concerning all 9 spei ies in (he ge-
nus Ldinhis and three in Stroinhiis i.S hiilla iRoding,
1798), .S. (lilatatus Swainson, 1821, and ,S', mnicr (Riid-
iiig, 1798)).

■Slatc'inenl ol' thai aeters and defiiiilioiis of eliarac-
Icr .states: Inlorination was coded into eliaracti'i' states
representing fi3 characters: 19 concerning the radula, 7
coiiceniing the soil aiialoinv. and ■■>7 (.onci'i'iiiiig llie slicll
isee Ix'liiw and apjiendixl.

|. H. Stoiif. 2001

I'-.K'v yi

Tahlo 1. Species ot the hiinil\ Stniiiiliulae cxaiiiincil niiiseiiin eatal()ij;iie uleiillficalioii, rdllcelioii kicalides ami sample sizes (n).
As a conse()ueiK-e ot rarih" ot sciiiie nl llir speeies iii\c)lve<l, only subsaiiiples ol speeiineiis were liaiulled (iireetiv to evtruet iiif'or-
matioii tor some characters (e.g., to olilani iiironiiatioii concerning the radnla oi' Lriiiihis lainhis. on!\ a sin>jle sjiecinien of the hvo
comprising AMSG C.'5063T3 were snhjected to scaniiinii electron niicrograpliN ': other specimens and Mierature were examined to
obtain information lor other characters.

r.r

Stn>iul)us ilil(it(ilii\ .Swainson 1S21
Stroinhiis hnllii (Roding. 179S)
Sintinhus voiiicr (Rikling, 1798)
Ijiiiihis liiiiihis (Linne, 17.5'S)

Liitnhis Iniiiriitii ( nniiiplirex. ]7S(-i)

Liiinhis cnirdlii (Link. 1S()7'
Ijiiiihis iiiillrprdd (I.inue. 175S)

Lambis ili'^itiilti (Pern. 1811)
Liniihis ri>liiista (Swainson, 182
Ldinhis si-iirj)in^ (Lnint', 17.58)

l.<iiiil)i\ iliini'j^ni (Linne, 1758)

Depositon nunilu'i

AMSC
AMSC
AMSC
AMSC

ANSJ

ROM

AMSC

ANSP

ANSP

ROM

A.\SP

ROM

AMSC

ANSP

IK )M

IIOM

.\\IS(

AMSC

ANSP

ANSP

AMSC

AMSC

ANSP

ANSP

ANSP

ROM

; (:.i()f-i.377

; C:3()fi41)2
; C:3()fi398
; C306373
246931
Ace 1990-039
; C30fi4()(i
237062
'21.5318
Ace 1989-051
'212819
Ace 1987-043
; 0306371
' 22S924
.\(c 199-0.39
1995-001

; (;.506:)75

; (:,306374
206,391

39879

306374
; 0306372
212429
' 201470
223975
Ace 1987-0.55

( olUttioii loe.ilit\

.\nstralia

l'liilip[)ines

(.'oral Sea

New (Jninea

Philippines

NLI

Ooral Sea

New Caledonia

Marianas

Oman Mnscat

Zanzibar

Philililiines

Philippines

I'hilippines

NLI

Tanzania

Societs Islands

New Guinea

New (iuinea

Zanzibar

New Cluinea

.■\nslralia

Zanzibar

Palaii Lslaiids

Philippines

iiulo-l'aeihe

3

1

2

2

2*

2

1
2

1^
1

2A

4

1

8^

1

1

1

1

1

V

1

1

3

1'^

1*

1

A\-\ specimens (i.e. shell onl\ ).

photographic plate of specimen inchideil Ml \bbott lOISOor \liliott 1961.

\\.\nv\.\

Strombid species possess a taenoglossate radiila consist-
ing ot rows of se\'en teeth (figure 1): a central (median,
nr lacliidian) tootll Hanked on eitlu r side In a single
lateral tiiotli (uid Iavii marginal teetli (or imcini).

Characters 1-4: Tijpc of fimtli cusps. Absence (0) or
presence ( 1 ) of 4 t\pes of tooth cusps were defined and
coded as hiuarx character states: obtuse-tipped, apicu-
late-tipped, inilk'peiloid," and "cliiragroid' (figures 2-5).

Character.s 5-6: Xiiinhcr of cusps fliiiikiiiii cciitial
teeth- C^entral teeth possess a single medial cusp flank(^d
on either side li\ smaller cusps in a series (figures fi-7).
Absence (Oi or presence (1) of 2 and .3 cusps in the series
were defineil and coded as binar\ character states.

Characters 7-10: Number of cusps of lateral
teeth. Most strombid species possess Literal teeth with
a single large cusp adjacent to the central tooth and
smaller cusps in a series that extends towari! the inner
marginal teeth, .Vbsence (0) or presence (1) of 2, .3, and
4 cusps in the seiies were defined and coded as binar\
character states. One species, ,S. dilatiiius. li;is latei'al
teeth with 4 eijuixalent cusps; this condition also was
coded as a binan charac'ter state.

Characters 11-14: Suuibcr of cusps of inner uian^iu-
al teeth. ,-\bsence M)l or presence (1) ot specific num-
bers of cusps ot inner marginal teeth were defined and
coded as binaa character states: 4, o. fi, and 7,

Characters 15-19: \ umber of cusps of outer man^in-
al teeth, .\bsenei' lOi or presence (1) of specific nimi-
bers of cusps of outer marginal teeth were defined and
coded as binai"v charac'ter states: 4, 5. 6. 7. and 8,

Soft i^M^TS

Characters 20-21: I'/pe of mouth. Some strombid
species burrow into substrata, using their acrcmbolic
snouts to displace sediment (Sa\a/./.i, 199r), .\bsence (0)
or presence (1) ol a stromboid' and an elongate mouth
I figures 8-9) were defined and coded as l)inai"\ ciiaracter
states.

Characters 22-20: Sumber of .serrations of the oper-
culum. Strombid species possess a sii'kle-slia|ied oper-
culum that tills the aperture incompleteK. .\s most
strombids do, s]iecies oi l.ambis use the operculum as a
lexer during xaulting,' or leaping locomotion (Sa\';izzi,
1991). OccasionalK, such behaxior results in oxerturniug:
righting is accomplished 1)\- lifting the operculum oxer

Page 92

TIIK XAl'TILUS, \nl. 115, No. 3

Figure 1. Scanning electnm iiikroiirapli ul tarnoglossate
radula hpical of species of Strmnliidac [Ldiiihi^ iitillcpnhi: bar
= 150 |xm,).

llic Icll side (il tlic aninial. lowering the operciiliiin into
llic siilistratiiiii, and kicking lateralh' (Savazzi. 1991). A
siniilai' op<iculai motion has been ohseived as a defense
nicilianisin against fish and crah predators (Jnng and
Al)l)ott. 19fi7: Xh'Kin, 1973: Sa\az/i. 1991: Wrnieij.
pels, coinni.). Tlie operculiun is suiootii (i.e. has 0 ser-
rations) or (lenticulated with 7, <S, 10, or 16 serrations.
Absence (()) or presence (1) of specific nninliers of ser-
rations ol llic opcicnlinn were coded .is liinaiA character
states.

,Sni:i,i,

C;haracters 27-29: Number of ivhoiis. .'\lisence (0)

or presence ( 1 ) of 9. 10, and 11 whorls were defined and
I'odeil as liinan cliaracter states.

Character 30: Form oj protocoiich. The protoi'omh
of tlie shell of some strombid species is scnlptnred. Pos-
session of an nnscnlptured (01 or a scnlptnred ll) pro-
toconch were defined ami coded as distinct character
states.

Characlers 31-35: lijpr of sndpiurv (i/ aliorls. The
whoiis of the shell ol strombid species exliibit a variety-
of scnlptnics. Absence (0) or presence (1) of 5 t\pes ol
sculptuie ol whorls were defined and coded as binan'
character states: cords, lines, ribs (longitudinal, narrow
ridges; Arnold, 196.5), threads (slender surface eleva-
tions: Arnold, 1965), and beads.

Characters 36-38: Tiipe of sculpture of eoluniel-
1(1- The columella ol strombid species exhibits a \ariet\

Figures 2-5. 't;-p s ul > u,]/, ui i.ululai li'etli of some sp<'eies of SlKiinlildae ieliaraeteis l-Si. 2. .\pK iil.ile lippi'd [Slrnmhiis
haemitstoma SoweHn-. 1.S42; bar = .50 |xni): 3. Ohtnsc-fipped il.iiinhis tniinalir. bar = 6.50 jxin); 4. 'Millepeiloid i/, iiiillriirila:
bar = 1.50 (jliiiI: .5. '(^liirasiioid' i/,, elnr/i<inr. bar = 150 p.nil. Dirierences ariKiiii; t\]ies were deterniiTied on llie basis ol lonii.

H. Stone, 2001

Page 93

Figures fi. 7. \iiiiilicr nl t-iisps llaiikiii'j; iciiliai tirlli i>l
some species of Stromlndae (eliaracters 9-1 i). 6. Central tooth
of a specimen ol Striiiiihiis lariahilis Swainson, 1820 with 2 or
3 flankinij ensps (bar = 25 [xm). 7. Central tooth of Slrniiiliiis
inar^inatii.s Linne. 1758 with 2 or 3 Haiikiiiij; eusjis lone ol llir
tiiree ensps is iiulieated with a wiiite wcih^e; har = 75 |j.in).

of

sculptiirt's. Absence (0) or presence (1' ol a sniootli
srn-faee, lirae (fine, parallel grooves: Arnolil, U)(i.5). and
ribs were defined and coded as binan cliaractcr states.

Characters 39—45: Color of colitmcllii. 'Hie eoln-
mella of llie sliell ol stroinbid species exhibits a \ariet\
ol colors. Absence (0) or presence (1) ol 7 colors were
defined and coded as binan' character stat<'s: white,
black, orange, brown. [)niple, pink, and nianxc.

Characters 4(i-54: Color oj (ipfiiurc. The ap<'rtnre
of the sliell of sti'onibitl species exhibits a sariet)' ol col-
ors. Absence (0) or presence (1) of 9 colors were defined
and coded as binan character states; white, \ellow. |inr-
ple, brown, rose. tan. niau\'e. cream, and orange.

Figures S-9. lypes ol month of some species ol Stromhidac
tharaeters 39—12). H. 'stromhoid;' 9. 'elonsiate'.

C;haraeter 55: ionjuc oj sij)lioiuil <(in(il The si-
phonal canal ol (he shell ol some stromljiil species is
twisteil abonl i(s own axis. Possession of an untwisted

(0) or a twisted ( 1 ) siphonal canal were defined and cod-
ed as tiistincf character states.

Character 5(J: Ciimihirc oj sijilioiitil cdrud The si-
phonal canal ol the shell of some strombid species is
cnnedi in the plane ol tlie aperture. Possession of a iion-
cnnftl (0) or .i (.niAcd I 1) siplional canal were defined
and coded as distinet eluiiaeler states.

C^haracters 57-()3: \iiiiihcr of circum-apcrftirnl pro-
jcrlioits. Shells ol lanibid sjiecies pnnide some ol the
most striking examples ol sexual dimorphism among ma-
rine gastropods. Shells ol liMnales are larger than are
those of males, and. in .iddition, tliere are differences of
shape. For example, eirinm-a|iertural projections ema-
nating from shells ol leniale Laiiiliis laniliis are directed
dorsalK. when-as those ol male L. Imnlji.s are ilirectetl
posteriorK. During copulation, each partici|ianl positions
its shell so that its stromboid notch (a parabolic impres-
sion at the anterioi |)or(ion ol tlie .ipertnre oi the shell
of most strombid species i is adjacent to that of its part-
ner. Tills spatial arrangement ol eircnm-apertural pro-
jections enhances close positioning of shells and may la-
cilitati- coition (Abbott. 1960. .Absence^ (0) or presence

(1) ol specific nnmbeis ol eirenm-apertural projections
were defined and coded as binan' character states: 1, 5.
(x 8. 9. 10. and II,

f:ladistie Analv-si.s: Chidistie anaKsis ol the data i.see
abo\e and :ippendixl was conducted using the computer
program IleniiigS(') (Farris. 19SS). designating the three
species ol' Stroinhus as outgroujis las the\' are co-iainilial
with species ni' I.iiinhis: .\bbott. 19fi0i and invoking tlie
implicit eniniiei':itioii option ie iwliich determines a
maximnm of 100 etpnilK inost-[)arsimonions clado-
granis). Fach of (lie (hree species of Slrotnhiis was in-
N'oked as the priman ontgionp in (hree separate aniJy-
scs. .\ut;i|ioiiioq)hic charai'fer stati-s were omitted from
aiiaKses to a\oid inflating consistency indices (Wiley et
al.. 1991 I. To provide a measure of node support, the
bootstrap resampling procedure was conducted (San-
derson. 1989; ilillis and Hull. 1993: Felsenstein and
Kishiiio. 1993; Saiulei'sou. 1995; Caqienter, 1996) ii.sing
(he i'inin)u(er program l>an(loni Cladistics (Siddall,
1997): (.'lades were obtained In cladistic analysis of tiie
original data matrix; the percent occuneuces of partic-

Page 94

THE NAUTILUS. \ol. 115. No. 3

iilar flack's thus (k-fiiicil in i-ladom'aiiis rfsiiltin'.i lioin
fladistic anaKsi.s ol data matrices created 1)\' rcsaniplinsj;
were determined (this procedure differs from tliat tie-
scribed in Fel.seustein, 1985, wliicli \ields a "liootstrap
e.stimate(l pli\Iogeii\"'. It was assumed tliat characters
were unassociated and charact(>r states were indepen-
dent.

RE.SULTS .\ND DISCUS.SION

Two most-parsimonious clailonrams were ohtainctl
(length = 93, consistency index = 45. retention inde.x
= 46). oni' if eitlier Stroinhns hulUi or S. dihitahis was
designated as the prime outgroup and one if S. vomer
was (figures 10-11). Each of these cladograms cimtaineil
a clade that included .ill 9 species of Lainhis. one in-
cluding S. vomer and the other inchiding S. bitlla and .S.
ililiiUilns. Bootstrap \alues ranged from 0 to 38%.

C^hulograin: In a ciadistic anai\'sis. cliaracter states of
each cliaracter are considered with respect to character
states of each of the otiier ciiaracters, and the cladogram
ohtained represents the most parsimonious grouping of
potentialK mutualK" e\clnsi\e Inpotiieses of clade mem-
bership. The stmctures of the cladograms containing
species of Lambis were predoniinantK' determined h\
character states associated with the railnla and shell
(which comprise 56 of the 63 characters; figures 10-11).
In particular there is a tendency tor characters concern-
ing the radula (characters 1-19) to be represented by
s\napomoiphic states that determine the basal structure
of tlic clatlogranis and tor ciiaracters concerning the
shell (characters 27-63) to be represented b\' sxnapo-
moiphic states that determine terminal structure. There
is a tentk'nc\ for characters concerning the soft bocK to
be represented b\ autapomoqihic cliaracter states.

One cladogram is completeK pectinate (figure 10),
whereas the othei' includes a I'lade containing species of
Lambis and Stronibiis (figure 1 1 ). Within the completelv
pectinate cla(k)grain. S. bulla. S. diliilatus. L. erocata (a
member of the subgi'uus Lambis}. aiul S. vomer se-
cjueiitiallv are sister groups ol all other ta\a: within tfie
incoinpleteK pectinate cla(k)grain, S. vomer is the sister
group ol all other taxa. and /.. rrocala is the sister group
of till' clade comiirised ol .S, build and S', dilalafus (this
clade was determined b\ Hie alisrnce ol tricuspid lateral
teeth (character 8). licxacnspic diiler marginal teeth
(character 17). beaded whorl sniplnre i cliaiaiter 35),
and lirae (character 37) and the presence of smooth
whorl sculpture (character .36) and a brown colored col-
umella (character 42)).

The classifications obtainable from (he cladograms
(figures 10-11) are inconsistent with prexious classifica-
tions of the faiuily Strombidae and its member genera
(e.g., Abbott, I960; 1961); Lambis is paraphvletic. and
Sirombus is polypliyletic. The classifications also are in-
compatible with current snbgeneric assignments: the
subgenus Lambis (L. hnnbis. L. erocala. and L. Innieata)
is ]iol\pli\letic. whereas the subgenus Millejies (L. seor-

j)ius. />, uiillejieda. L. dinilala. L. robusta. and /,. vio-
laeea) is parapli\letic and includes the lone member of
the subgenus C.liira<ira iL. ehiragra). Because prexious
classifications of strombid species were established non-
cladisticalK on the basis of simihir information, the in-
coinpatibilit\ of these hypotheses with previous classifi-
cations results from the pre\iousl\ undocnniented \'ari-
ation obsc'ned and the li\pothetico-dedncti\'e testing in-
\c)l\c'd in ciadistic anahsis.

Since the publication of .-Abbott's (1961) monograph,
two new species of the subgenus Millepes ha\e been
described soleK' on the basis of inionuation concerning
shells; Lambis araelinoidcs Shikama. 1971 (Shikama,
1971) and L wheehvrit^hti Greene. 1978 (Greene.
1978). Kroneiiberg (1993) concluded that both taxa refer
to Iwbrids of L. tntneata sebae (a member of the sub-
genus Lambis) and L. millepeda (the tvpe species of the
subgenus Millepes). as hybridization between members
of their respective subgenera occurs (e.g., L. seoiyius
.\eoii)ius X L. eroeata erocata. and L. millepeda X L.
lambis) and shell moiphologies of hvbrids are interme-
diate betxveen those of the two putative parent species.

The subgenus Millepes is characterized bv ". . . elon-
gate apertures bearing well developed lirae. bv a si-
phonal canal which is either straight or cuned to the
right, and bv the presence of sLx to ten labial digitations"
(Abbott, 1961). Kronenberg (1993) suggested that, be-
cause species of the subgenus Lambis have elongated
apertures, variablv shaped siphonal canals, and 6 circum-
apertural projections, the presence of lirae might remain
as the onlv diagncjstic character state of Millepes. How-
ever. L. tniucata sebae can exhibit lirae on the outer lip
(Kronenberg, 1993). (liven this obsei"vation and the fact
that hvbridization betyveen members of the subgenera
Millepes and Jjiiubis occurs. Kronenberg (1993) sug-
gested that these two taxa should be svnonvmized and
the concept of Millepes as a subgenus should be aban-
doned.

,\bbott ( 1961 ) offered a c;iution;in- remark concerning
svstematics of Lambis: "... from ;i biological standpoint,
some woikers might wish to consider them a subgenus
of Sirombus." On the b;isis of this caiitionan' rcMuark.
Krcinenbergs (199.3) ol)sen;itic)iis ;iiid conciusions, ;ind
the cladograms obtained, the validity of current geueiic
and snbgeneric- classifications of Slro)ubus and Laiubis
are dubious ;ind, consecjuenlK. the cnirent ciassi(ic;ition
of the familv Stiombickie is nnteiuililc.

Reassessing liypollieses of morpliological ey olulion
and function ol circuni-a|Jerlural projections: The

ehidogiiims ciinlaining species of Laiubis (figures 10-1 1 1
mav be used t(i ic';issess hvpothc"ses ol moipholo^ical
evolution oi ciic'um-apeiinral projections. In considering
distributions ol cluiraeter states representing numhers of
circum-;ipertural projections on terminal nodes as "at-
tiibntes" isensu Deleporte, 199.3), inlening most-parsi-
monions cliaracter st;ite trimsformations. and iuteipi'et-
mg bivmching |iattirns lo reprc>sent reconstrnciions of
pliv l(}^cnetic liistoi-v (bec'ause the (intiiionp eiiterion lor

J. R. Stone, 2001

Pasre 95

10 6 1

9 5 8/9 10/11 6 (.

t<lC/5>«JC/5'^"«jKj^^^.^ nJ

, 1 27. 2S

1,14.I9,2.S,4(),51

27,28-0.37-0

J

2«)-0,3(l,48

13-0,18,31.35-0,42-0

. 11-0,16-0,59-0
J 2-0,6,22,55,56-0

9,11,13,20,21-0,54-0

J 8,1 7,35,36-0,37,42
J 11-0,12,33-0,39-0,59

8-0

10

1 1

0 6 6

6 9 5 8/9 10/11 6

^ *• *»«w "^ sj ^ •*■ **^ '"^ •«*• ^ •

^ ^ >t

i^ ^ ^ — :^

l__JIj3(),55-0,59

. I 27,28-0,37-0

J 29-0,30,48

^

J 13-0,18,

31.35-0,42-0

11-0,16-0,59-0

2-0,6,22,55,56-0

9,11,13,20,21-0,54-0

59

11

Figures 10-11. ( lidii^ranis inMiltiii'4 rioiii clailistii- anaksis iildala .Irsciib.-d in Materials and Mctliods' tliat iiicliRk- spt-dcs
cunvntK classified as Lmiiliis (eliaracters are indieated li\ iminliers, and cliaraeter states arc 1 unless specified with --O). .Svnapo-
nioiphic cliai-acter states are indicated on inteniodes, exeejit loi' lliose shared In S hiill,L S dilatatus. and /,. crocata (S-O. lT-0.
35-0 36 37-0 42) and those shared 1>\ S. hnlhi and S dilalalus ill, 12-0, 24,33, 59-0). .Vntapomoqihies are omitted for clarit\'
(S. huUa: S 27 31 50, 55, 57: S, dilalalns. 10. l(-l-0. 17. 30. .32. 34-0. 3S, 39, 46, 4S, 49. 54-0; .S. vomer. 23, 24, 2S-0. .38, 41, 57;
L. crocata: 7. 22; L. scoiyius: 12-0. 25. 27, 43. 4S; /. lamhis: 1. 15. 36, 37-0, 40, 50, 51, 53, 54; L. millcpcda: 3, 8-0, 28-0, 34-0,
44, 52, 61; /. c/iJ/YVfiw: 4, 22-0. 26. 43. 44, 46, 58; L. dh^itata: 45. 52. 60. 61; /.. violacca: 62, 63; L. rolmsKi: 2. -5-0, 9-0, 11, 42,
48-0. 53. 56; /. tnwrala: 1. 14, 19. 2S. 46. 51). Specific nnuihers ol circuni-;ipertnral projections are indic;ited on terminal nodes.

Paee 9fS

THK NAUTILUS. \ol. 115. Xo. 3

12

c1 c2 c3 c4

t1 1 1 1 0

t2 1 1 1 0

t3 0 1 1 1

t4 0 0 1 1

t5 0 0 0 0

Fij^iires 12-13. Hcconslnictiiin cvnlutiDii (il allrilmlcs A clailislic ilal.i iiiatiix loiisistiiiL; nl tlic ."> t,L\a 1 1 5 ami 1 cliaractiTs cl-
4 is ii.scd to construct two cliulosjranis on tlic Icnuiiial nodes ol wliicli cliaractcr stati'S ol cliaractiM' 2 arc distiihulcd as 'attrihutos"
(scnsn Dcleporle, 1993) for historic reconstruction. 12. The cliaractcr states of cliaractcr c2 are included in the cladistic anal\si,s.
13. The character states o!' character c2 are exclnded. The two cladoi;rauis dilTcr. (.Iciuoustratiui; that. In exclndiusj attiihutcs from
a cladistic analysis, dillerent clados;rauis conld he used to test hxpotheses coucerniug the same group of taxa. using the .same
characters. lucludiug attrihntes is the more consistent procedure logicalK.

J, H. Stone, 2001

Pm- 97

(.■liaracter-statc aidiiitj; is iTulfpciulciit dI cxoliilionaiA
processes, the eodinti ot eacli apoiiKiipliN is iiulfpciKlcril
of the stnicture ot the entire elatlistic data matrix, the
cladoiirani ri'sulting from cladistii- an.iKsis. and the pli\-
logenetic inteipretation ot tliat eladoiirani: tlins. there is
no loijieal eirenlarit\ in inclnthng attriliutes iis eharaeteis
in elatlistie analyses — i,e,, apomoipliies are "piimaiA
statements," anti piniogenetic Inpotlieses are "seeond-
arv statements" isensn Deleporte. 1993); in laet, l)\ i'\-
elndinij an attrihnte, difterent ciadograms wouhl lie used
to test hspotheses concerning tlie same gronp ol la\a.
using the same cliaraeters (figures 12-13; Deli'porte.
1993")). T\\f most-parsimonious Inpotlieses eoneerning
the nniiil)er ol eircum-;ipertural pi'ojei'tions present
upon their origin are e(jui\ocal: t)n the basis ol the loin-
pleteK' pectinate cladogram, the ancestor ot all the spe-
cies possessed a set comprised ot either 0. 1. or (i cir-
cuni-apertural projections; on the basis ol the iucoui-
pleteK peetin;ite cladogram, the ancestor ol all the spe-
cies possi'ssed a set comprised ot either 1 or fi,
Subse(juent to the origin of this prominent fe;itm"i\ tlie
number ol eii"c'um-;ipi-itnial projections was modified in-
dependentK with the origins ot L iiiillcjHdii. L rliini-
gra. L. (li^ifdld. ;ind L. viohicra^

These clailograins also can be useil to lorninlate li\-
potheses of functions ot circuni-apertural pi'ojeetious.
Circum-apertui'al projt-ctious lia\e been shown to eonter
protection against crushing (Palmer. 1979), and it has
been postulated that the\' have evoKed ;is a response to
predation (\ermeij, 1989). This hxpothesis coulcl be test-
ed. In' comparing the evolution ot circum-;ipertunil pro-
jections with till' ex'olution ot crushing appar;itus ol
predators, each inferred troni independent cladogranrs.
In particular, a correlation should exist between nnniber
(and strength conferred hv possession' ol cirenm-aper-
tural projections ;ind cmshing strength ol pretlatois. The
hvpothesis that circinn-apertural projections piciviih' sta-
bilitx' on substrates (Sa\azzi, 1991) could be ti'sti'd In
considering distributions of character states ri'presenting
nuiribers ol ciiciuii-apeitur;il projections and luibitatsou
the terminal nodes of the cladograms and mler|iretiug
them phvlogeneticalK, Perhaps a correlation exists be-
txveen number (and stabilitx proxided In' possession) ol
circum-apertural projections and the txpe ot substrate
preferred b\' strombid species, suggesting a renuirkable
adaptation of foi'm to function or \ice-\ers;i. 'Hie li\-
pothesis that cirtuni-apertnral projections provide
means ol circuuneutiug geometric constraints dui'ing
ontogenx' (Sa\';i'/zi. 1991 ) could be elaborated ;in(l ti-sted
bv integrating infornuition concerning ontoi.ien\ into a
phx'logeuetic context. ,\ correlation might exist betxveen
numbers of circum-apertural projections and 'time-to-
maturation,' which would be discemable b\ eousideritrj;
distributions ol iippropriate character state's on the ter-
minal nodes of the cladograms. FiualK. the possibilit\
that enhanced stabilitx' on substrates as a consequenci'
of increased drag conferred b\' the possession ot circum-
apertural projections could be tested from within an his-
torical context. b\' considering biogeographic;il disti'ibu-

tions. tests ol llnid-d\ lumiics, and the bi'anchinii pattern
dipicted b\ the cladograms. Correlations among en\i-
romnental, h\(lrod\namie, and inferred temporal infor-
mation would indic;ite whether numbers of circum-ap-
eituial [irojeetions ha\e e\oKcd in response to difterent
h\drod\namic environments encountered during evoln-
lion. ( au'i'entK, testing of these hvpotheses is delaved In
insutfii-ient information concerning predation, lack of
knowledge ;iboul the evolutional histories of predators.
sc;int docnmeut;ition of lite-histories, habitats, and de-
velopment ol sti'ombid sjK'cies. and the paucih' of data
concermug hvdiodv n;iinie propi'rties ol their environ-
ments.

.\( KXOWLKDCMKXr.S

.Specimens mentioned in this p;iper were provided bv
curators D. C.'alder and K. ( ,'oates iRoval Ontario Mu-
seum: f-lOMl. I. Loch e\ustraliau Museum ot Natural
Histon'). and C. Hosenberg (Acadenu' of Natural Sci-
ences. Phihidelphia; .\NSP); information concerning
those specimens was provided bv .\1. Kitson (.ANSP) and
\i. Zubowski (HO.M); and additional specimens and in-
formation were provided In .A. Baldinger and T. Kausch
(.\gassiz Museum of Companitivc Zoologv'), P. Mikkelsen
(then at the Delaware .Museum of Natural Ilistorv), and
E. Lazo-\\'asem (Peaboch' Museum of Natural Histon).
Illuminating assistance with scanning electron micros-
copv was pnnideil In P. Kalm. 1^. \'illadiego. Dr. K. Lin.
and B. T .Moose |r ( .'onstruetive eonnuents concerning
(.'laclistic analvsis and suggestions lor supplving support
uii'asiu'es tor cladogram nodes were pnnided In 1). (.'-.
Keid anil two r<'viev\ers. Kditorial recommendations re-
lated to reporting of results were provided bv J. H. Leal.
Moral support ;in(i inspiration wci'e pnnided bv NL Tel-
lord, Financial sui)port w;is providetl bv a Malacological
Societx' of Lontlon (."I'utenan Research Crant, a C^on-
chologist of .\meriea Research CIrant. Natural Science
and Engineering Research (Council ol Canada (NSERC)
Individual Grant 4696. an Ontario (iraduate Scholarship.
an NSERC' Postiloctonil Eellowship. a Swedish Natiu-al
Sciences Research Coimeil Postdoctoral Project C.rant.
and a ('anadian Institutes of Health Postdoctoral Eel-
low shiii.

LITKR.VrURE CITEI)

\|ili(ill. H. T 19(-)1). 'I'he s^eniis StrKinlius in tlic Indo- Pacific.

Indo-l^icific .Molliisca 1: .'J.-J-l-ie.
Alihiill. R. T. f9B]. The genus Laiidm in the lM(l()-l';icific.

Indo-1'acific Molhisca 1: 147-174.
VriioKl W !I. 1965. ,\ glossaiy of a thousand-aiid-onc terms

iiscil HI eoiieliologv. The Wliger 7 (supplement): 1-50.
C ;;irpcnt(r, |. M, U)9(i, I'ninfnrmativc hootstrappiiiii. C^ladistics

12: 177 ISl,
Deleperte, 1' 1993. Cliaraeters. attriliutes and tests of evolii-

liciiuii'v scenaricis. Cladistics 9: 427-432.
I'.irris. |. S. loss, Uc)tn'r^S(>. \crsioii 1..5. Program and Doeii-

iiient;ituin. Port |elleisoii. New York.

Page 9S

THE WrTILUS. V(,l. 115. \(

Greene. J. 1978. \ new .species of Lanihis (Molliisca: Stroni-

hitlae). La Coneliijjlia 10: 110-111.
Feisenstein. J. 19S.5. (Confidence limits on plnlogenies: an aji-

proacli nsin<i the bootstrap. K\olntion 'W: 78.3-791.
Feisenstein. |. and 11 Kislnno, 19!).). Is tlicre something wrori'j;

with the bootstrap on plniogeniesy .\ repK to Hillis and

Bnll. .S\stematie Biolo<r\ 42: 19.3-200.
Ilillis. D. M. and J, J. Bnll.' 1993. .-Xn empirical test ot boot-
strapping as a method lor itssessing confidence in plivlo-

geni'tic anaKsis. .S\'stematic Bioiogx' 42: 182-192.
Kronenberg. (.',. C. 1993. On the identih ot L/iii>his ulircl-

nriiihli Cireene. 197S and L ciriiiliiiDidi-'. Shikam.i. 1971.

\ita .Manna 42: 41-5(1
Maddi.son. W. P.. M. J. Donoghne. and 1), I! M.idilison. 1984.

Ontgronp ;inal\sis and paisinionx, S\ sliniatic Zoologx 33:

8.3-103.
McKm. \ (;. 1973. Seashcll I'aradc. Charles E. Tnttle. Knt-

land. .3(i9 pp.
\i\on. K, ('.. and |, [. Da\is. 1991. PoK niorphic taxa. missing

\ahi('s and cladistic anaKsis. C^adistics 7: 233-241.
Palmer. .\. H. 1979. Fish predation and the evolntion of ga.s-

tropod sliell scnlptnre: experimental and geographic exi-

dence. Exolution 33: 697-713.

Platnick. N. 1., C. E. Griswold. and J. .\. Coddington. 1991.
On missing entiies in cladistic anaKsis. ("ladistics 7: 337-
:54:5

.Sanderson. .\1, |. 1989. CJmtidi'nce Innits on ph\logenies: The
bootstrap revisited. Cladistics 5: 11.3-129.

.Sanderson, M. J. 1995. Objections to bootstrapping phvloge-
nies: a critique. Systematic Biologv 44: 299-.32().

Sa\az/.i. F.. 1991. Constructional moiphologx ol stronibid gas-
tropods. I,ethaia24: 311-331.

Se\er. J. O. 1994. StRicture and optics ol the eve ol the hawk-
uing conch. Stroinhus mniiuis (L. 1. Journal of Experi-
mental Zoologx- 268: 200-207.

Shikania. T 1971. On some noteworthx marine gastropoda
Ironi southwestern Japan (III). Scientific Reprints of Yo-
kahama National Unixersity 8: 27-.35.

Siddall. M. E. 1997. Random Cladistics. N'ersion 4.0. Uni\ersit\
of Toronto, Toronto.

Wrmeij. C. |. 1987. Evolution ami Escalation: .\n Ecological
Histon ol Life. Princeton L'niversitx- Press, Princeton, 527
pp.

\\'iley, E. O.. D. Siegel-Causev. D. R. Brooks, and \'. .\. Funk.
1991. The Complete Cladist. The Uni\ersit\ of Kansiis
Museum of Natural Ilistorv Special Publication 19.
Lawrence, Kansas. 158 pp.

Appendix. Data m.itrix with coding ol characters states used ni the ckulistic anaKsis ol species ol Lamln:
Numbered (.haracters are explained in the text. 0 = alisence. 1 = presence. ? = undetermined

Stromhtis l>uUu
Stroinhus (lildtiiliis
Stromhtis toiiicr
lAiiiihis hiiiihis
lAliuhis Irtiiiccitii
lAliiihis ciociitti
lAiiiihis millcpcdfi
lAiinhis (liiiitiita
lAimhis i<)l?iist(i
lAimhis scorpius
lAiiiihis rioldccci
humhis chirai'ni

I 2 3 4 5 6 7 8 9 10 1 I 12 L3 14 15 16 17 18 19 20 21 22 23 24 25 26 27 2S 29 30 31 .32

'^ '^ ■-- ':' I (10 I (Id 1 (I (I 0 0 I (10 0 ■-■ '■' '■' ;^ -■ ■■' '-- 1 I I ' 1 0

0 10 0 1 (I 0 0 0 1 I 0 (I 0 0 0 I 0 0 (I 1 0 0 I 0 0 0 I I 10 1

0 10 0 1 (I 0 1 0 0 0 I (I 0 0 I 1 0 0 0 10 1 I 0 0 0 0 1 0 0 0

1 0 0 0 I 10 1 I 0 I I 10 1 1 10 0 10 10 0 0 0 0 1 10 (I 0
10 0 0 1 I 0 I I 0 0 I (I I 0 0 1 1 I '.^ ':' 1 0 0 (10 1 1 0 '-' 1 0
':' ':" J 'r' I (I 1 0 0 O 0 I 0 0 0 I 0 0 0 '^ '^ 1 0 0 (I 0 0 1 1 0 0 0
0 0 10 1 I 0 0 10 0 1 I 0 0 0 I 0 0 I 0 I 0 (I 0 0 0 0 I 0 (I 0
'J 'J "J 'J 'J 'J 'J 'J 'J 'J '-* 'J ■j V 'J '-> 'J 'J 'J V '■' '-* ■-* '-' '-' '-' 0 I 0 '"' "^ f^

0 1 0 (I 0 10 10 0 1 1 0 0 0 0 1 10 10 1 0 0 0 0 10 0 1 '^ ?
(I 10 0 10 0 110 10 10 0 1 1 (I 0 I 0 0 0 0 1 0 I 1 I 0 ? ?

? ':" '■' ':" '.-' y ? ? y '-' '^ ':' ? ? ? ':' '^ '■* '.^ '^ '^ ':' ':' ':' ':' '^ 1 0 0 ? 1 0

(I 0 (I 1 1 1 0 1 1 0 (I 1 0 0 0 0 1 10 1 0 0 0 0 0 10 1 1 'r' 1 0

33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63

1 1 0 1 (I 0 I 0 (I 0 0 0 0 0 0 0 0 10 0 0 1 1 1 1 (I 0 0 0 0 0
I 0 0 I (I 1 10 (I 0 0 0 0 10 1 I 0 0 0 0 0 0 1 0 0 0 0 0 0 0
0 1 I 0 i 10 0 1 10 0 0 0 0 0 0 0 0 0 0 10 1 I (1 0 0 0 0 0
(II 1 10 0 0 10 1 0 0 0 0 0 0 0 1 I 0 I 1 1 0 0 0 1 0 0 0 0
(110 1 (I 0 ':' ':' ':' ':' ':> '^ 'r* 1 0 I 0 0 1 0 (I 0 0 0 0 0 1 0 0 0 0

(110 1 (I 0 ^ '^ '■• ■-' ;■' ;■' '^ 0 0 o o o o o o i o i o o i o o o o

(I 0 10 1 0 0 (I 0 1 0 (I 0 0 0 (I 10 0 1 (I 0 1 0 0 0 (I 0 1 0 0

? y ? 0 1 (I 0 0 0 0 0 (I 10 0 1 II 0 0 I (I 0 1 0 0 0 0 1 1 0 0

? ? ? ? '^ 'r' 0 0 0 0 (I (I 0 0 I (I 0 0 0 (I 10 0 1 0 (I 1 0 0 0 0

? ? ? 0 1 0 0 0 0 1 I (I 0 0 0 I 0 0 0 0 (I 0 0 10 0 1 0 0 0 0

0 1 0 ? '^ 'r* ':• '■' ':' ':' ':' ':' ':' 0 0 I 0 0 0 0 0 0 1 0 0 0 0 0 0 11

0 10 0 1 0 0 0 0 0 1 10 1 0 0 0 0 (I (I 0 0 'J ':' 0 1 (I 0 0 0 0

Stroinhus hulld
Stnjnihus clilattilus
Stroinhus vomer
lAimhis laiiihis
Iximhis truiicutd
lAiinhis croidid
iMinhis miUipcdd
Luinhis diiiitdtd
lAimhis rohustd
lAimhis scorjiius
lAimhis lioUncd
Iximhis chiraera

THE NAUTILUS 1 15(3):y9-104, 201)1

Pagf 99

Rediscoveiy of Canidia dorri W'attebled, LSSCl witli clisciissioii
of its systematic position (Gastropoda: Neo^astropoda:
N as s ai"i i dae : Nassodc )nfa

Yuri I. Kanlor

A. N. Severtzo\ liistitiiU' (if I'lcililcms

of E\'oliiti()ii
Russian Ac;ul('iii\ ol Sciences
Leninski Prospect 33
Moscow 117071, RUSSIA
kiinti )r(?'n ialac( i-se\ i n , n i sk. n i

Richard Neil Kilburn

Natal MiiseiiiM
Pietcrniaril/liniii
SOUTH AFRICA
dkilliiunl" nnisa-oi'il.za

ABSTRACT

Canidia doni W'attehlecl. lSS(i, dcscrilii'd lioni Kao-lial La-
goon (near Hue, N'ietnam) hut until now known onK Irniii its
orisjinal description, was rediscovered in lower pails nl n\irs
at Phan Ri, centra! Vietnam. Examination ol iiKirplidloijA and
radula ol the .species revealed, that it slionkl lie allocated to
the laniih' Nassariidae and, according to shell iiioipholog\, to
the genus Na.ssoilontn H. Adams, 1867. This is the first eon-
firmed record of genus Nassodontd outside IniHan waters. The
onlv other Asian species of the faniiK Nassariidae known to
inhahit fresh or hrackisli waters were found in Lake Uliilka and
other back-water areas in India.

Additional An/ um-tls. Casfnipod, hrackisli water, .Asia, \'iet-
nani-

INTRODUCTION

Diiriiig a \isit to the I'islieries llni\frsit\ in Nlia TraiiLi;
(Central Vietnam), a,s part of the Tiopieal .Marine Mol-
lusc Pro'j;rain (TVIVIP), the autliors came across a sam-
ple ol an iiiiiisiial iieogastropod. These spi'einieiis pos-
sess a deep liasal spiral sidcus, similar to that found in
the Pseu(.loli\idae. The)' were showni to Dr. N<j;uN-en
Ngoe Tliaeli, a mollusk specialist and shell dealer in N'lia
Trang, wiio recognized the species and oilered to collect
additional samples with precise data. \ sample was siih-
sequentlv provided to the authors with tiie note that the
species inhaltited tla- lower parts ol N'ietnamese rivers.
Examination of the radula rexealed that the species lie-
longs to the famih' Nassariidae. It was coni-lnded thai
the species is congeuteric with and closeK related to
Nassodoiild in.siis.nis H. Adams, 1S67.

While I'.xamining collections of freshwater neogastro-
pods in Muse\nri national d'histoire natnrelle, Paris, the
senior author came across two svnitvpes ol Caindia dorri
Wattehled, 1886, desciibed from Lagune de Kao-liai, \'i-
etnam. The figured sviitvpe (Wattehled, 188fi: pi. 111. fig,
5) differed markedk- from the seconil one. wliicli in turn

exactK matched our specimens. PortnnateK. Iiotli s\ n-
tvpes contained a dried l)od\. which allowed us to ex-
amine its radulae.

Since Canidia dorri seems not to lia\e been reported
since its original description, we here give a detailed re-
tlescription on the basis of both t\pe mateiiid and re-
centK eollectetl spi-cimens.

SY,STE.M.\TICS

Class Castropoda Cu\ier. 1797
Order Neogastropoda Wenz, 1938
Superfainilv Buccinoidea Rafinesf|ue, 181.5
Famil\- Nassariidae Iredale, 1916
(Melius .Vr/s\iir/i)///^/ 11. \daiiis. 1867

T\pe species
1867.

h\ iii<)iiot\p\ I; Xasscnhnilii insi'^nis 11. .■\danis.

Remarks: .Mthongh Ceriioliorsk\ » 1984: 199) regard-
ed Sassodoiita as a subgenus of Xassarins Diunerii.
1806. we consider i( lo be a lull genus, characterized b\'
tlie presence of a conspicnonsK deep basal groove (sul-
cus) and b\' its distincti\e radula.

The radula oi' Sassodonl a. as exemplihed by thi' spe-
cies here described, resembles that found in the genera
Biillia and Biiccinanojis (see Cernohorskv, 1984: figs 78-
90), rather than tvpical Xassariiis. in the prominent de-
velopment of accesson cusps on the lateral plates and
in the less pectinate appearance of the cusps on the
racliitlian lioweNcr, ac'cesson* cusjis occur in at least one
t\])ical Xassariiis (see Ceruohorsk-v 1984: fig. 31). and
these characters should be regarded as homoplastic.

Altlioiigli Xassodonfa was originally described in the
Biiecinidae. Smith (1895) transferred the genus to the
Nassariidae basetl on shell characters. The presence of
a nudticuspidate rachidian tooth of the radula. as illus-
trated here, confirms this family allocation.

The t\pe localit\- of Xassodonia insi^iiis was given as
the Peilio River, China (which is possibly the modern

Pa<4e lOO

THK NAUTILUS. \oI. 115, Xc

iMl^iircs 1-13. ^\h-\\s(>\ Sii.ssoihiiild. 1-}J. .\ris\,Hloiila don-i (Wattcblcd, l.S,S()). 1. Apcrliir.il. ,iikI 2. tlmsal \ic\\.s ol fisuu'd s\iit\pc,
\l\ll\. 3. AiKM-tiiral. and 4. dorsal views of syntvpc, MNIIX. 5-9. Spcciiiu'iis i'loin I'liau Hi, central N'ietnani, 5^ Apcrturid. 6.

Y. I. Kantnr and R. N. Kilhum, 2001

Pa.'c 101

Hai He River, wliicli enters the sea at Tieiisien). How-
ever. Sniitli (1S95) pointed out that tiie actual laliel ac-
conipan\inu; the h(>lot\pe read "Peihoi" and referred to
it as ha\ing been collected together with Vcloiita [tlie
corbiculid genus Villorita Gray. 1834J. Noting that the
latter was an Indian genus and that undoubted examples
ol'N. insioiiis liad been collected in Intlia, Smith <iuerie(l
the Chinese origin ot the h()lot\pe. Preston il91fi) and
Cemohorskv' (1984) confirmed the occurrence ol .\'. in-
signis in India, living SNnnpatricalK' with a second ta\on,
Nassodonta <ir(iveh/i Preston, 1916, which Cernohorsk\
.s\non\'mised with .V, iiisignis. It nia\- be noted that tlu'
present material trom \ ietnani greatK' I'xtends the oxer-
all known distribution ot \/isso(loiit(i. ,is prexiousK de-
fined b\" Cernohorsk'x.

Na.s.fodiiiiln (li>rh (W'attebled. 1SS6) new combination
Figin-es 1-S. 1.3-1(H

Caiiidia (lorri Wattchled. ISSfi: 56-57, pi. \\. fit^. 2.

T^pe material: Two s\nt\pes. Museum national
d'liistoire Naturelle (figures 1-4).

Type locality': Lagune de Kao-hai (near Hue),

Material exaniinecl: Sxiitxpes, 20 spei-imens lioiii riv-
ers at Phan Ri. now stored at; Zoological Museum of
Moscow State Unixersit)-. ZxMMU No. Lc-2517], 25172;
Natal Museum L5452, The Natural Histon Museum,
London. BMNH 20000.391; Museum national d'liistoire
naturelle, Paris. MNHN unnumbered; National Muse-
um of Natural Histon; Washington, DC. USNM 9()532fi.
905327; .\cademv oi Natural Sciences. Philailelphia
ANSP 403196; Austrdian Museum C.3S6612; National
Science Museum, Tok"\(). NSMT Mo 72fiSS; Zoological
Institution. St. Petersbiug. ZIN 595S4; Unixersitx ol
Fisheries, Nha Trang, \'ietnam.

Distrihulion (Figure 21): Central X'ietnani, liem
Hue to rixers at Phan Ri (al)out 150 km south ol Nlia
Trang). Aiklitional material xvas collected in the lower
parts ol' lixers. at a depth of about 3 meters.
Description: Shell thick, oblong-oxate. usuallx xvith
subcvlindrical bodx xxhorl (broadly fusiform in figured
s\nt\pe) and loxx; obtuse spire; w^iorls distinctlx shoul-
dered, spire somexxhat cxitoconoid, apex mamillate. Pro-
toconch eroded in all specimens.

Aperture oblong, lanceolate, constricted anteriorlx. in-
ner lip xvith xxide smooth callus, its outer edgi' slightix.
but distinctlx raised, xxithout parietal nodule. Outer lip
smooth interuallx, uotclied in anterior portion, wheie it
is cut bx basal sulcus; outer lip strouglx thickened behind

Figiir*' 14. < )pei(.-iihiiii (it specimen in figs. 5-7, length 5.'

nun.

cilge. but not torming xarix. Siphonal canal short, mod-
eratelv narroxx-. dorsallv forming a fiiirly shallow notch.

Shell surface slightly glossy spire vx'horls xxith strong
prosocline axial ribs, in transxerse section rounded and
xxidc 1 than iutei-vals. Axial ribs 9-10 on first and second
xxhoii, on spire extending from suture to suture but on
later xxhorls tlexeloping into nodules, xxhich become ob-
solete mid-dorsallx. In figured sxiitxpe axial ribs are pre-
sent on tlie last xxhorl, there totalling 10.

Base of last wlioil at lexel of parietal columellar junc-
tion xxith a tli'tp, as\ ietricall\- cut iurroxx. Fasciole

conxex xxith groxxth lines onix, .\rea betxxeen fasciole and
basal grooxe xxith 1-3 spiral ridges.

Periostracum smooth, tightlx ;idhering to shell surface,
color straxx-olixaceous. Shell xx^hite beneath periostrac-
um. xxith inconspicuous axial zigzag sinuous brownish
lines, seen at dorsal shell surface, more pronounced im-
mediatelx- aboxe the sulcus. Zig/ag lines max- be seen on
a|)erturai surface of shell. ;is well as dorsallx; although
thex are alxxaxs better iironouneed on latter. Sometimes
tliese lines are reduced to oblique dots above sulcus.

Operculum (figure 14' elongate-oxal. occupxing
slightix more than Vi of the aperture length (including
the canal), xelloxx; semitransparent. xxith terminal, clock-
xxise-coiling nucleus. (Growth lines numerous and thick-
ened. See table 1 for shell measurements.

Analdint/: Moiphologx of one female specimen from
Phan Ri Rixer (figures 5-7' was examined. Due to its
sl.ite of presenation. we were not able to studx' anatomy

dorsal and 7. (il.li<ine dorsal xiexx'S, ZMMU Lc-25171, SL = 14.9 mm. Arrow indicate the !)as;il sulcus. 8. Apertund view. Natal
Museum, L5452Ari764, SL = 14.2 mm. 9. Apertural view. Natal Museum. L5452-T!7(i4. Si. = 13.0 nun. 10-13. Nassodonta
insioim H. Adams, 1867. holotxpe, the Natural Hi.storv Museum, Lond.ni. BMNH 1S7S.1.2S.42S. 10, .\]iertnral, 11. lis^lit lateral.
12. dorsiil, and 13. ohliiine dorsal xiexxs. Arrow indicate the hasal sulcus. All shells at the same scale.

Faw> 102

THE NAUTILUS. \o\. 115, Xo. 3

Figures 15-20. Haclulae of Nn.ssii(li>iila durh. 15. Dorsal \i(\\ ol iiii(l-|«)rtioii ol nidiila of fitjiircd s\nit\pe, MNHK, Hi-IT.
Dorsal \-ie\v of tlio niid-poilioii ol ladiila of s\iit\p(', MMIN. l.S-2(). Hadiila dI tlir spcciincii on \''vj,s. 5-7. ZMMU Uc-25171. 18.
Dorsal view of the niid-poilion, 19. Kinlil lateral (45°) \ii'\\. 20. l.alcial \ icw ol lateral teetli to sli(i« iiideiitatioii on inner side.

ill detail. Tlic rear ciul of the foot posses.ses small, paired
tentacles, altiioiitrji in preserved speeinieiis these were
iiiiieh eoiitraeted. Head with iiiediinndiiinj; eniiieal ten-
tacles with lai'iie lilack eyes at tiieir bases. Anterior part
of the foot is piiimented with small, wiilel\-.s])aced hlaek
.speckles, while the mantle is liea\-ilv pigmented,

Probo.scis in the retracted position — \.2 mm long

(().2S Sl,^ and I ,o mm wide. poorK' pigmented. .Sali\ai"\'
glands [laired, not Insecl. medium-sized. \'aK'e ol Lei-
lileiii small, prononneed. Mid- .Liid posterior oesophagus
was too [loorK pi'eseiAcd foi' stud\.

Hadnla ol fignrt'd s\nt\pe (figure lot .3.5 nmi long
10.25 SI, and 0,.i4 ALi. ~ .350 |jlui wide iO.()25 SL and
0.0.'54 Al,). coniposi'd ol aiiout 75 rows ol tet'th. Lateral

Y. I. Kantcir ami R. N. Killiiini. 2001

Page 103

Nha Trang

G type locality
^examined material

Figure 21. Distiiihiitioii ot \(i>.M>(li)ittii dnir't in Miliiaiii.

teeth with 4-fi cusps (number of cusps \arics e\fn on
adjacent rows), outermost cusp — 2.5 times lont^er tliaii
innermost. Intermediate cusps either shaipK pointed or
bifiu'cating at their tips. Innermost cusp with 7-S den-
ticles on its lateral side. Rachidian with 11-12 cusps,
central cusp serrated (number oi cusps xaiiable e\en on
adjacent rows). Basal plate e\enl\ and dec pK notched
alone anterior edse.

Radiila ol scconil s\ i it \pc j figures Hi- IT) .3.9 mm long
(0.3.3 SI, and O.Wi .\{.). '- .3.50 |jLm (0.029 Si, anil 0.0.32
.\Li. comjiosed of 7.3 rows of teeth. 4-5 nascent. Shape
ol teeth is \cn similar to that of figured .s\nt\pe. Tlie
specimen Irom idian Hi (.siiell on figures 5-7) hax'e \'er\-
similar radula (figures l.S-2()). 4.7 nun long (()..32 SL and
0.44 .\L), - .355 IJ.III wide (0.024 SL and 0.0:5:5 AD,
composed ot .S5 rows ol teeth. 3 nascent.

Nariability (Table 1): Species laiiK miilorm in shell
^liape. The figured s\iit\pe diflers lr<iiii all other speci-
mens examineil in lia\iiig a more swiilleii last whcjrl and
ill the complete absence of thickening of the outer lip
abo\e the edge of aperture. .■Mtliougli fairk' large, this
specimen appears to us to l)e immature, from its thin
ami still simple lip. Moqiliolog\ of the upper part of the
aperture suggests that the thickening of the lip is in its
initial stages, whereas in the otlu-r specimens examined
it is formed at tin' eml of shell growth.

.■\11 other available specimens are much alike. 'I"he
most \ariable character is the degree of thickening of
the outer lip mentioned abovf. Zigzag lines ma\' be seen
on the apertural surface of the shell, as well as on shell
dorsum, although tlie\- ari' alwavs inor(> prononnci'd on
the latter Sometimes these lines are reduced to oblicjue
dots abo\'e the sulcus.

Remarks: The main diiierenci'S between A', dorri and
.V, insi'^ms (figures 10-13) (including iV. (jraf(?/f/i) are the
much lower spire ol the former, presence of strong axial
nodules, shouldered whorls, strong constriction in the
base ot the outer lip. {\\v smooth outer lip and the lack
of a parietal nodule,

DISCUSSION

The species under consideration was originalK' described
in the genus Cunidia 11. .\dams. 1862 (T\pe .species [b\
monot\p\] C. fused II. ,\dams. 1862). The name Cauidia
appeared to be twice preoccupied. Oossmann (19011
iironosed tile substitute name Aiiiiiloiiif. Ancntomc is

Table 1. Slicll iiieasiireiin'iits (iiini) of Xiiwoiltniln ilmri

l'"i'4iircil

S\ iit\ |)i

Cliaracter

s\iit\pe

.-)

K.iii'j,e

-\\orage

II

Shell lenndi (SL)

14.1

12.0

12.0-14.9

13.69

0.93

Body whori length (BWL)

11.6

11.0

10.6-12.8

1L61

0.(54

."Aperture length (AL)

10.3

9.7

8.7-10.8

9.86

0,73

Shell width (SW)

9.5

8.0

8.0-9, (i

S.S6

0,61

BWL/SL

0.S2

0»2

0.81-0.94

0.85

0.04

AL/SL

0.73

(LSI

0.70-0.81

0.72

0.04

SW/Sf.

0.67

(l.(S7

0.61-0.(57

0,(55

0,02

Nunilicr lit spiral ri(lt;es l)et:\\een

fa.sciiile and lias.il groove

2

1 \cn
weak

1-3

2,42

0.:32

Xiunlier of axial rilis on first teleo-

conth wliorl 10

Number of ;ixial ribs mi seinnil teleo-

eiiiieli wlidil 12

9-12

9-i:5

9.2

10,7

1.26

Page 104

THE NAUTILUS. \'(

15. \(). 3

generalK' c'oii.sidered to he at most a .suhgenus ol Cica
A. Adam.s. 1855 (e.g., Thiele. 1929). 'I'lic t\]ie species,
Canidia fiisca. was never illustrated, and tin' loeation ol
its tApe material is uiikiumn. Two specimens (suspected
s\iit\pes) from (^ami)odia. identifieil as "C. fusca" from
the Cuming collection are stored in the collections ol
BM(NH), no. 2()()()l;31fi, and were examined hy the au-
thors. Nevertheless, the\' do not match the original de-
scription and therefore are not t\pes. Thus the real po-
sition oi Canidia remains uncl(>ar The other species that
was originally attributed to the geims is Melanopsis hc-
lenac Meder, 1847, which cleai"i\ belongs to Cica.

Therefore, allocation ol dorri in the genus Cica
(Anentome) ( = Canidia) is certainU' wrong. .A.t the same
time, the shell uioipholog\ is ven similar to that of .\V;.s-
sodouta insif^iii.s H. Adams, 1867. including such an mi-
usual character as the l)asal sulcus. This induces ns to
attribute the species to Nassodonta witli some certaintA.

This is the first confirmi'd record ot genus Nassodoifa
outside Indian waters, despite the original t\pe localit\
of China wxen for N. insi<'nis. Accordins; to Dr. Thach
the species is abundani in ii\crs ot central Vietnam
around Nha Trang and I'han Hang, .although N. dorri
(like N. insii^nis) appears to inhabit the lower parts ol
diese rivers and is probabK' found in brackish waters,
the salinit\' of its habitat nei'ds to be investigated.

The onlv other .\siaii species oi the lamilv Nassariidae
known to inhabit fresh or brackish waters, are Nassariiis
{Pi/ginafuassa) siihconst rictus (Sowerln; 1899), N. (P.)
oiissaensis (Preston. 1914) and N. (P.) fossae (Preston.
1915) from Lake Chilka and other Indian back"waters
(see Cemohorskv, 1984).

.VCKNOWLKIK'.MENTS

The authors would like to express their thanks to the
n,\\Ii).\-spons()red Tropical \hu-ine .Mollusc program
and its director. Prof. Jorgen Hxlleberg. lor pro\iding
(he o])portunit\ to attend their workshop in Nha Trang,
X'ietnam. to l)r Tliach for suppKing us with the material,
to Mrs. Kathie W'av and Dr. John Tavlor for the loan of
liolot\]")e of Nassodonta insiiiiiis and speeimcTis attribut-
ed to "Canidia fusca".

l.lTKH.Vn HI-: (TTKD

.Ailaiiis, H. IS(S2. Descriptions of some new s^encni anil .species

ol shells from the collections of Hugh (aiming. Esq. Pro-

ceeilings of tiie Zoological Societ\' oi London. 25: 383-

3,S5.
Cenioliorsk-v W. O. 1984. Sxstematies of the faniilv Kassari-

idac. 1-iulletin of the .\uckland Institute and M\iseum 14:

i-iv + l-.35fi.
C'ossniann. .\1. 1901. Ess;us de paleoconcliolotjic comparee. 4.

.Author's edition, Paris, 293 pp.
Preston, 19US. Report on a collection ol maniu- .Mollusca Iroiii

the Coi'liin and Enmir backwaters. Record ol the Indian

Museum 12: 27-39.
Siiiitli. E. A. 1895. Notes on Nassodonta iii.siu.iiis. Proceedings

of the M:ilacological SocietA' of London 1: 257-258.
'Hiielc. j. U)29-1934. Handlnich der svsteniatischen W'eicli-

tierkunde. 1(1): 1^376 [1929]; 2(1): 779-1022 [19.34].

Gustav Fischer. Jena.
Wattebled, G. 1886. Description de mollnsques inedits de

rAnn;ini Recolte du capitaine Dorr anx ensirons de Hue.

|nuiiKil de Gonchvlioiogie 34: 54-71.

THE NAUTILUS 115(3):105-114, 2001

Paiif 105

Systenuitics and ecolog) oi' Gdstrocopfa (Gastrocopta)
rogersensis (Gastropoda: Pupillidae), a new species of land snail
from the Midwest of the United States of America

Jeffre) C>\ Nekt)la

Department ol Natural ,uul Applictl

Sciences
Universit\ ol Wisriiiisiii-Creen Bay
Green Bay. \\\ 54."ill ISA
nekoIaj@uvvt;b.eili I

Brian F. C<jlcs

42(1:2 Sci.ttif Siiiitli l)r
Irtlrisoii, AK 72()7i-J ISA

ABSTRACT

Gastniiojitd /(i^iTsc/is/.s, a new species of reecnt gastropod
mollusk (Pupillidae. Gastroeoptinae) is deserihcd from the \i-
cinit\' of the Ozark Uplift and Paleozoic I'latcaii ("nriitless
Area") in the niidwesteni USA. The structure ot the angulo-
parietal "tooth" in G. rogersensis consists of two discrete sub-
parallel lamellae borne on a rectangular callus, distinguishing
it from all GastrocDpta procera (Gould. 1840) subspecies and
variants, inchuliug Gastrocopta proccra mcchiuiii (Hanna and
Johnston. 1913), Gastrocopta proccra riparia (Pilsbn. 1912).
and Gastrocopta proccra sterkiana (Pilsbr\-. 1912). Moipho-
metric analyses demonstrate that e\'en at sites of co-ociur-
rence, C. ro'^erscnsis shells are significantK' (P<0.0(K).5) smaller
than G. procera- Additionally, while G rogersensis exhibits no
\ariation in shell size with latitude (P = 0.8T6). a liighK sig-
nificant lP<0.00()5! cliual \ariation e.yists in G proccra. G ro-
gersensis populations appeal' restricted to undisturbed calcar-
eous bedrock outcrops in areas that escaped W'isconsinan gla-
ciation. The limited range, habitat specificiti.. and potential firt-
sensitivity of this species suggests that it should be gi\en a high
priorit\ for [■onsen ation.

Additional kn/ nords Gaslroropla proccru. uiorphomctncs,
biogeograph\. ecology niidwesteni LISA.

INTRODUCTION

Tile genus C^.aslrtirojitd (Wnllastiin. 1S7S) ediiiprises a
group of pupilliil mollusks ot nearly global distribution
(Pilsbn-. 194S). In tlie Americas, this genus extends into
die nearctie where at least 18 recent species occur east
of the continental ili\itle (Hubriclit. 19S5). Because ol
variabilitv in apertural tlentition aiul sliell size. Gastro-
copta (Qaslrnrojita) pi'occra is one ot the most taxoiiom-
ically challenging memliers ot this grouji. Pilsbr\ i 1012.
1948) regarded G. proccra to consist ol lour wcakk dil-
ferentiated taxa: G. procera. Gaslrocopta proccra
>ncclun<ii. Gastrocopta proccra sterkiana, and Gastro-
copta proccra lonii rijuirid- Ho\ye\er. Hubriclit (1977)
considered G p nicclunaj s\iion\iiious with G proccra

and ele\ated tile remaining two loniis to spe^cific rank.
Untortunateiw no data was presented to snjijiort tliese
conclusions.

During studies on the recent land moiiusks ol tiie
midwestern US.A. we located 19 Gastrocopta popula-
tions from .Arkansas. Illinois. Iowa, and Missouri that
could be reailiK' distinguished Ironi Ga.stmcopta proccra
and its subspecies via a consistent suite ot characteristics.
e\en at sites of co-occurreuee. .An adcHtional 10 popu-
lations ot this form were documented from nortiiuestern
Illinois and soutliwestern Wisconsin tbrongli examina-
tion ot G. proccra material at tlie Field .Museum ot Nat-
ural IlistOA (FMNH). Inspection ot G. proccra material
from the Academy ol Natural Sciences ot Pliiladelpliia
(.\NSP) reyealed that a specimen from Rogers. .Arkansas
also represented this form. This specimen had pre\ ionsK
been identified and illustrated as Qlastrocopta proccra
mcchn<ii (Pilslin-. 1948. figs 493:4-5). Pilsbiy (194S) did
not specificalK' discuss tliis specimen. e\eu though con-
spicuous differences can be seen bet^yeen it ami tlie otli-
(']■ illustrated G p uicchin^i specimen lop. cit., figs 493:
1-3), which w;is stated b\ him to lu' "identical" to the
t\pe.

In this paper wf describe tliese tlistiuct populations
as Gastrocopta rois^crscnsis new species and comment on
its relationship to the Gaslrocopla proccra c-omplex.
moiphometricalK compare it to G. proccra. and consid-
er its biogeograph)' and ecology.

.M.ATERIALS AND XHmions

•Study Populations: \11 populations of Gastrocopta
collected by the aulhors from tiie central US.A were e.\-
amined. Specimens oi' Gastrocopta proccra seiisu lato at
FMNH and .ANSP were also examined. Included in
these were (i\v Piisbi-y (1948) figured specimens ot Gas-
trocopta proccra incclu>i<s,i trom Rogers, Arkansas and
Soiitli Dakota. .AdditionalK, the holotxpe of G p
nircluniii (USN.M 226395) was examined.

Page 106

THE NAUTILUS. \ ol. 1 15. NO. 3

Fijjiires I-S. .Sc-aiiiiin^^ electron niieroiiraphs ol i',<istroci>])lii fiif^crscusis ami Gaslnicopta proccni. 1, 3. Gust rucopta nx^ersi'iisis.
L\\(,B 3914. Fults Hill Prairie Nature Presene. Monroe Coiintx. Illinois. USA (9()°iri5" W. .3S°9'19" N). 2. CastroropUi ro'^cr-
si-nsis. UWGB 1061. Maquoketa Sontli Glade, Cliiiloii Couiilx.^ Iowa. USA (90°39'5" W. 42°U12" N). 4. Caslrocopta prorcra.
U\VC;B 3916. Fnlts Hill Prairie Natnre Presene, ,\Ioin.ie ( (Mnilx, Illinois, US.A (9()°1 1 '15" W. 3S°9'19" N). 5. Castroropta pmrrrii.
UWCH 575, Jnniper Hill Sli.il.- Chide, Flo\(l ( :<.unl\ low.i, rs.\ (92°59'2" W, 43°3'1()" M, 6. Castroropta prorcra inrrlun^i.
liolot\])e, I'SNM 226395. l.oiin Island. Plieljis Counts. Kansas. 7. Castroropta nK^rr^icnsi.s. angiilo-parietal lamella, U^^'GB 3914,
Knits Hill Prairie Xaliire Preserve, Monroe f:oiiiit\-, Illinois, US.A (9f)°I I ' 15" \\', 3S°9'I9" \), H. Castroropta prorcra. angulo-parictal
lamella, UWGB 3916, Knils Hill Prairie Naliiie Presene, Monroe Connt\ Illinois, US,\ i9()°l I ' 15" W, .3S°9'I9" N).

M(>i-|)li(»iiic(|-ic aiiaUses: liidix itiuals were iLS,siij;iu(l
to citlicr Gdstrocoptd new species or Ga.strocopta pru-
cera ha,sed on apcrtural laiiiclla eoiifiguration, Twciits-
fi\-e Gnstrocopta ir-w ,spfC'ies and 24 G', jiidcrra |io|)ii-
lations Ironi the states of Arkansas, Illinois, Iowa, and
W'iseoiisiii wfre used for slicll nioiplioiiictrie aiiaUsis

it<il)lc I', Included were .ill 9 known stations at wliicli
tliesc ta\a co-occini'ed I'lie geograpliie coordinates lor
eai'li population was delcrniiued usiiiti; a Trinilile liaud-
licld (d'S, apin'opriatf US(;S 7,5 luiuute topot;;raplni'
maps or nel.inuie (^a/etlccr, and coiixcrted to UT.M
/.oiie I (■) (.■oordiu.ilcs iisiu'j; .\|{( 'I \ l''( ),

J. C. Neknla and B. F. Coles. 2001

Fatif 107

Table 1. Ldcatiim aiul ci)llt'ctiiiii iiil(iniiali(iii hii kimwn C.(isliiicoj)Ui ro<icr\t'nsi\ sites and iiirasuicd (.'.ustrniojita j)niriTi: sites.

with ninillins dl shells used in iiKupl etne aiiaKses \n \ i(|)reseiits known (.' nincisciisis sites iiiiin where ni) sneeiniens were

measured

Site

Leeatien

Cdlleetiir

( 'olleelion iinnilier

^ \K asiiud

c;

sciisis procci'd

Arkansas

Biixhr ( 'niiiili/
Ni)rl(irk
.SalesMJIe

Benton Cnnnli/

Ri liters
BoiHw ( 'iinnli/
Cinriill C'linntij

Bea\er Dam

Table Boek Lake

liiilil>rn(hncc ('onnli/
Caishni.ui \

Izdict Coiinli/
Calico Keck Kast
("aliee Heek W est

Milili^nn ( 'nuitllj

\\ ill II I iw .Springs Park
Sciu'iij (.mint II

Harriet E

Leslie S

Marsliall \\\'

Marshall S

Stone Connti/
.Mliscin

BarloDt Ht'creation .-Vrea
C'alieo Hoek Sentli
Senth Side S

Illinois

Calhoun Connli/

Franklin Hill
jiiokson ( 'oioitij

Kiiiu;s k'ern IdntI
}o])(ii:wss Conntij

Flizaheth
Miiili^int Connlij

Alton
Monroe Conntif

Fonntain (^ap

Fiilts Hesene

Rtinclolpli i'onntij
Chester
Prairie dii Mother

I<»«a

.\llinniikee (.'onnlij
I'ish Kami .\hiinids

92°lfi'44" W, .">(-i l:V22" N
92°16'52" W. :Wl:V()2" \

93 Dd'OO" W. .5(1 29'(Hr \

94°( )!'.%" W. 3(i°19'59" N
93°4(idS" W. :5(S°2.S'22" N

Brian C^oles
lirian C'oles

Pilslin 194S
(ieori^e Walsh

Brian Coles
Brian Coles

9r47'27" W. 35°53'5S" N Brian Coles

92°0.S'14" W. ,5tS (Ki'4.S" X linaii Coles

92°n,S'55" W. .3(S'=()7'I)I" \ Brian Cok-s

93°43'55" \V. 3B"1)9'07" N Brian Coles

92°29'42" W.
92°33' 19" W.
92°41'39"\\'.
92°35'41"\\;

55'='59'().S" N

35''49' 15" N

5°57'51" N

5°54'21" N

92°1)7'23" W. 35°5(-i'35" N
92"15'1S"\V. 36°()1'16" N
92°08'30" W. 3fi°()6'22" N
91°36'46" W. 35°4()'(K)" N

Brian Coles
Brian Coles
Brian Coles
Brian Coles

Brian Coles
Brian (a)les
Brian Coles
Brian Coles

9()°36'3S" W, 39'03'57" N led .\ekola

9()°15'33" W. 3S=22'3fi" X |ed Xekola

9l)°iri5" W, .3S'II9'19" X Jed Xekola

89°53'06" W. 3S°5(i'42" X |ed Xekola

9()°H'56" W, 3.S°(lfS'2S" X lelT Xekola

199(i/(V2.3
l99(S/,5/i(l,4

kiiinre 493:4-5

1995/10/12.2
199V6/(i.l

199S/4/19,2

1995/S/5.3

1995/ 1 0/1 3.2

1999/10/24.2
199S,/5/13.i
199.S/5/I6.2
199S/5/31.2

1997/7/12.2
1997/7/17.2
1999/4/4.4

L:\\CB 3.S0.S

89°26d5" W, 37''3(i'(12" X |efT Xekola UW'CB 3S4(i

90°09dS" W, 42 19'59" X John Slapeinsk) FMXll 2S(>S35
90°13'3fi" W. .3.S°54'51" X Jeff Xekola UWCB 431 ]

10
75

X
X

32

X

24

4
13

33

1

2

3

12

42

31

FWCB :5939 25

FWCB 3914 IWCB 39iri 45 42

UWCB 42(i7: IWCB 4269 9 1

1'\\(;B 3S94: t\\(:i5 .3S9fS 43 24

91 = 17' 11" W. 43°27' 12" X Jeff Nekola UWCB 5366: 536S

Paa> 108

THE NAUTILUS, \ol. 115. No. 3

Table I. (."(iiitiiiiicd.

Siti

Location

( .'ollcctor

(Collection nnnihcr

# Mi'asurcd

c;

nificr- G.
sciusis procera

('lai/li)ii Cituntij

Tnikc\ HiNcr Moiuuls
CliiilDii Cntnilii

Ma(|iiiik('t,i Sdntli
DiihiKjiic ( iiiiiilii

Rooscvcll i\(i;iil
Fhii/d ('(Uiiilii

Jnnipcr Hill SIi.lIc Clatlc
Jiuksim (\mnli/

Hamilton (ilailc
\\ iiiiii shirk ( iniiili/

Decorali (Clailc
Missouri
Tdiicij C'imiili/

Hollistcr
Wisconsin
Buffiild CiHiulij

Landfill Koad
Criiufonl ('oiiiili/

Lcitncr Hollow

Rusli ( 'reck

Cniiil ( 'diiiilii
llc\\i'\ lltl'.;lils
Zinnncr

LdCrnssc Ciiiiiili/
I'Apcriiiirnt.il I'arni
I li\on

I'icnv ('imiili/

Ilagcr Cits
Trcnipcfilc/in ('ciiiihi

Rnidy's Hlnir
\rrniiii C.oiuitij

Battle Hlnir

N'icton

9i°02' 1 1" W, 42°42'45" N jelT Nekola I'WCB fi46S

90 :59'()5" W, 42"()1 '12" \ Jell Nekola UWCB fil42

9n°44'3()" W, 43°32'55" N |<.|T Nekola I'WT :B :3rs;5

92°59'02" W. 4:3"(13'l(r \ JelT Nekola UWCB 575

9()°34'()S" W, 42°04'23" N JelT Nekola UWCB 3732

91''46' 10" W, 43°1.S'55" N ji'lT Nekola U\\'(;B 6315

93°L3'41" W, 36°37'()()" N Brian Coles

91='52'45" \V, 44n5'56" N J.inies Tlu-ler KMNIl 2S5717

91°()5'()5" W, 43°13'03" N
9r'()7'54"W, 43°21'56" N

9I°()1'14"\V', 42°44'03" N
91 ()2'50" \V, 42°50'3()" N

9rO()'47" N\; 43°50'12" N
91' J2'(K)"W, 43°49T4" N

92°31'36" W, 44°36'2()" N

|anies Tlieler
James Tlieler

lames Tlieler
lames Thelei'

lames Tlieler
|ames Tlieler

lames Tlieler

91"2.S'59" W. 44=01' 12" N J.imes Iheler

9]°12'3S"\\'. 43°27'36" N
91°12'45"W, 43°29'26" N

|anies Tlieler
lames Tlieli'r

KM Nil 2SW)7(i
KM Mi 2S5.S24

FMNIl 2SfiK31
F.MNH 2S5(>S0

FMNII 2S5fS70
FMNII 2S5761

I'M Ml 2S5920

I'MMI 2S573,0

FMNII 2Sfi049
FMNII 2S5S43

31
18

16

6

8
11

13

8

4

Vm small p()[)iiluti()ns (<4() iniliv uiiials) all iii.itiire,
iindainagc'd siiclls were measured. V\)V lariicr popnla-
tions. a random sample oi'aiiproNiiiuiIeK 45 inidamaf^ed,
adult siiclis wa.s selected. Sliell liei'j;lil and width was
measured in increments of 0.01 mm nsinn a dissectimj;
microscope with a calihrated ocular mii-romeler. Maxi-
mum dimensions were recorded lor each shell. Shell
lieiijht was measured Iroin the lip of llie [iroloeoneh lo

(he liase c)l (lie lip. while sliill width was iiieasiiri'd Iroiii
the l"i'J,ll(-lll( ist lliai'<j;ill 111 llle .ipel'tllle In Hie lell-miisl
iiiare;iii ol tlie lind\ whorl.

Dillereiices iii sliell hei'^hl and wuhli Inr lindi C.ds-
iroroptd new specii's and (.^d.slriH'opId j>roccrd were an-
ak/ed \ia lull 2-wa\ .ANON'.Vs in which ta\on and <j;eo-
'^raphii' location sened as independenl x.irialiles. Be-
cause ol natural ii;ronpiiit^s in iiccurrenee. pi ipiilalions

J. C. Nekola aiul B. F. iU

201)1

Paec 109

wvYc assiij;iu'cl to oiu' ot tliree <j;e(inia[ilii(-' regions: l^a-
It'o/.oic Plateau, Sdiitlit'iii Illinois, ni' O/arks. Diflcrcnccs
in slu'll li('i'j;lit and witltli witliin tlic 9 .sites ot co-ofcur-
renci' wrrc also (lotunu-ntctl nsiiiL;^ hill 2-\\a\ ANO\'As
in wliifli ta\on and sitf seni'il as tlic indc'iii'iidont \aii-
ahk's.

Tlic rt'iitrai tendencies in lliise rrlationsliips were
liiapliitallv represented \ia liox plots. In box plots, tlie
I'entral line I'epiesents the nieilian ol the sample, the
inart^ins ot the ho.\ repre.sent the interijnartile distances,
and the fences represent 1.5 times tin' interi|uartile dis-
tances. For data having a Gaussian distrihution. approx-
iniateK' 99. oC^ ol the data will fall iusiile ol the fences
(\'ellenian antl Hoa^lin, 19S1). Outliers tallin'j; outside ot
the fences are shown with asterisks.

The strength of clinal \ariation in shell heiL:,lit \s. lat-
itude (as e.xpressed in UTM Zone l(i coordinates) was
estimated for both species using least-sijuaies linear re-
gression. UTM coordinates were used to preclude po-
tential bias originating Irom ust- ol polar-cooidiiiate lat-
itude coordinates.

Scanning eleclron microscopy: Scanning i leclKui
micrographs ot Gastrocopta rogcrsoisis and Ga^lrticiipUi
procent, taken with a Hitachi S-24fiON Scanning h'.lec-
tron Microscope m \-SEM Mode (10 Pa; 22 k\) with
a backscatter detector and no. 2 'j;anuna coirection.

Habitat associations: Tlie ph\sical habitat and asso-
ciatetl plant i-oiuniunities were noted during field col-
lection of sites documented li\ the authois, Hiis intoi-
mation was determineil tor otlii'r sites through museum
records and/or the published literature leg., Theler,
1997).

SYSTEMATICS A\D DISCUSSION

FamiK Pnpillidae Turton, 1S31
SubfamiK ( iastiocoptinae Pilsbn. UJlS
Genus Gdstnnoptu W'ollaston. 1S78
Snligeuus Giisl rocoplii Wollaston, 1S7S

Gii.strocoplii /ogcr.sc/i.v/.s ni-w species
(Figures 1-3)

Gastrociipid pron-ra iiicfhin^i Pilshn, ly4S. fiu;iin' |platc| 493:
in part, (ink spcrinuii in ftt^iires 4-5.

Diagnosis: Gustrocopta nt^H-rsfusis is similai" m torin
to Giistnmipta pmccra but is distinguished In the shape
ot the angnlo-parietal lamella. In G /-cgcr.sc/js/.s the an-
gular and parietal kjbes form two discrete, offset, sub-
parallel ridges borne on a rectangular callus (figures 1,
2. 7). In G. procvni tliese tx\'o lobes comergt'. creating
a triangular structure (figures 4, 5, S).

Description: Shell elongate-oxoid with a weakK
conical s[iire, brown, weakK' striate, 1.77-2.5S nun tall
(mean = 2.11 mm) X ().S1-|.05 mm wide (mean =
0.92 mm); fi whorls, the last 2 of approximatt4\' equal
width; sntiiri' pronounced. The aperture is elongate
and rounded with a non-continuous peiistome, Tlie

wcakl\ reflected lip is ol lighter color iIkiii the rest of
the shell, .mil is stiiin'_;l\ reinforced with a sliallf)w sul-
cus immedialeK bclimcl 'file a]iertiire has 5 lamellae,
file ;ui<,;nIo-pariiial consists of two discrete, approxi-
inatelx straight, siili-parallel lamellae borne on a rect-
angular cillns with the angular portion oritjinating
near the iimclion ol lip ami bod\ whorl. The colii-
niellar lamella is bilobed. the upper prcniiinent and
the lowii nodiifii I he upper p;il;it;il l.imella is short
and p];ice(l 111 liiini oj tlii' aii'j,ulo-parietal. The lower
palatal is long and deepK inserted behind the angnlo-
parietal. rile basal lamelhi is short, colnmnai'. and in-
serted ill front of the ;iii'j;iili)-paiiel,il.

Type material: ltoIot\pe ifignres t. 3). 1-\\1,\11

29fia51, 2.1b mm lentil X 0.96 mm widtli, Jeff Nekola
leg. 11 Jul. 199.S; Paratxpes: FMNII 296657, 20 speci-
mens collected with the holot\pe at Fults Ilill Prairie
Nature Presenc, JetT Nekola leg,; Florida Museum of
Natural Histon 2S5352, 5 specimens collected with the
liolot\pe at Fiilts Hill l^rairie Nature Presence, [olm
Shipcinsk-x- leg.; FMNII 29655S, 10 specimens, Galico
Hock West, Izard Conutx, Arkansas (92"8'55" W, 36°7'I"
N), 15rian Coles leg; FMNH 296559, 10 ,specimens.
Sales\ille, Baxter Countw Arkansas (92°16'52" W,
3fi°13'2" N), Brian Coles leg.; F.MNH 296660. 10 .spec-
imens, Prairie clu Rocher, Randolph Coimt\, Illinois
(90°1'56" W, .3.S°6'2S" N), Jeff Nekola leg.- F.MNH
296661, 10 speciinens, Maquoketa South CJlade. Clinton
Conntx, Iowa (9(n9'5" W, 42°ri2" N), Jeff Nekola leg.
FMNH 2S5730. b) specmieiis. lir;i(l\'s Bluff Trenipe-
;ile;in (.loiintw Wisconsin |9P2S'59" W. 4t 1'12" N).
billies Tlieler le',^,

Txpe localit\: Fnlts Ilill Prairie N.itnre Presenc
(90-11 '1.5" \\. 3S"9'19" N), Monroe County, Illinois,
l'S.\, approximatcK .'^^ km SI-" of I'iilts along Bluff Road;
on dix limestone ouli.idps. imdei bed Cedar [Jiiuipenis
iir<ii)ii(lU(i L.' ;it tlie crest of a bluff. We selected this
localit\ as it is cenlralK located within the known r;mge
ol Gdslrocopid I'i'iU'rsi'W^is. ;iiid exists within a ]irotected
iiatur:Ll itrc'a. .AdditioiialK, the loralitv is included in part
of the nmij;e ol ( .' fiiiur\tii\is where the new specit'S
occurs s\ nipatricalK with Giislrocojihi pi'iicfni

Et\'molog\: 'flie specific name honors the town of
Rogers, .Ark;iiis;is. 'fliis is the collection location tor the
specimen upon which the first published illustration was
b;ised. F\cn though we luixe been iinabli' to relocate
them in Rogers, extant po|iiil;itioiis ;u'e known within 40
km ;it Withrow Springs St;ite Park ;iiid l^ea\cr Dam.

Subgeneric allocation: 'I'lie ;ipproximatel\ straight
angnlo-parietal lobes ;is seen in basal \iew, colnimiar
basal lamella, and brown shell color indicate that fir/.s-
ln>n)j)ta rogcr.vri/.v/.s belongs in tlie subgenus Gasf rorop-
lii SuperficialK, its ;mgnlo-p;niet;il lamella r(>seinbles
that of sexeral species in Gaslrocopta subgenus Iinuicr-
sidciis. notabK Gastrocoptd InlinncUala (Sterki and
Clapp, 1909) and Gdsl nx-opld ddUiaiid (Sterki, 189S'
(PilsbiT, 194S; fitiure 490; 1-4'. However, members ' i

Paa- 110

THE NAUTILUS, \ol. 115. No. 3

Ozark Populations

Southern Illinois Populations

Paleozoic Plateau Populations

f
2

1 1
T

275

-

250

^

.

225

-1

*

200

1 7C,

J

*

1.15

-

-

108

i

101

*

-

I

in

0S4
087

■ ^

-

i

-

•

I

r2 -0600

Taxon p < 00005

Locabon [) < 00006

Taxon X Location
Injeractiwt p < 00006

r2 -0602

Taxon p < 0D0O5

Location p < 00005

Taxon x Localicn
InteracBon p < 00005

Figure 9. Box-plot (liai;r;iiii ol \;iriatioii in dislrocopid n<!H-rscii\i\ and (.".asl rocitjtiii jiroccni slicll licit^lit ami width wit
Ozarks. soiillii-rii Illinois, and I'ali-ozoic I'lalcan, Slatistkal results arc liascd on a 2-\\a\ A\()\.\ witli iiitt'iaetioii.

tilis suhgeuus are cluu-aeterized In lia\iiiif angular aiitl
parie-tai lobes that are curved or bent at their distal ends,
an elongate basal lamella whose long axis is parallel to
the lip, and a clear to white shell color.

Moipln)mt'trics: Shells lidin 414 Castnu-opta n)<ui--
semis (146 from Paleozoic Plateau populations, 121 rnnii
southern Illinois, 147 from the Ozarks), and 343 Cw;.s-
troropld priH-cra (88 from Paleozoic Plateau populations,
177 irom southern Illinois, 78 irom the Ozarks) were
measured. Ciomparison of these demonstrated that C
rogersensis averaged 2.1 I nun in luiglil whereas G. /;/"(<-
cera averaged 2.40 mm iligure 9). This difference was
highly signilicant (l'<().{)0()5). .\dditiouall\-. the dilfer-
ence in height l)etweeu Ozark and southern llliiidis |i(i|)-
ulations was greater than for Paleozoic Plateau inipula-
tions (P<0.0005). Similar trends were noted in shell
width (Figiu-e 3), with C ri>gcrsensi.s averaging 0.92 nun
and C prorcni averaging 1.02 mm. This dilTerence was
highly significant (P<0.00()5) and also varied between
population centers (P<0.()()()5), with niaviiiniiii diver-
gence occurring in tlie Ozarks.

One-hnndred and forty three of thi' measmed (xV;.s-
trocoptd ro'^crscnsis and 113 of tlie measured Gtislro-
copta proccra shells originated from sites of co-occnr-
rence. When analvses were limited to these stations.

higlilv significant differences (P<0.0005) were still noted
in both slu'll lieight and width (figure 10). .\dditionallv.
a liighlv significant interaction bet\veen site and shell
height (!'• 41.0005) and width (P = 0.002) was noted,
with maximimi tlivergenci' occurring betwei'ii the two
species in (he Ozarks.

Liiu'ar regression ot slu'll height vs. L'T.M \-S coor-
dinates (table 2) demonstrated that while Ga.strocopla
procera shell height stronglv (r- = 0.375^ and signifi-
cantlv (P<().0005) incieased towarils the south, no clinal
variation occurred in Gastrocopta rogcr.scii.v/.s (P =
().87(i; 1- = 0.000), Because of this, differences in shell
size are less marketl between C^ n><icr\cii\is and Grt.v-
trocopta procfi'd in the iiortii. It is not clear w hv these
taxa lespond dillerentlv to iilentical env iroimiental gra-

(Ill'lltS.

Cieograpliio distribulion: The 30 known stations for
C.iisl rocoplii i(i'j_rrsi'ii\is apjiear restricteil to three dis-
tribution eeiilers: (he ( )zark iiplilt ot northern Arkansas
and southern Missouri, southwestern Illinois, and the
i'aleozoie i'laleaii or Driflless Area": see Prior, 1991)
ol northeastern Iowa, northwestern Illinois, and south-
western Wisconsin (figure 111. The majoritv ol known
sites in the ( )zarks are restricted to limestone bliills near
the ii|)per western I Hentoii. ( .'arroll and Madison conn-

C. Nekola and B. F. C:()les. 2001

Paee 111

Shell Height

Shell Width

-p 250

225

2,00

*
+
*

* .

*

* I 1

rog©r3 (Tocora

Taxon

r2 - 0,690 :

Taxori p < 00005

Site p < 00005

Taxon x Site
Interaction p < 00005

Taxon

r2 - 0,619

Taxoa p < 00005

Site p < 00005

Taxon x Site
Interactiorv p = 0002

Figure 10. Box-plot diiiijriim of \aiiation in C.dstnx-opta ro^erscnsi.s and Gastroiu^jthi proccru slicll litijit .ind width williin site:
ol co-occiirifiice. Statistical results are based on u 2-\\a\ ANOVA with interaction.

ties) and eastern (Baxter, Izard, and Stone counties)
U'liite Ri\'er and its trilmtaries. In sontliwestcrii Illinois,
C. roocrscnsis is limited to a fiO km extent ol limestone
lilnffs alone; the Mississippi Ri\"er in Kandolpli and .Mon-
roe comities. Tlie Paleozoic Platean populations lie with-
in 50 km ot the Mississippi Ri\er in |o Daxdess Countx
(Illinois), Allamakee, Clinton, Dnl)U(|ue, and Jackson
counties (Iowa), and Chawtortl, (wanil LaCrosse, Trem-
pealeau, and Vernon counties (Wisconsin). The localized
distribution of G. Wiierscnsis contrasts markedK' with
Gastrocopta proccra jjrocera, which is widespread
throughout nnich ot the eastern and midwestern USA
(Hubricht, 19S.5).

Even thougli we ha\'e documented land snails at o\-er
700 sites in tlie region, as neither our own collections
(nor those museum collections that we ha\e examined)
fnlK- co\er this landscape, we cannot uni"(jui\()call\' state
tliat G. ro<s,crscnsis is limited to onlv these tliree distri-
butional centers. This is paiticularK' true in tlie soutli,
where undercollection in southern Missouri ma\' well ac-
count tor the apparent disjunction between Ozark and
southern Illinois populations.

The doiniiiant distribution tor localized midwi'stern

Table 2. Siininian' statistics for regression ot sliell liei'Jit \s.
UTM X-S location for Gastrocopta roiicrsciisis vs. Giislnuoptu
proccra. The best-fit slope is calculated as tlie iinii cliansje in
sliell lieit;lit per 10" meters.

Species

G ni'^ersciisis

G Procira

11

415

.34:3

slope

-O.OO.j

-().:567

standard error

0.021

0.026

t

-0.1.56

- 14.2S9

p (2-tailed)

0.876

<().()( 105

r-

0.000

0.375

USA PoKgxridae and Zonitidae species (Huhriclit, 19S5:
Emberton, 199.5) is t\picall\' centered on the north-
western Arkansas (^zarks [e.g., luflfcliihiis eclentaliis
(Sampson, 1SS9), Paracitiva siiitpsoiii ( PilsbiA', 18S9),
Stciuitirirui hihnisniit (Bland, 1S62), and Vctitridciis
liritisi (Pilsbn, 1S92)]. Howe\er, other localized inid-
western distributions also exist [e,g.. Discus iiiacrUiitticki
(F.C. Baker, 1928), Eiwhcmotrcma hnhriclili 1, Filsbn.
1940), and Tiinilopsis discoidca (PilsliiA", 1904,)). G«.s7;v)-
copta roiicrscusiK possesses one of these latter patterns,
being limited to the Ozarks and die Paleozoic Plateau.
Its range is most closeh' matched b\- that oi Vertigo iiwr-
(iincccusis WiuDevender, 1979 which exhibits an almost
identical distribution (Frest and Fay, 19S1: Hubricht,
19S5: Fre.st, 1991: author's unpublished data!. Both of
these regions are nnderlain b\ carbonate bedrock, lia\e
a I'ugged terrain, and escaped glaciation during the Wis-
consinan. llnlike G, r(i<iiTscnsi.s. howexer, \' iinr/iiiic-
cciis/s is restiicted to cool, mesie carbonate ciitis if rest,
1991).

Taxononiic remarks: l'ilsbr\ il94Si dilTereiitiatcd
liiiiiis and subspecies in the GdslrDCupln pmn'ra group
based priinariK- on the degree of separation ot the an-
gular and parietal portions ol the angulo-parietal lamella,
grading from (Idstrocopta pruccni loriii riparia (least
separated) thi'oiigh Castnunpta pruani prorrra and
GastrocDjitd pnu-rni stcrkidiin to Gdsiyoropld pwccra
)uccJtiniii (most .separated). Gdstnicoptd p. niccliiii<j,i was
also thought to differ trom the nominate species bv lie-
ing sliorter and lia\lng a thicker and more convex lip
(Baker, 19.39; Franzen ami Leonard. 194.5: Pilsbn,
194S). Subsecjuent workers ha\e considered G, j>.
iincluii<j,! as simpK a variant of G. procvra (Leonard.
19.59: Hubricht. 19S5). Hubricht (1977; 1985) differen-
ti;ited the remaining taxa as distinct species ha.sed pri-
nuirilv on the position and slope ol the lower i.)ai.ii.''

Page 112

THE NAUTILUS. \'(i

15. No. 3

Figiii'o I I. I )istnliiitu)ii c]l Ciisliin oplii roiuisrnsis.

tootli, ranging from C. prorcra (nio.st dci'pK- inserted)
tlirougli G. slcrkiaua to C;. r'qxirid (least deepK inserted)
hut failed to gi\e adecjnate reasoning lor tiiese eoneln-
sions. We lia\c Found tliese features to lie liigliK \ari-
ahle, witli continuous xariation ap]iai(iilK exisliug liotli
within and l)et\\cen po])ulations. i''urtliei' nioqihonietric
and taxononiic iincstigations will he neeessan to help
determine the appropriate t;L\ononiie categories within
this group.

Despite continued ainhignitv over llie slaliis of lorins
within the Guslrocopid prorcra complex. Gti.slrocojila
rofyrscnsis new species is distinct in all asjiects. It differs
most conspicuousK In ha\ing the aiigul.ii' jioition ol the
angnlo-parietal lamella arising near the lip and rinming

paralli'l and separate Iroin the more deepK set jiarietal
portion. While the angular and parietal portions show a
xariahle degree of tlistal separation in G. prorcra. in all
cases tlie\ fuse at approximateh' mid-length. .Svmpatric
populations of (I. ro<j^crscnsi.s and G. prorcra do not in-
tergrade in tliis feature. Additionallv, C. ro^ersensis in-
dixidnals appear consistentK smaller (ca. 0.25 mm) than
G. prorcra. Less striking differences inchnle the distinct-
ly reflexid lip of G, rogersensis as seen in profile (figure
3), antl its somewhat longer and more deepK' inserted
lowi'r palatal lamella.

Based on these criteria, examination of the holotxpe
of Gaslroropiu prorcra mccliingi (USN.M 22(i395, figure
fi), and the ANSP figured specimen of G. p. incclungi
iVoni South Dakota (Pilsbiy, 1948. fig. 493: 1-3) rexealed
dial lioth fell well within the normal range of xariation
lor the Ga.stroropta prorcra complex. Thus, we concur
with Leonard (1959) and Huhricht ( 19S5i that this t;L\on
should he regarded as a SMion\ni of G. prorcra. How-
ever, the figured ANSP specimen of G. p. larrhiugi from
Rogers. Arkansas (Pilshn', 194S, fig. 493: 4-5) conforms
ill all respects to Ga.stroropta rogersensis. The reasons
lor the oM'riooking of the uniqueness of this specimen
In PilshiA are likeK tx\o-fold: first, the specimen fell
within his concept of G. p. mcrluiigi as it has a \en'
pronounced separation between the angnlo-parietal
lohes. Si'cond, as no other G. rogrrscii.sis specimens exist
in the ANSP collections, there was onl\' limited oppor-
tunitx' for him to obsene the other consistent differences
that e.xist between it and G. prorrra.

Because of this confusion, Ga.stroropta rogcr.scn.sLs h;is
remainetl undescribed even though .specimens have hke-
l\ existed in collections for oxer 60 X'cars. Baker (1939)
referred to Ga.stroropta prorcra mrrlungi from Illinois,
xvithin the known range of G. rogcnseihsis. His draxvings
of this taxon appear similar to G. rogcrscn.si.s. but are too
crude for definitive confirmation. Hutchison (1989) list-
etl G. p. incclinigi from Fountain Bluff in Jackson C^oun-
tv Illinois, approximateh' 35 km to the SI'" of the south-
eiii-most known Illinois G. rog('rscnsi\ pcipnialion. Ex'en
though wf ha\i- not been able to exannne tliese speci-
mens, the reported location and habitat make it likeK'
thai these also represent G. rogcrseii.si.s. Tlieler (1997)
enconiil<'red (.' ni^r/'sc/is/s in his sun'evof xvestern Wis-
consin bedrock glade land snail faunas, but identified all
iiidi\ iiliKils as (.'. proccra.

PrererrctI baliitats: \ll known iiojiulations ol Gastro-
I'opta rogrr\cn\t\ are liiiiiti'd to xerie or dn-mesic cal-
careous rock (iiilcrops. ( )zaik and soiUlieiii Illinois pop-
ulations were encountered on wooded cliffs or did
cresls, as at the Ixjie location. The i^aleo/.oic Plateau
popukilioiis are restricted to bedrock glades (see Tlieler.
I!)!)7 lor images of the BiacK' Bluff site). In these liah-
ilals, individuals are liiiiiled lo organic aifumiilations in
xerie, s[)arsel\ vegetated iiiicrosites, where short-statu-
red grasses and forbs such as Agoscris rii.spidala (Pursli.)
Kaf. .\uilropogoii sropariu.s Michx., Ai-iciiicsia raiidalii
\iielix,, ('arcx alxlita i^ickn.. Carrx ricliaid.soiiii B.Br.

1. C. Nekola and B. F. Coles. 2001

Page 113

Cdstlllcja .scs\ililli>r(i I'lirsli. Miililiiihci'tiiii ruspuldla
(Nutt.) R\(lli.. and \iol(i i>riliilii \ .. (xciir.

in the !\-\ic\\ c)l occupied names lor ol<l-\\<irld C'.ii.sfro-

tOjlltl.

CONSERX'ATION RECOMMENDATIONS

Because ol its limitctl geot^raphic raugc and liahit.il spcc-
ificih. Gdstroccplii /■o^crvciisis sliouUl he considcicd ol
cousenation importance. K\t'u within its distrihutional
centers, populations are sporadic anil tend to hi' limited
to hiti;li-(|ualit\ sites. Ironicallw a lurtlier conieiii loi' its
continued sunixal is the occurrence ol uian\ populations
(includinif the t\pe and most oi the Wisconsin stations)
within nianam'tl nature resenes. Prairie anil hedrock
glade habitats throughout the midwest U.S.A are geuer-
all\- sulijected to intense fire management, as it has heen
l)elie\ed that reintroduction of fire will increasi' tlie vigor
and di\ersit\' of tlie grassland biota (e.g. Curtis, 1959:
Paulv. 19S5i. Becau.se of this. Theler (1997) stated that
fire management is essential to maintain the habitats
usi'd b\- C, roi^crficusis in Wisconsin. However, fire is
known to be deleterious to main natixi- prairie plant
(Hill and Piatt. 1975) and arthropod (Swengel. 199H;
Haqier et. al. 2000,) species. Such negative impacts likelv
also exist for G. mucrscusis. as individuals icside in
tliatcli and organic duff layers that are removed through
lepeated fire episodes. As reeoveiv of these lavers takes
over a dozen years in xeric grasslands (Gibson and Hul-
bert, 19S7), the frequent use of fire management (<10
year return intenals) mav limit the amount ol appin-
priate habitat and significantly reduce the size of G. n>-
^crsen.sis populations. Suggestions of this can be seen in
Theler (1997), as the most frequently burned sites (e.g.
Rush Creek) also have the smallest G. rogcrsois/v and
G. procera populations. The number of recovered shells
per unit volume of soil litter in these managed Wisconsin
sites is 2-3 times smaller than that obsenetl in nearbv
inibumed Iowa sites (e.g. Maquoketa South and Roo-
sevelt Road). Such trends should not be suqirising. as
land snails are highly sensitive to fire (Frest and Johan-
nes, 1995). We have obsened 50'% reductions in rich-
ness and order-ot-maguitude reductions in abundance of
land snails between ailjacent munanageil and fire-uian-
aged grasslanils in northwestern Minnesota (author's uu-
publisheil data). Thus, ovenise of fire management In
consenation groups may pose as great ol a threat to the
surAiv al of G. ro'^crscnsis as habitat loss.

ACKNOWLEDGMENTS

fo Sutto was institmiental in the measurement of Gas-
twvoptd roocrsensis and Gastrocopta proccni shells.
Matt Barthel was responsible for the SEM imaging of
specimens, l^r. Robert Wise and the Universih' of \\is-
consin-Oshkosh kindly allowed use of their SEM facili-
ties. The malacological cmation staff at FMNH (espe-
cially Jochen Gerber and Margaret Baker), ANSP, and
NMNH (especially Warren Blow) kindlv provided access
to specimens, facilitated loans, and pnniiled specimen
numbers. Peter Mordan and Robert Cameron assisted

LITERATURE CITED

li.iker. F. C. 1939. Ficklliook of Illinois land snails. Illinois .Nat-
ural liistorv Suncy Manual 2. Illinois Xalinal llistorv
Suncv Division, llrhaiia, l(i(i [)[>.

Cairtis, J. T. 1959. i'lic Wuclallon of Wisconsin. I nivcrsitvol'
Wisconsin Press. Madison. (m7 pp.

I'.iiilierloii, K ( ', HiT) \\ li, it shells do not tell: 145 million vears
ol evolulion in \iiilli Aiiicrica's poKsrvriil land snails, with
a revision and ioiiscia.iIioii pnoiilies. .Malacolo<iia 37: 69-
110.

I'ian/en. li. S.. :iii(l .\. IV l.eoii.inl H) 13 I lie iHollilsca ol tiic
Wakarnsa river vallev. Liiiversitv ol k.ins:is Siiencc Bul-
letin 29: 36.3-439.

Frest. T. J. 19VJ1. Sunniiiin st:itns reports on eight species oi
candidate lanil sn;iils Iroin the Driltless .\re:i I l':ile()/.oic
Plateau), Upper .Midwi'sl l'iiL:il Ke])ort. (^oiitniet #301-
01366. USFWS Heuioii :V Ft. Snellmg. 5-4 pp.

Frest, T. j. :ind F. 1' l'":iv, U)S1 \(7Yig(» mcrameccnsis (Pul-
nionata: Pupillidae) Ironi the \iat;ar;iii I'"se:upnieiil. Iowa.
The Nautilus 95: 33-37.

Frest, T. |. and E. |. Johannes. 1995. Interior Cloliiinhia Basin
iiiollnsk species of special concern. Final Report, (.Contract
#43-OE()0-4-9112, Interior Columbia Basin Feosystem
Management Project, Walla Walla, 274 pp.

Cabson, D.]. and L. C. Ilnlhert. 19S7. Effects of fire, topog-
raphv. and vear-to-vear ilimatii- variation on species com-
position in tallgrass prairie. N'egetatio 72: 17.5-185,

Hanna, G. D. and E. C. Joimston. 1913. A Pleistocene mol-
luscan fauna from Phillips (^ounh. Kansas. Kans;« Uni-
versity Science Bulletin 7: 111-123.

Harper, M. G.. C. H. Dietrich, R. L. Larimore and P A.
Tessene. 2000. Effects ol prescribed fire on prairie .Ar-
thropods: an enclosure stndv. Natural .\reas [onrnal:
32.5-335.

Hill. G. R. and W. f Piatt. 1975. Some effects of fire upon
a tallgrass prairie plant commimitv in nortlivvestcni
Iowa. In: Wall. iM. K (ed), Prairie: a Multiple \ievv.
University of North Dakota Press. Grand Forks, jip,
103- 11.3'

Hubricht, L. 1977. Thirteen new species of land snails Irom
the southeastern United States with note on other spe-
cies. Malacological Review 10: 37-52.

Hnhriclit, L. 19.S5. The distributions of the native land mol-
lusks of the eastern United States. Fieldiana. new series,
24: 1-191,

llntcliis(}ii. M, D. 19S9. .\ snni-v of land snails in southern
Illinois. FiiKil report. Noiii;:mie \\ildlil<- (."onservation
Fund. Illinois Dep:u1ment ot(:ons<avatioii. N;itmal Her-
itage Division, Springfield. 21 pp.

Leonard, A. B. 19.59. Handhook of <iastro])ods in Kansas. Uni-
versitv of Kansas .Museum of .Natural Ilistoiy .Miscella-
neous Publication 20. l,:ivvrence. 224 pp.

Nekola, J. C. 1999. Terrestrial gastropod richness ol'ciulxiiiate
cliff and associated habitats in the Great Lakes region of
North .\iiierica. Malacologia 41: 231-252.

r;iiil\, W- R, 19S5. How to manage small prairie fires, Dane
Gounh Environmental (lonncil, .Madison. 30 pp.

Pilshtv, H.A. 194S. Land Molliise:i of Norlli America (North
(if Mexico). Aeademv of Natural Sciences of Philadelphia.
.Monograph 2(2): .521-1113.

Page 114

THE NAUTILUS, Vol. 115, No. 3

Prior, J. C. 1991. LiUulforTiis ol Uma. l'iii\crsit\ (il lima Pifss,

Iowa Cit\. 153 pp.
SwcnijC'I, A. B. 199fi. l\H('C'ts ol fiiv and lia\ iiianageiiK'tit on

the ahuiKlancc of prairie hiittcrliics. Biolo<;ical Con.ser-

\ation 76: 7.'3-S.5,
Tlieler, |. L. 1997. Tlic inodcni terrestrial ga.stropod (land

snaill fauna of western Wisconsin's hill prairies. The Xan-
tilus 110: 111-121.
X'ellenian, I'. F. and D.C. Hoa'^lin. I9S1. .Ajiplications. ba.sics,
and eoniputini; ol exploraton data anaKsis. .■\dtlisoii-\\'cs-
le\ Press. Reailint;, .354 pp.

Book Review

Catalo'^ue and Bihlio^raphtj of the Marine
Shell-hearing MoUusea of Japan. Type
Figures.

Sltiuilrlii Hi'^o. Paul Cnlloiiion. and Yosliiliiro Goto.
2001. Elle Scientific Publications, O-saka, 208 pp.
$325.00.

Tlii.s important wcirk is now complete with the pub-
lication of the volume of figures. The te.xt volume v\as
reviewed in these pages i Petit. 1999) shortK' after its
publication. The book is mii(|ue in that all figured spec-
imens are name-bearing t\pes. As the authors state in
the introduction, it became evident during the produc-
tion of the te.\t \olume that the identity of many species
was uncertain. .\s onlv t\pe specimens are figured, read-
ers ma\ be assured that the figures match the names.

Almost 2()()() t\pe specimens are figured in color. Each
page of figures comprises 16 excellent, informative illus-
trations. The arrangement of this data is, of coursi', ex-
plained in the introduction that should be read before
the !)ook is used. Data is arranged in five areas, the most
important being the eiiti'v munber that corresponds to
the species number in the text volume. The suffix "s" on
a number indicates that the name of the tvpc fignrt'd is
treated as a jmiior svnonvm in the text voliuue. .'\s an
example, there is a figure of the holotvpe ol Zeifra iiii-
trifonnis A. .^dams, 1860, and also three figm^es of tvpe
species of nomina that appeal' in most literature as svn-
onyms of Z. milrifdnitis: Z. zomiln ((^onld. IS6()i, /. ral-
klicosta (Habe, 1960), and Z. siibiilrca (E. .\. Smith.
1879). When viewed together il is clear that all loui' are
not conspecific. On tiii' other hand, the figures ol Ihtc-
cinuin midori Ilabe and Ito, 196.5. and B iirdcinnluiii
Dall. 1907, appear to represent the same species. Tlicre
are numerous examiiles to be found ol both cases, 4'his
book dramatically illustrates the importance of having
figiu'es of type specimens.

The second bit of data is a scale bar used for speci-
mens mider 10 nnn, and the third is a measurement in
millimeters for larger specimens. Of little interest to the
casual user is the fourth l)il of information that is an

indication useil if the figure is a composite image. .^
composite image mav be used for a shell whose shape
precludes bringing the entire specimen into good focus
at once, in which case two photographs are taken and
combined.

The final liit of data is termed the "main data box"
and contains the specific name and generic placement
used in the text volume; the autlinr and date of publi-
cation: tvpe status: museum name ami registration num-
ber. The rules for designation of tvpe specimens have
alwavs been somewhat confusing and were recently
made even more so bv changes in the fourth edition of
the ICZN. The authors list the various kinds of types
recognized, the acronvms for which are used in the
"main data box," and give a description of each.

The only negative aspect of this production is the fact
that onlv some of the taxa listed in the text volume are
represented bv figures of txpe specimens. As explained
bv the authors, prioritv was given to species originallv
described from [apan as their limited resources preclud-
ed inclusion of all species. Manv well-known Japanese
species originallv described from elsewhere bv the ear-
lier authors (Linnaeus. C.melin, Lamarck, and others)
are therefore absent.

An appendix lists adtlitional species names for the
Catalogue, errata for the Catalogue listings, errata and
additions to the bibliographv, and additional and emend-
ed Japanese names. There is a complete index. .\s with
the first volume, tliis book of figiu'es is superblv pro-
tliiced. It is .\4 in size, the text printed on cream stock
papei- and the plates on lieavv glos.sy stock, and bound
in n;uiiascene cloth with a gold-stamped leatherette
spine. .\s is standard in Japan, it comes in a slipcase.

LlTi: H.VTURE CITED

I'elil K. K, 1999. Book review: Catalogue and bibliography ol
the marine shell-liearing Molhusca of Japan. The .\antilus
113: 102.

Hieliard I,. IVIit

l'( ) liii\ ,i<l

\orih Mvrtle Heaeh. SC 29.597 US.\

r.e,pe(it("all,nel

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rHE NAUTILUS

Volume 115, Number 4
December 18, 2001
ISSN 0028-1344

A quarterly devoted
to malacology.

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EDITOR EMERITUS

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National Museum of
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Field Museum of
Natural Histon'
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North Carolina State Museum of

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Laboratoire de Biologie des
Invertebres Marins et Malacologie
Museum National dUistoire Naturelle
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College of Charleston
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F"lorida Museum of Natural lli-sini-v
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History
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THE€^ NAUTILUS

CONTENTS

Voluinr 1 13. Siinihcr I

Drccnihcr IS. 2001

ISSX 0028-1344

D;nifl C. Reid New data on tlic taxonoiiis ami distriliiitioii of tlic ijciiiis

iMormiii (iiilfitli .iiid I'id^con, 1S34 ( Castropoda:
I,ittnriiildaf in liidci-Wcst Pacific mangrove lorcsts 115

BaiTA Rolh Pli\l()'j;cn\ nl pncuiiuistdinal an-a ini)rpl[c)luti;\ in tcncstrial

Pulnionata ((Gastropoda) 140

Richard Diierr Ciijimcaissis chipt'liiiid. ,t new s[iccics i ( ;asli-o|)oda:

(.';Lssiilae I from the Mioi-ciic t!liipola I'^onnalion ot
nortli\vfst(.'ni Florida 147

Note

Francisco J. Garcia Correttioii ot a mistake introduced in tlic description ol an

Jesus S. Troncoso aeolid nndilirancli ol tlie ij;eniis Favoiinns V,\a\. 1S5() 150

Book Review 151

STATEMENT OF OWNEIiSl 1 1 1' MANAGEMENT AND CIRCLILATION

I'nl.licalioii Title. THE NAUTILUS

I'lililication No.. ()()2S-l:344

Filiiit; Date, N(neiiil)er 14. 2001

Is.sue Fre(|ueiR'\'. Quarterl\

No. ol l.ssiifs I'lililislicd .\imiialK, i'Our

.\iii)ual SiihscTipticiii Price, US $5fi.00

(Joniplt'tf .\Iailinij; .Address ol Known Olfiet' ol I'lililieation, 3073 Saniliel-( !apti\a iload. Sanihel, PL 33957.

Complete .Mailinij; .\dtlress ol Ileadtjiiarters. same as 7.

Full Names and C^omplete Mailiiii; Aildresses of Fulilisher. Tlie Baile\-Mattlie\\s Sliell Museum. 3075 Sanibel-

Captixa Road, Sanihel, FL 33957

I'xlitor, Dr. [ose H. Leal, address as aho\e.

Ma)ia<j;ing Editor, (Christina Petrikas, acklress as aho\e.
10. Owner, Shell Museum and Educational Foundation, Inc.. address as above.
1 I, Known Biindli<iklers, Mortgagees, and Other Seciuit\ Ilolilers Owning or Holding 1 Percent or More ol Total

.\inount ol Bonds. Mortgages, or Other Securities, None.

12. The puipose, hmction, and nonprofit status ol tliis organi/ation and die tax exempt status lor iederal income
tax puq^oses lias not changed during the pit-ceding 12 mouths.

13. Publication Title. THE NAUTILUS

14. Issue Date lor Circulation Data Below, October 31, 2001

.\\erage Single

15. Extent and Nature of Circulation 12 months Issue

A. Total Number of Copies 543 540

R. I^aid (Circulation

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THE NAUTILUS 1 !5i4 il 15-139. 2()()1

Page 1 15

thr u;cMnis

New data on the taxononn and distribution ol

Littoraha Griffidi and Pidijjeon, 1S34 (Gastropoda: Littorinidae)

in Indo-West Pacific nianeroxc forests

Da\id C;. Reid

Di-partiiiciit cil Zooloijx
Till" Natural llistnn Miisruin
Londdii SWT -inn IK
dgrteiiliiii.ac.iik

ABSTRACT

This papc'i' adds iiiw data cm niaimi()\c-asM)(iati-d spcrics ol
Littomrii: in tiic liidi>-\\Vst Pai-ifk- ifjoii thai lia\c arcuiiiii-
lated since tlif l.iMinomic rt'vision cil tlic i^ronp In Kcid
(19S6a). One new speeies is deserilicd lidiii llie nortiieastern
iniliaii Oeean and L. -sinensis iPliilippi, IStT is dislinijnislied
from (lie similar I., aiiiriihila ( Pliilippi. IS4(i ' with \\ hieh il \\as
pie\ionsl\ ennhiscd. Siij;nifieant extensions to the distribution
ranges ol 12 speeies are repoited The penis, paraspenuatozoa
and radnla ul' I,, ili-licdliiln (\e\ill. 1SS5' are deserihed I'orllie
first time and additional inlonnation i;i\en on the geographieal
variation ot />. filos/i (So\\erb\'. LS.32' in Austialia. New htera-
tnrt- I post- 1985) on the entire genus IJilortirin is Imelh re-
\ie\\etl.

IXTHODIC TION

The hiiiilK' Littorinidae is aiiinii'j; llie iikisI well stiuliecl
o( marine gastropod groups. loi- it is ol woildwide dis-
trilmtioii aiul its nicnihers are iisiialK alinndaiit, aeees-
sil)le anil easlK eollecteil in the intertidal /.oni' (see ri'-
\ie\vs h\- Me(,)nai(l. 1996a, b). Ifntil some 30 vears ago
the gronp was noted lor a eonliised ta\onom\ based
mainl\- on bigliK \ariable shell eharaeters. However,
sinee then tlie applieation ol teelnni|nes ol line anatoinv,
scanning I'leetron mieroseopx and moleenlar biolog\ has
resnited in refinement. Within the most well known sni)-
lamiK, tlii' intei-tidal and mono|)h\letie Liltorininae
(Reid. 19S9a), recent stndies recogmse 1'59 speeies in
eight genera: 1 MclaHiiiphc iHosewater, 19S1; Keid.
l9S9a): 11 Pcasirlla (Reid, 19S9!): iieitl and .\!ak. 199Si.
2 Maiuwari)iiiia (Reid. 19S(Sbi, 10 'rvclahus (iiosewater.
1972; Reid and Geller. 1997). 1 Cvnrhriiis i Rosewater.
1972: Reid, 1989a>. 39 Littomria i Keid. UJSfia. 1999a.
b. this studv). 19 Littoriua (Reid, I99fil and 5(i Xoililit-
foriiui (Bandel and Kadolsk-A. J9S2: Keid. 19S9a, in i)re.ss
a. 1)1. For the snblamiK Littorininae there is at least a
prelimiiian pli\ logen\ of the genera (Keid. 19S9al anti
S(jme species-le\el ph\logeni(>s (e.g. iieid et al.. 199fi;
Reid, 1999h), and fnitlier moleenlar phylogenies are

now beiiiL; ])rodnced, I'lir <ill these l.)9 species the re-
prodiieti\c moile and habitat are known, and lor almost
all ol them the geograjibical distribution is recorded in
detail. This, therelore, rejiiesents a remarkable dataset
among marine invertebrates: a lelaliveK well-worked
monoplnletie gronp ol woildwide i but predomiiiantk
liopieal) distribution. Once a more robust plnlogenetic
hamework is achieved it should become a model svstem
h)r the studv ol m:ieioevolntionan proces,ses (e.g. IJt-
loriiKL Keid et al.. 199fi). .Meanwhile, it is important to
understand the svstematies ol these speeies accurateK
and to record their distributions as preciseK as possible.

The modei'U concept ol the genus Lillcnirid as a
mouophvletic grou]) was estai)lislied b\ Handel and Ka-
dolskv 1 19821 and Reid (l9Sfia. 19.S9a): JH-fore this the
species were generalK included in IJIIohiw. a large
poKpliv letic as.seiublage. The i^enns is almost exciiisi\el\'
tropical in distribution and the highest diversilv is found
in the Indo-W'est I'aeifie hiogeographie region. M the
species level, tlie classification ol this group imdervvent
a eonsiderabk' change following a revision In Reid
(19S6a) of tlie Indo-Pacifie species associated with man-
grove habitats. Using a range ofevidenc-e from analomv
(in particular of penis, parasperniato/oa and pallial ovi-
duct), radula. siieil inoiphologv. Ii.ibital, distribution, and
copulation Iretjuencv. it was shown that 20 speeies could
be discriminated whereas onlv 3 had been conimonlv
distinguished |ireviousK. Since this revision, new collec-
tions have accinnulated. .\s a result, some of the earlier
conclusions nmst \k- revised and three additional species
can be recognisi'd in the region. The new material also
adtls significant extensions to the distributional ranges ol
some ta\a. It is the aim ol this paper to describe and
icdescrihe t\vo of the additional taxa (the third will be
deserihed elsewiiere, Stnckev antl Reid, in preparation)
and to document some new obsenations and distribu-
tional records. It should be used in coujimction with the
earlier work (Reid. I98fia).

.\t the time of Reids (I9S6a) svsteniatic revision, oth-
er aspects of the hiologv- of these tropical species had

Page 116

THE NAUTILUS. \ol. 115, No. 4

received relati\{^l\' little stiuK. llowcNcr, in the \eais
since then the\' ha\(> iccciNcd iiiiuli more attention.
Among several reniarkalilc tcalurc.s ol iJlloniria are the
a.ssociation of mam species witli mangnne trees, leading
to studies of their /onation patti-rns. diet and predation.
Some members are oxoxixiparons. and nian\ sliow
spawning or feeding migrations with hniar and tidal pe-
riochcitv. A striking color poK inoiphism of the shell is
shown l)\' species occurring on mangnne foliage and
these ha\e enierged as a model s\steni tor the stud\' ol
the niainteiuuKc of poK tiiorphisiii In natural selection.
There has also heen new work on ultrastructure, egg
capsules, radular plasticitv. parasitism and genetics. This
new literature will he lirief]\" rexiewed here.

NEW LITERATURE ON LITTORARIA SINCE 19S5

Reid (19S6a. 1989a) listed 36 known species oi' Litto-
raria vxorldwide (one of whicii was unnamed), but de-
t;iiled descriptions were restricted to those 20 found in
association with mangrove trees in the Indo-West Pacific
region. Since then the si.\ species occurring in the East-
ern Pacific region ha\e lieen described (Reid. iy99a; see
also Reid and Kaiser (2()()1 ) for t^vo new records of Indo-
West Pacific species in the eastern Pacific) including the
unique nonplanktotrophic L. ahcrniiis (Philippi, 1S46)
and the new taxon L. ro.scwatcii Reid. 1999. In addition
to the single new species described in the present paper
and the recognition of two ta,\a within L. (ii-ticiilnta (Phi-
lippi, 1846)' (as used by Reid, 1986a), a further new-
species from northern Australia is being describeil else-
where (Stuckev and Reid, in preparation; see W'armoes
et al., 1990, and Reid. 1992a, for additional regional tax-
onomic accounts). This brings the total for the genus to
39 species.

A preliminaiA phvlogenctic anaksis of all Litlorar'ui
species then knowii, based on iladistic analvsis of mor-
phological characters, was done In Reid (1986a). This
has since been revised and updated (Reid. 1999b). The
new phvlogenv was not well resoKed, but supported the
monophyly of the subgenera Pahistorhui and Litioiinop-
sis, and suggested thai the subgenus IJtlonir'ui was a
paraphvietic assemblage; a significant change to the pln-
logenctic classification of the genus was the combination
of five mainlv rock-dwelling Indo-Pacific species (L. pin-
tado (Wood, 1828), L. inanrilitnui (Lamarck. 1822), L.
alahmta (Philippi, 1846), /.. coccim-a (Gmelin, 1791). L.
undulata (Gray. 1839)) in (he basal stibgeinrs Frotolitt-
orariri. wiiich had previousK held onK /.. piiitddo (Reid.
1989a). The fossil record of Ijlltmind is limitt-d. but
better than that of Other littorinid geneia. probali!\ bc-
cau.se preserx'ation is more likeK- in niangroxc cnxiion-
ments than on high-energ\' rock\ shores. The known fos-
sil species were listed i)V Reitl (19991)) and discussed in
the context of vicariant events in the biogeograpliie his-
toiy of the clade. A similar histors-, of Tethvan origin ;uid
subse(]uent \icariance, has been proposed tor niangroxcs
by Ellison et al. ( 1999), who cited the worldwide diver-
sity gradient of IJtloraria species in support. Ivxtinetions

of marine invertebrates during historical time are rarelv
reported; L. fiammca (Philippi. 1847) was included as
one of onlv four possible examples among 'gastropods
discussed bv Cariton (1993).

Regrettablv little genetic work has \et been undertak-
en on Littoraria. Several studies of allozvine variation
within the Australian species have confirmed that the
species defined bv nioq:ihological criteria are indeed dis-
tinct (John.son and Black, 'l998; Schmidt. 1998; M.
Stuckev, personal conmiunication). Levels of intraspe-
cific genetic variation have been assessed using alloTA'me
markers in L. anguUfera (Lamarck, 1822) (Janson, 1985)
and in the hvo subspecies of L. cingidata (Philippi, 1846)
(Johnson and Black. 1998). DN.-\ sequences have so far
been published for the 16S ribosonial RNA gene (3 spe-
cies, Reid et al., 1996) and 18S rRNA gene (1 species,
Winnepenninckx et a!., 1998^, but have yet to be used
for phvlogenv reconstmction within the genus. This
should be a priorih'.

Although the genus Littoraria is well known as a char-
acteristic inhabitant of mangrove forests, this is not tine
of all the species. There appears to have been an evo-
Intionan- specialisation to the mangrove habitat, since
basal species occur exclusively on rocks, or on both rocks
and drifhvood, whereas only the more derived clades
occur on trunks and foliage of mangrove trees (Reid,
1999b). The three-dimensional structure of the man-
gnne habitat, and frequent .svmpatric occurrence of sev-
eral Littoraria species, has stimulated numerous ac-
counts of comparative zonation patterns. These have
been carried out in Australia and the western Pacific
(Cook et al. 1985; Reid, 1985; Boneka, 1994; Catesbv
and McKillup, 1998), Japan (Ohgaki, 1992), the Carib-
bean (Gutierrez, 1988) and the eastern Pacific (Blanco
et al.. 1995; Blanco and Cantera. 1999; Cantera et al.,
1999). In general. Littoraria species do not seem to be
restricted to particular tree species, but are found on
characteristic substrates (bark or foliage), at txpical tidal
heights and in tvpical horizontal zones (seaward or land-
waril fringes) within the forest; furthermore, the assem-
blages of species on continental margins and on oceanic
islands are different (Reid. 1985. 19S6a, 1999a). Some
of the tvpicallv mangrove-associated species can also be
found on shclteicd rock-v shores, including L. aiiindata
(see Crowe, 1997), L. sinensis (Philippi, 1847) (accounts
of distribution of 'Littorina scahra' on roek-y shores of
Hong Kong and southern China probabK refer to a mLx-
tnre of these tAVo species, e.g. Ohgaki. 1985; Yi and Li,
1988; You, 1990) MvXL.flaia (King and Broderip. 1832)
Nionlinhd ;uid Alves-Costa, 2000). Ecological data on
llidsc species tli;it ;nx' found mainlv on rocks and drift-
wcHid. such ;is /. unduhila and L. coccinea. are scarce
il'diilieek et al., 1994; Sacchi, 1994; Ohgaki. 1998).

Within the mangrove environment those s|ie(ies in-
h;ibiliii'^ liuiiks and roots ;it lower levels on the trees
nii'j;r;ile \erlicallv with the tide in order to renuiin above
die w.iler level (Yip[). 1985; (;utierrez. 1988; Ohgaki,
1992; Jensen, 2000). Similar behavior has 1h'(mi reported
on rockA- shores (Ohgaki, 1993; Svane and Pringgenies,

D. G. Reid, 200

Patif 117

1997) and displacfiiieiit i'\periiiu>nts ha\c lircii cairifd
out (Antwi and Aniexan-Ahimfi. 1987). In tontiast. spe-
cies hpicalK- Ibinul at higher lewis among tiic foliage of
mangnnes are not contaeted l)\ the higli tide, hut be-
come acti\e during rain and iieaxA' di'w (Little and Stir-
ling, 1984; Kohlmeyer and Behont. 1986; Ohgaki, 1992).
Lihowria irrorata (Say, 1822) occurs in salt marshes in
the soutiieasteni United States, where i( chmhs tlie
stems oi marsii grass and descends to teed on the suh-
.strate at low tide. This behavior has lu'cn investigated in
relation to lioth thermoregulation (McBride et al., 1989;
WilHams and Appel, 1989; Henn- et al., 1993) and a\oii!-
ance of predators (Warren, 1985; \'aughn and I-lsclier
1988, 1992; Di.\ and Hamilton, 1993; Du\al et ;il.. 199-4).
This species has been eniplo\ed in studies of trail fol-
lowing (Stirling and Hamilton, 1986; Tankersle\-, 1989,
1990;' Robbins and Hamilton, 1996). Unhke ;ill other
species of the genus, it is entireK- template in chstri-
bution and hibernates during winter (Paul et .il 1989).
Aggregation beiiavior has been described in /. tl'ii'i "u
Toch,- shores in Brazil (Moutinho and AKes-Costa.
200(;)).

Littoraria species sulfer high Ie\els of pre(hitioii. a|i-
parently mainK h'oin aquatic pretlators, accounting toi-
their avoidance oi submersion i)\ the rising tide. In man-
grove forests portunid crabs take snails ;it and below the
water surface, while some grapsids are able to climb sev-
eral metres up into the trees (.Maruthamutim et al.,
1985, 1986; Borjesson and Szelistowski, 1989; Reid,
1992b; Boneka et al., 1998). Predaton- fish ma>- also be
important predators of juxenile Litforaria and species
from lower \ertical le\els (Hughes and Jones. 1985;
Hughes and Mather. 1986; Borjesson and Szelistowski.
19S9; Catesby and McKillup, 1998; Duncan ;md Szelis-
towski, 1998). in salt marshes botli portunid crabs and
predaton- gastropods (.A/c/o/igcofl) attack L. irrorata
(Warren.' 1985; West and ^\■illiams, 1986; Dix and Ham-
ilton, 1993; Schindler et al., 1994). There is limited e\-
idence of predation b\' birds on Littoraria species in the
mangnne canop\- (Reid, 1987; Cook and C;ad)ett, 1992).
An unusual predatorv association has recentK' been re-
ported in southern Queensland, where U\t> species ol
sarcophagid flies are a significant cause ol moii;ilif\ in
L. filo.sa (Sowerbv, 1832) (McKillup and MeKillup, 2000;
McKillup et al..'2()00; P;ipe et a!., 2000). In \ietnam.
South Kast Asia and India the larger Littoraria sjieiies
are gathered localK lor food In fiumans i i\asm;itliaii and
Shanmugam, 1988; Poutiers. 19981.

On tropical rock'\' shores Littoraria h;i\e been re|)oi1-
l| ed to graze on diatoms, microalgae. c\;inob;Rtt'ri;i ;ind
r bacteria, but in mangro\e and saltuKush h;ibitats addi-
tional sources of food are available. Littoraria species
appear to be opportunistic grazers, ingesting ;i range of
fungal, algal, detrital and plant material and their role in
food webs has been examined (Kemp et al.. 1990; (>urrin
et al., 1995). Several investigators have stressed the likely
importance of marine limgi in tlie diet ol Littoraria in-
habiting both mangrove trees (Kohlmeyer and Behont.
1986; Christensen. 1998) and grass (Newell and Biir-

locher. 1993; Barlocher and Xewcll. 1994a. b; (iraca et
al.. 2000!. Whether mangrove pkmt tissue is a significant
food source is delnited i Ohgaki. 1990; Jensen. 2000; Lei'.
2001) and the composition of the diet ma\- depend upon
vertical zonation and tree spcfies '[ensen. 2000), The
mechanical function of the radul.i has not been studied,
but it has been shown tluit the form of the radular teeth
differs consistentlv in specimens of the same species
samjiled from rock and wood snlisti;ites. impKing that
tooth .shape is phenotypicallv plastic (i{(id and Mak.
1999). Growth rates can be r;i])id in tropic'al Littoraria
species, most reaching :isvmptotic size in six months to
one vear (Maruthamuthn and K;isinath:in. 1985; Bnrgett
et al . 1987; Cruz. 1989; Boneka et al.. 1997; Jensen et
a!.. 1999; Jensen, 2()00>. Breeding and population char-
;icteristics have been reported in several Littoraria spe-
cies I Bern. 1986; Maruthamutim ;incl K;isin;ilhan. 1986;
Yi ;md Li. 1988; Herjanto and Thomas. 1VJ95; Jen.sen et
al.. 1999).

The mtraspc'cific variabilitx' of littorinid shells has
becMi a connnon Ciiuse of taxouomic conhision in the'
past. In a uioq)hometric stucK of L aiiiiiilifcra through-
out its Atkmtic range. Merkt and Kllison i 1998) reported
strong intnispecific variation in shell shape, but argued
tli;it this was a conse(|uence of ecopheno^vpic effects
rather than genetic differentiation. However, an earlier
study of the same species over a more restricted area
had reached the opposite conclusion (Janson. 1985).
When considering variation betAvc'cn species, it has often
been obsened that shell thickness decreases in species
zoned at higher levels on th(> trees, reflecting adaptation
to the severity of acjuatic predation at lower le\els (Cook
et al., 1985; Borjes.son and Szelistowski. 1989; Duncan
and Szelistowski. 1998) and the danger of dislodgenient
in the c-anopv (McMahon. 1985).

In manv Littoraria species the most striking aspect of
shell variabilitx' is the color jiolvmorphism of the shell.
Indeed this feature has stinnilated the most active field
research on (lie gcMms. so that species <)\ Littoraria have
become recognised as a model svstem for the studv of
visible genetic polvmorphism ((.'ook. 1992). I'nfortu-
natelv. earlv studies suffered from the confiision of sev-
enil different svm]-);itric species (Hughes and Jones,
1985; Hughes ;md .Mather 1986). It has been obsened
that there is a consistent correkition between monomor-
phic- (;ilthough still variable) shells and a low-level hab-
it;it on trunks and branches, wiiereas species inhabiting
foliage are polvmorphic (Cook, 1983, 19S6b), although
mider exceptional circumstances polvmotphism has also
beiMi reported in a trunk-dwelling species (C^ook ;ind
Bridle. 1995). The three princi]ial moqihs are vellow.
brown and pink (or Orange) and each appears to be
cnptic against different parts of the \isuallv varied back-
ground of the mangrove canopv, or to mimic elements
of the foliage (Cook. 1983; Reid. 1987). Assuming that
the polvTnorphism is genetic in origin and not selectiveK
neutral, there are a number of mechanisms bv which it
could be maintained in natin-al selection (Reid. 1987:
Cook and Gadiett. 1992). Evidence has accunnilated for

Pasjo lis

TUP. NAUTILUS, \ol. 115, No. 4

D. (;. H(>id, 2001

Pai'c 119

sisiiiifiraiil iliaiim's in iiioipli licc|urnrics ac((ii(liii'4 to
tlic cdiriposition of the l)ack>j;n)iiu(l (cij;. tree species ami
al)uii(!aiR-e 1)1 li)lia<ie; Heid, U)S7; (.'ook and (iaiiietl.
19S9; Cook. 19y()a; Schmidt, 199.S) and tlieiv has heen
one nianipnlativc field e\perinit>nt tiiat sutisiested the op-
eration ol lre(]nenc\ -dependent (apostatiel selection
(Reid, 1987). \e\eitheless, the identit\ of [lossilili' \isual
predators is a matter ol speculation: hirds. erahs and
even flies ha\e been su^^ested (Reid. 19S7: Cook and
Garhett. 1992: McKillnp'et :il.. 2000 i. The differences
in thermal properties ol the niorphs iiia\ result in l>e-
havioral ilillerences that contrihute to site sck'ction and
background matching (Cook, 19S(il); Cook and Free-
man, 1986: Reid, 1987). The color inoiphs of /,. jiiillcs-
cens have also been showni to differ in shell strength,
size, variance and growtli rate iCook et al . 1985: Cook.
19901): Cook and Kenxon. 1993: Bonek:i. 199(r. Honeka
et al., 1997).

ReIati\eK' lew new anatomical studies ha\e been e:n-
ried out since 198fi. Egg capsule shape luis been docu-
mented in a number of ailditional species (Bern. 19Sti:
IIo, 1987: -\lak, 1995). The llagelhun-like structure of
the paraspermato/.oa is a character ol phvlo^enetie sig-
nificance (Reid, 1999b): this has bctMi named the pscn-
dotrich and has been tlu' subject ol struetnr:il :md on-
togenetic stucK (IleaK ;md |;uniesou. 1993: iiuckland-
Nicks et al., 2000). The lunctions ol parasperniato/o;i
have been rcxii-wed hv Buckland-N'icks (1998: 1-inck-
land-Nicks et al.. 1999). Eertmau (1996) described the
ultrastructure ol the gill filaments ol L (iiiicuhtUL

MATERI.\L AND .METHODS

Shell dimcTisions weie uie;isured with \erniei e:ilipers lo
0.1 nun. Shell heiglit (H) is the m;L\innun dimension
parallel to the axis of coiling, shell bre:ullli iB' the ma.\-
imuui dimi'iision peipeiidicuhir to II. and (he length ol
the aperture (LA) the greatest length from llie junction
ol the outer lip with the penultimate wliorl to the an
tenor lip. Shell shape was quantified :is the r:ilio II B
and H/LA (rekitiw spire height. Slfi.

For geni'ral :iceounts ol the m:Lle and lem;ile :iUiitom\
of Lifforariii, and methoils of stniK. see Beid i 198(ia:
1999a). Li\ing animals wi're relaxed in 7.5'/f maguesium
chloride in fresh water, fixed in lO'/f sc:iwater formalin
and stonxl in SO^r ethanol before examination. Banis-
permatozoa were I'xamined Irom the seminal \esicles of
specimens fixed :ind storetl in lO'f sc:iw;iter lormalin.
The relatixc radular leu<fth was the total rathilar k'ntith

di\ ided b\ shell height. B;idnkie were cle.iued b\ so:ikini4
iu a h\pochlorite bleaching solution ,it room tempera-
ture' for about 5 min. rinsed in distilled water mounted
on ;i film of pol\Aiii\l acetate glue ou glass, allowed to
(In in air and coated with gold and palladium before
(.■xamination iu :i scanning electron microscope. I'nworu
jiortions ol radulac were xiewed in three orientations: in
st;uKkir(l H;it view iVoni x'erticalK' above the radula (to
show sh:ipes of teeth), at an angle of 45° from the front
end ol the railula (to show sluqies ol tooth cusps), and
at ;in angle of 45° from the side of the raduki (to show
relief).

Inslitiiti<)nal ahhre^iati<>lls:

.\.\1S. .■\ustr:iliau Museum. S\dne\

ANSI' .\c:idem\ ol \:itnral Sciences ol Phikidel]ihi:i

BMXJl. \:itural lliston .Museum. London

\l\ll\. Museiun \;iliou:il d llistone \alnrelle. Baris

\S\1 1 National Science .Museum, 'lokxu

\.\1\\ National Museums and (Galleries ol Wales. ( .':u-

din

UNllL. Xatioiuial Natnmhistorisch .Museum. Leiden

USWI. National Museum of N:itur:il lliston. Smithson-

i:in Institution. Washington. D.C.

Z.\l A. Institute ol 'laxououiic '/oolog\. I'liixcrsitx ol .\m-

stei'dani

SVSI EM AilCS

iJI/oriniii {I .itioriiKtpsis ' scabrti ; Linu.ieus. 175S)

Distribution: New leiords: M;iscareue Islands: Ro-
drit;ues iNMW'. N'ietnam: Lo Bixcr. Nha Trang
(B.MNin. Chiu:i: S:ui\:i. Ilain;m Iskiud BMNIH. Tai-
wan: Tan Shui (BMNTL. |;i[)an: .\mami () Sliima iKxu-
shu I'nixo'rsitx h Inoda. Ishigaki Shinui i BMNT I h Chichi-
jinia :md llah:ijim;i. ()ii;is:ix\ai'a Iskinds l''nkndii. 1993).

I .iltoniiiii • Lilldhnopsis- hilcd I'hilippi, ISI7)

Distrihntioii: New records: Nieluiun: Lo Bixcr. Nha
Trang (fUlNll Indonesia: l,emb:u'. Lombok ; BMN II i.

l.iiloniriii ' lJII(>i-iii<>psis< piillcscciis i I'hilippi. IStfii

Distribution: Nexx records: \ietn:mi: Lo Rixcr Nha
Trang (BMNH). China: Sanxa. ll:iin:iu Iskind BMN 11).
T:iix\an: Tan Shui (l^MNTIl. |apan: Na/.e. .ATuamioshinui
iKxiishu Unixcrsitx): Kabria Bax, Isiii<'aki (B.MNTI).

Figures 1-13. SlielK i>\ LitUtnnid iHiiiiiilciisis new species and IJttoriirin intermedia 1-11. /.. hcii'^alriisis. 1, 7. 15:itii Mauii;;.
Peiiani;. M:ila\sia il5.\INH 2()()1()1 lo: 1, niafc; 7, Icmafd. 2, 4, 8. I'aratxpcs: Hare Island. Cailf of Mannar, Taniilnailu, In(li;i
(HMNH 2001(11 t(l; 2, 8, female: 4, male). 3, 6. Tntieoiin. Tamilnadu. India (BMNII 20()0()7S3: females). 5. Sun<.»ei Mcrhok, near
Pantai Merdeka, Kedah, Malaxsia i BMNII 2()01()11S: malcL 9. .-\i) Nam Bor Phuket Island. Thailand ( BMNII 2()(II()| 17: lemalel.
10. Ilolotxpp: Hare Iskiiid. Culf of Mann:ir Tamilnadu. India iB.MNII 2()()1()I14: male). 11. .-^o Nam Bor Phuket Island. Thailand
IBMNII 2001011(1: male' 12, 1.3. /, iiitcnitidiii: ll:ire Island. ( ailf of \lann;ir Timihwidn. In(li:i iBMNll 2i)0007."il 12, male;
13, leuialel.

Page 120

THE NAUTILUS. \(.I. 115. No. 4

\\'estern Australia: Bt'rkt'lr\ Ki\(r. W coa.st Joseph Bon-
aparte Gulf (BMNH); Maii<iro\c Bav, (^ape Range
(BMNH). Mariana Islands: .\leri/o. Cuain (BMNH).'

Liltorciria {Littoriiiojisi.s) hcn^dlcnsis new species
(Figiu-es 1-11. 14- US, 20-22. 34. 35)

Lilloriiia acahra — XicUcn. Ii)7(i: 1-4. ti<j;. l.'\ (in part, iiitliulcs
L. saihni (Linnaeus, IToS). L. pallcsccns (Philippi, lS4(i)
and prohahK /.. iiilcniirtliri (Philippi, 1S46)).

Littoniiia inlcnncdUi — Heicl, I9S5: .'j^-fi.S. fiijs 9, 11 (zonatioii)
(fig. 9 includes /,. iiilcniicditi ll'liilippi. 1.S46)). Cook and
Garbett. 1989: 5. fig. If. iv (penis) (in part, iuclndt's /,.
intermedia (Philippi. lS4fi)).

Litturaria (Uttoiinopsi.s) intennediu — Reid. 19S6a: 124-135.
figs 43b. 44e, f, 45c, d. 4fib (jienis), 46k (parasperniatozoa)
(in part, includes L iuliiiiu-diii (Philippi. 1S4G)).

EtMTiologA-: From the Ba\ ot Bengal, of which the
known distributional range" ot tliis species spans tlie
southern part.

Types: Hok)t\pe BMNH 2(1(110114 (Figure 10); 17
dn- parat\pes BMNH 20010140 (Figures 2, 4, 8); 100
paratopes in ethanol BMMI 20000755; 4 chv paratopes
USNM 1000887.

Type locality-: Hare Iskuul, fiulfol Nhnuiar, Tamilu;iclu.
India.

Material Examined: 32 lots; 15 penes; 7 sperm sam-
ples; 4 pallia! (niduets; 4 radulae.

Shell (Figures 1-11): .\dult size range 12.3-28.6 nun.
Shape elougate-turbiuate ( H/B = 1.49-1.86; SH =
1.72-2.16); whorls modei'ateK rounded, suture im-
pressed, periphen ol last whorl angled, olteu with a
raised rib; relativelv thiu-slielled. Mature lip sometimes
sliglitK fiared in males, \arices rarelv formed. Columella
wide, pillar straight or slightK eon\e\. e\ca\ated. Sculp-
ture of (8)9-10 primars spiral giooxt's on spire whorls,
equalK' spaced; priuiiu^x groo\cs remain as incised lines
or up to 0.2-0.3 rib width on kist wliorl, often sliglitK
deeper and wider posterioiK (where occasionallv pos-
terior groove mav be 0.5-1 rib width); iutenening ribs
remain undivid(>(l until last whorl, where 2—1 posterior
ribs (excluding rib adjacent lo sutnri') become di\ided
by a central impressed line: peiipheral rib usualK' raised.
occasionallv carinate; somelimes ;» u;uii)w liblel inter-
polated in 2-3 posterior grooves; b;isiil ribs Huer; tot;il
ribs on last whorl 21-35. Surlaee gloss\. spinil mieros-
triae faint or absent. Frotocouch 0.35 nun diameter, 3.5
whorls, with spiral ribs and sinusiger;i notch. (!olor \;u-
iable; ground color criMUi. pale \ellow or oelu'e; [):ilt(in
of dark browii daslies on ribs, on spiic whorls ilashes are
usually axialK' aligned to form obli(|ne or /ig/.ag stripi's.
but on hist whorl dashes become less discrete and break
up into diffuse blackish to red-brown motlling; often ;i
broad paler zone on middle of base; p:illcrn sonietinies
faint, so that shell appears ncHow with grcN-brown mot-
tling. Coliunella pmple brown to dark violet; ;iperture
with e.\terior pattern showing through aTid not obscmcd
b\' the thin whitish callus.

.Vniinal: Headioot. operculum, pallial oxiduct and rad-
ul;i ( Figui'es 34, 35) do not differ significantK from those
ol /,. iiileniwdia as described b\' Reid (1986a). Penis
I i'"igui('s 14-16. 20) bihu'cate; large dark brown glan-
dulai' disc carried on long branch of base; smooth bkide-
shaped filament with mucronate tip. separated from
wrinkled base bv constriction. 0.4—0.5 total length ol pe-
nis; entire vas deferens from tip ol peni;il hiament to
prostate closed as a duct. Paraspermatozo;i (Figure 21;
l\eid. iy86a: fig 46k) 14—26 |j.ni (including rod jiieces):
rod pieces eoniposetl ol a bundle of several uaiidw el-
I'uients. or I'arcK single, usually projecting from cell; cell
hlled with large rouiid granules. 0\"o\ iviparous; embn'os
brooded between gill folds in mantle c;i\it\ until M'liger
stage; development planktotrophic.

Di.stribiiti<)n: Habitat on branches ;iud foli;ige ol
mangroves (Avicennid. Rhizopliora. Somicriilid) ;ind
m;u"itime trees (Pcmplus), up to 3.5 ni above groimd.
iilwavs above water level; most frequent at seaward edge
of mangrove forests, but e.xtending far back into forest
(see Reid. 1985: figs 9, 11, for zonation; as 7. iiilcniw-
ili(i); onlv rarely found on rocks. Fomid in modenitelv
turbiti and estnarine localities as well as on nearshore
iskuids. Range (Figure 22) from hulia to western Nhilav
Peniusnki :ni<l northern Sumatra. Records: India; Chau-
pati Beach, Mumbai (USNM); \'engurla (USNM); Man-
dovi EstnaiA', Goa (BMNH); Netravata R.. Mangalore
(USNM); Ciochin Harbor (ANSP); Alkppev (B.NINH);
Tuticoriu (BMNH, USNM); Krusadai I., GulY of Mannar
(BMNH); Portonovo. Tamilnadu (BMNH); Chennai
(BMNH); Port Blair. Andaman Is (BMNH). Burma;
Thavawthadaugvi Kvun (Elphiustone I.), Mergui .\rch.
(BMNH); K;iil;m Kvun (King 1.1. Mergui Arch.
(BMNH); Lanbi Kvun (Sullivan I.), Mergui Arch.
(BMNH). Thailand: Goh Contee, Ranong (MCZ); 70 km
S R;mong (BMNH); Ao Nam Bor. Phuket I. (BMNH);
Puhiu Tang;i, Butang Is (USNM). Makivsia: Merbok Es-
tuan (BMNH); Batu Maung, Penang (BMNH): Matang
EstuaiT (BMNH). Indonesia: Belawan. Deli. Sum;ttra
(RNIIL); .Shiabaug, Simeulue I. (RNHL).

Remarks: In his discussion of L. iiilcniicdin Heid
(1986:1: 134) noted tliat specimens Irom southwestern
Thaikind. Penang and India hail thin-walled, olten large
shells with :ui irregnku eoloi' pattern. This lorm occurred
logelher v\itli tvpic;il /.. intermedia ;it Phuket Island.
4'h:nkuul. \el w;is not considered distinct, since it was
ckiimed that intermediates were present and bec;nise no
imiitomie;il differences were obsened. .Mthough the
shells can be iloselv similar and ;iiiat(nnic;il differences
are slight, new evidenci' supports recognition of this
lorm as .i distinct species. This was first suggested bv
11(1(1 (ibsci-viitions at the t\pe localitv (Hare Island, (uill
III MaiiiKii'. Ileie. three species occurred together on
slnnbs (il I'rinphis oveiluuiging a sheltered sandy lagoon
behind a fringing I'eel. There w;is a eV-AV /onation: /,.
ililcnnedirl w:is jireseiil on tiiinks :it ,i level ol 0 oO cm
;ibove the high \\:ilc|- iiunk: L. scahni oeeiipicd ;i /one
.lO-tiO em iibovc lii''li vwiter on ti'iinks :m(! briinches.

D. (;. Hcid. 2001

Paa-

whereas a third speeies [L. hciiiialcusis) oeciiiicd nn
lioth ioHage and l)ranclies from a lici'^ht ol. '50-100 em.
The tliree speeies eould lie separaled li\ sliell eliaracters
(see helow > and, most siy;nifieantK. all three were cop-
ulating onK with eonspecifies. Siil)se(|ueiit anatomical
examination loimd a consistent dilleicnce in the penis
ol L. h(iiij_iil(iisi.s and /,. iiitcniii-diii. iU'cxamiiialion ol
nnisenni colk'ctions has i\'\eale(l s\nipatric occurrence
(mixed samples) ol L hcii'^alcnsis and /., hilcnncdUl at
the tollowinu; additional localities: ( l.ille. Sri l,anka
(BMNH); two islands in Mei'^ni \rcliipelaii,o, Hurma
(BMXIl); Ao Naui Hon Phuket I.. Thaihuid (l5M\lli;
Fulau Tanj^a, Bntang Is. Thailand il'SN'Mi: Sinahant^,
Simeulne I.. Indonesia (RNHL>. In <'acli casi' separation
ol shells was strait^htlonvard. with no iutermeiliates. and
males from Ao Nam Bor were iilso tlistiu<^uislicd nsin<;
the same penial characters as at the t\|)e localitx.

The ditferenees between L. hvw^alcnsis and /, iulcr-
mcclia are sununarised in Table I. The shells ol /, in-
termedia show considerable geographical and ccotxpic
variation (see Reid, 1986a, and below), so the descrip-
tions appK ouK' to specimens from within the 'jjeograph-
ical distribution ol L. hcii<ialciisis and Irom the uian-
groxe habitat. For comparison, sliells ol L. iiilciincdia
from the t\pe localit\' ol L, hcw^/iliiisis are illustrated
(T'ignri's 12. 13). The shells ol L. I)(ii<i(ilciisis are usualK
larger, up to 28.6 mm. tliose ol L. iiitcniicdifi larcK i'\-
ceeding IS nun (although the maximum si/c ot liic spe-
cies as a whole is actualK" greater. .32 mm: Hcid. 19S6a).
Thev are also of thinner texture, with more sharpK an-
gled periphen, than those of L. iiitcniicdia. Sexual di-
moi"|ihisni is evident in both species (Figures 1-1.3).
males iieing smaller and with a relativcK lower spire and
larger aperture (Reid. 1986ai; howexer. males ol /> hcii-
gf//<';i.s/.v often have a more elongateK patulous shape,
reminiscent of a Surrinca (compare Figures 5 and 12),
The overall lolor of the shell is ochraceons or vellowish,
with brown pattern, in L. hciiiialcusis: in contrast, that
of L. iuli niicdiu is grev with black to brown p.ittcrn. In
detail, the dark pattern ot dashes on the ribs is similai"
on the earlv whorls of both species, but on the final
whorl the dashes of Z^. iutcrmcdid icmam nioie discrete
and axiallv aligned, whereas in L. hcii^iilciisis the pattern
becomes more diffuse and mottled. In both species
there appeals to lie a direct effect of substrate upon
shell color, presnmablv mediateil through diet i Riid.
1986a). Shells from Rhizopluini and Vcinjiliis are ilarklx
patterned, whereas those from Aiiirniiin and Soiiinniliii
are conspicnousK paler. OnK' L. intermedia is known to
occur on rocks and these too are usualK iclati\el\ ])ale
in color.

The ouK anatomic.il difference that has been ob-
sened is the shape of the penis: that ot /.. heie^aletisis
has a more robust filament and a larger glandular disc
borne on a longer branch of die base (although the pro-
portions of the penis depend upon the degree ol relax-
ation, .see Heitl. 19S6a: fig. 46). In all animals examined
the penial glandular disc of L. />eii>:aleiisi.s was dark
brown, whereas that of L, intermedia was cream or vel-

lowish. There mav ,ilso be a dilleicnce in the parasper-
mato/.oa: in /, henfialentis the rod pieces nsuallv project
(Figure 21; Hcid, I9S6a: fig. 46k). but in L. intermedia
thev seldom pierce the oval outline ol the cell (Rtid.
lOS6a: figs 46i. j) (ba.sed on 7 and 5 samples respec-
tivclv).

There is little information on the comparative habitats
ol L hen<^aletisis and L. intermedia since the txvo have
ouK icccniK been dislintinished in the field and louTid
in svnipativ. I'lie available observations indicate that /,.
heniialcnsis is lound higher on the trees, throughout a
greati-r width ol the mangrove lorest and that it is more
tolerant of turbiil and estuarine conditions. l'"or exaTiiple,
at the nmddv bav ol \o Nam Bor, Phuket Island, Reiil
( I9S.5: fig. 9' recorded /. 'intermedia' Irom throughout
a mangrove st.uid SO m wide. tliidn<4h to the laTidward
/.one and at heights ot up to .3.3 m above the gronnil.
Both species ari' jiresent at this site (collections ol Hcid
in BMNII) and since in other ])arts ol its range L in-
termedia sensn slrirla is known to occur mainlv at the
seaward edge and at low levels on trees, it is likelv that
the high-level anil landward records applv to L heiejji-
Icnsis. M the niuddv. turbid site ol Batn Manug, Penaug,
onlv /,. hcirj^aliiisi'^ was present and here Keid (1985:
fig. II. as L. inlcimediii ' lei'orded it Irom througliout a
belt ol Avicennia 10 m wide and from up to 2.0 m above
the ground. Likewise in the iiearbv Merbok Estnan L.
hen^alensi.'i (Kenrred alone and Irom seav\ard edge to
landward lring(> (peisonal obsenation ', The tvpc localitx'
of L. heniialensis is a sandv island with a binding reef
and clear water: here i)otii species occurred and L hen-
gr;/(7(s/.s was found above the level ol /,. intermedia aTid
on botli foliage and branches, whereas the latter was
pieseiit onlv on trunks (and has rarelv been lound on
leaves elsewhere. Reid, 1985 1. Litti>rar'ta henufdens'is has
so far been recorded almost exclusivelv on trees (one
spei inien on .i shelteied rock'X' shore near Ranong. Thai-
land: H.MMI. whereas L intrrmedia is coininonlv
found on both trunks and sheltered roekv shores (Reid,
I9S5: I9S6ai.

Rcitl iI9S6a: 57-58) discussed a distiuiiion between
Littoraria species with "oceainc' lUid 'continent;il' distri-
bution patterns, representing extremes on a gradient of
ecological tolerance. The former referred to a suite oi
iiabitat characteristics including clear water, nonn;il sa-
linitv, narrow mangrove fringes or seaward edges ol
broad mangrove belts, and offsliore islands. In contrast,
fontinental' li:ibitats were eharaeteii.sed bv turbid, es-
tuarine \\ater. eutrophic conditions, and broad forests on
continental margins. It is clear that on this continnum
/. heniSftlensis extends to more ■continental' habitats
than /, intermedia. For exampli'. in the .\ndaman Sea
both species have been recorded Irom the .Andaman Is-
lands. Mergui .\rchipelago. Phuket Island and Butang
Islands uk'tails given above): thes(> are all 'oceanic' sites.
However, at a muddv site on Penang and in both the
Merbok and Matang Estuaries, and in the vitinitv ol
f^anong. on or close to the mainland of the Malav Pen-
insula. /,. hennalensis occurred alone (large collections

Page 122

THE NAUTILUS, Vol. 115. No. 4

D. C. Hi'id, 2001

by D.G. Reid, E. Asliton and |.l^. Ta\Iiii- rcspcclivcK :
;J1 BMNH). Similarly, in India /-, hcii'^alritsis has been
recorded over a lar^e extent oftlic eontinental coastline,
from Mninbai to (^liennai. From islands in Mic ( ailldl
Mannar and Irom (ialk- in Sri Lanka. In ((inli.isl. m llic
same region /.. iiitiniudiii has lu'en recorded onK from
islands in the (ailFolMannar. from (ialle and from Trin-
comalee. Sri Lanka (see below), all of which apjiear to
be 'oceanic' sites. ( )n a wider geographical scale the dis-
tribntion ol /. hiUnncdui extends aci'oss the Pacific to
the islands ol Hawaii antl PoKnesia. again icllcctin<j; the
'oceanic" character ot this species.

The northern limits of L. bcwj/ihusis in the 15a\ (if
Bengal are not clear, owing to the lew available collec-
tions h-om northeastern India <uid Bangladesh, Sd far.
onh' two lAttonirki species ha\e been recorded Imm
Bengal ant! Bangladesh, L. lUlicdIiiIti and /. nirhiiiit-
stoina (Reid. UlSfia; see below).

The man\ moiphological similarities between /, Ihii-
ffilcnsis and L. intermedia lea\e little donbt that the two
are sister species. Likel\- s\iiaponioqihies inclnde the
blade-shaped penial filament with mucronate tip and the
bmidles ot narrow roil-pieces in the paraspermatozoa.
both niiii|Me in the genus. The closed penial \'as defei'ens
is a likeK s\napomoipli\' of a clade comprising /.. hcii-
grt/(7is/v. /. iiiti-nucdia. L. suhiittata R(id, 19Sfi and L.
philippimui (Reeve. 1S57) (Reid. 19991)).

Confusion ol /> hciej^dlcnsis with other speeies is less
hkeh. In southein Iiidi.i and the sdutlieiii .\ndanian Sea
its range overlaps with that ot L. scahrti and the two can
be found svinpatricalK' in relati\el\' 'oceanic' habitats (in-
cluding the \^-pe locality in the Gulf of Mannar). IJtto-
raria sicuhra is easil\- recognised b\- its wide, white col-
umella. Ijltorarin jxillcsceiis (Philippi, 1S46) is even
more oceanic in cliaracter than L. scnhni and has been
recorded sxinpatricalK with L hcii<^alcii\is onK in the
Andaman Islands, Penang, Butang Islands and Phuket
Island: it is distinguished b\ its colorful. poKniorphic
shell, lack ol secondan sculpture .md the rounded inner
lip of the aperture. In each case pi'uial ch.uacters are
diagnostic (Reid, 19Sfia). The distribution of L. hcii'^a-
Icnsis just touches that oi L dclicatuld. single s|)e( iniens
of the latter having been seen Irom the Cull ol .NLinuar
and the Andaman Islands, altliough most are Irom the
head of the t^av of Bengaf (see fielovv ), Sfiells ol these
tvvo species can be superfitiallv similar: both ni;iv be
thin-shelled, and veliow with a faint or dilluse p.itleru.
Nevertheless, tliev can :ilvvavs be distin'j;nislied bv the

more numerous prinian grooves i 1 1-14 i and much finci'
sculpture (35-50 ribs on final whorl 1 of /.. dcliailulti: the
columella of that spi'cies is narrow and excavated in the
tvpicallv vciA delicate shells, although it mav :ipproach
that ol /,. Ixu'j.dlcilsi'i in width in occasional thickei'
slu'lls. The shape ol I, dclicdiulii is subtiv diirerent; the
spire whorls are Hatter and the spire nsuallv slightlv tall-
er. Most importantlv. the |)enis ol /.. dclicatidd is entirelv
different (fiiiures 3f, 32. sec below).

Lilhiidiid [iJtloiiiuipsis! inlcnncdid i, Philippi, I S46)
(iMtiures 12. 13. 17-19. 22)

Svnonvniv as in Keid (1986a). excluding the entries now-
listed under /,. I}(ii'j,alcusis above. New references:

Littoiarid l.iltohnojisis' inlcnncdid — Hi'id. lyStia: 124—135,
figs 43a, c-i, 44a-d. g-i. 45a. 451) (protoconch). e. 45f
iradnla*. 46a. c-li ipenisi, 46i, j (paraspern)ato/oa), 46l-<)
iovidiicti, 47 idistiihntion) liii part, includes /.. hcnudli'ii-
s/.v )]cvv species '.

Di.stribution: Records and distiibution as in R(id
(]9Sfia: 132-1.14. fig. 47>, but excluding all records from
mainland India except those listed below, and also I'.x-
clnding records from Penau'^. Merbok estuai"V and Be-
lawan i the excluded records appiv to /,. hertfialcnsis, see
above i. The doiibtlnl iccoid fioni the (Galapagos Islands
I Reid. 19Sfia: 1-34 1 is now consideicd unreliable and
should be excluded i Reid and Kaiser. 2001'. New and
I'econfiirned records: Kenva: Kilifi R. estuarv iBMXIL.
Saudi Arabi.i: jedd.ih H\l\ll Oman: Masirah 1.
(ZMA); Khor .\l-|arama. Ras Al-lladd i BMML: Banchir
Khavran (BMXII). United .\rab I'^mirates: Fujaira
(B.M'MD: Has Al-Khaim ah Khor. Dubai BMXIL: Khor
Kalbii. Shaijah iB.MNll'. India: islands in Cochin Har-
bour. Kerala (ANSP): .Manali I. and Hare I.. Cnlf of
.Mannar (BMNIL. Thailand: Krabi BMMI . .Malavsia:
Pnlan (iava. Kota Kin:ibaln. Sabali iBMMli: Kudat! Sa-
bah iB.MXIli. Melnam: l.o H Xha Traug iB.MXII).
China: Bcigang I.. Hainan IJ.MXHi. Indonesia: Knta
Beach, Lo/nbok i BMXII). Philii)pines: Mandalec, S El
Xido, X Palawan i l5MXil). Taiwan: Tan Shni (BMXII).
|,ipau: Ishigaki (BMXII): Vakoshima (S. L'o/.nmi (-'olln);
.\maniioshima (S. Uo/.umi Colin); Sliirahama, W'akavama
I BMXII): Chichijima, Ogasawara Islands (Kukuda.
1993). Australia: Wooli Wooli \{.. Xew South Wales
i.AMS'. Tou<i;a: Sopu. Xnkn'alola. Tongatapu i BMXII).

Hemarks: Kcid il9S(ia' characterised this as a tvpi-

Ficure.s 14-21. Penes and |3ai'asprriii,ilii/iia nl I .illoidrin hi'ii'^dlcnsis new species and I.illiunrid inicnncdid I4-I(i, 20. I'eiies
of L. hcnoalcnsis. 14-16. Pa)at\pes; ll.ire Isfuid. Culldf Mannar Taiiiiliiadii. India (BMXII 2()()0()7.55; shell II = 17.0 mm, 16.4
111)11. 16.6 mini. 20. .^o Nam Bcir Phiikit Island. Tliaihiiid < BMXII 2()()1()116: shell 11 = 13.0 mm). 17-19. Penes of/., intermedia.
17, 18. Hare Island. Culf of ManiKin T.iiiiilii.idn. India (B.MXII 200007.54; sh(-II H = 12.S mm. 15.3 mm). 19. .Vo Nam Bor.
Phuket Island. Tliailand (BMNH 20010119; shell II = 13.4 inni'. 21. Paraspermatozoa fro))) one panilvpe of /., I>en>i(densis: Hare
Island, Gulf of Maiiiuir Taniilnadii, liidi;i (BMXII 200007.55'. Alihreviations; pli. penial l);Lse (wrinkled); pf. penial filament (smooth):
p2;d. peniiil <'laiidul;ir disc. Stipple on peiii.il ^l.iiidiifn disc- indic-;itcs dark hrovvii coloration; sliadiiii; at hitse ol penis indicates hkick
pigment in epithclinMi.

Page 124

THE NAUTILUS. \i,l. 1 15. No. 4

Tablel. Suiiinian' ol dillcrcnccs lictwccTi l.iltoraria iH'ii^dlfii.sis new species ami /. iiihTiucfliii (shell eharaeters appK to specimens
Irom inancrnne lialiitals in nortlieastern Iiulian Ocean onK ).

(IharacU'i

l.illiuiiriii /;eiii'(//(;is/.s

Litlonirid inleniwdid

Shell shape Larger [in 29 mm), thinner te\tiir<'. peripheral

anu;ulation with raised rili
Shell color Ochre to \ellow, with hrown pattern; on kisl

whorl the dark (lashes ar<' dillnse

Penis liolinsl lil.unent; brown gl.nnlulai (hsc home on

long hrancli of base

Paraspermatozoa Hod pieces project from cell

ll.ibitat More tolerant ol turbid and estuarine condi-

tions: occurs higher on trees and throughout
K)rest: lound on trunks, branches aud loliage,
rareK on rocks

Distribution Northe;isteni luilian Ocean. Ironi India to west-

ern Mala\ Peninsula anil nortlieru Sumatra

Smaller (to LS mm), more solid, more rounded

periphen
Ciiev. with black to brown pattern: on last whorl

the dark daslies remain discrete aud axialiv

.iligned
More slender filanicnt; cream glandular tlisc

borne on shorter branch ol ba.se
Kod pieces seldom project Irom cell
Prelerence for clear water; lound lower on trees

aud onK' common at seaward edge ol lorest:

lound on roots, tnmks and slieltered rocks

Occurs wideK throu<iliont Indo-WesI l':icilic re-
I'ion

callv oceanic species, with a wide tlistriliutioii from
South Africa to the Red Sea. tliroughout the Indo-West
Pacific tropics to Hawaii ant! Folviiesia. Within this area
he pointed out three ilistinctixe geographical tornis.
from the central Pacific, from the western Indian Ocean
(including the Red Sea) and Irom India and the Anda-
man Sea. It is demonstrated alio\e that the last ot these

is a distinct species, L. bcii^alcu.'iis. Its recognition re-
moves the anomah' of tlie occurrence of this 'form' in
tlie relatively continental conditions ol hroad mangrove
lorests in estuaries on tlie mainland shores ol the west-
ern Malay Peninsula. .\n obvious (juestinii is whellici'the
other 'forms' might also desene specific status. Accord-
ingly, much new anatomical materi;il has been examined

Figure 22. (ieographical distribution n\ Lilhiraiid hriii^iiliii\i>: wilh. loi couiparisou. llie icKinls ti\ l.iltoitiriii lulcniii'd'tii Irom
tiio same area (see text lor lists of records;.

D. (;. Heid, 2001

Paee 125

from tln'oiiiilidiil tlif raiiij;c (il lliis s|)ci'ics; iu naitifuiar
fifi |iciu's liavf Ix'cii drawn. I luwcs cr. iki (iIinIoiis dis-
fdntiimilic's liaxr hrcii loiiiid. I'lirllicriiiiirc. in areas
siicli as tlif Soloiiioii Islands. New (Caledonia and l'"iji,
where o\erlap in the raui^es dI a putative Indo-Malav
species (the 'tNpieal torni) and a putative c'ential I'aeitii-
species nii^ht lia\e been predicted, no e\id<'Tice has
been lound lor the svniipatric ociinience ol discrete
coiicliological or anatomical t\pes. Instead, the evidence
continues to suggest a single widespread species, tvpical
ol oceanic haiiitats, in wliiili shell cliarat-ters show small
and gradual changes (iu the color pattern, lre(|nencv ol
color morphs, and presence ol setondaiA scnlptnic
across tlie considerable geographical range, .Shell I'liar-
acters continue to siiow a strong coirelation with habitat
as wi'l! as geographv, as pointed out bv Heid i 1986a).
Thus, tlu'onghout the range, those specimens Irom rock\
shores (anil from certain mangroxc tree sjiecies such as
Alicciiiiid and Soniicnitiii) are paler in color and show
a more ilitfuse pattern, in comparison with the tv[-)icalK
dark shells tounil on Rliizoplmni trees, wliieli aic the
prevalent shell t\pe in tin- Inil<i-Mala\ and Anstialian
regions.

The new records listed abo\'e significaniK exli'iid the
known range of tliis species in .\rabia, the South ('liina
Sea. and southeastern ,'\irstralia. In |apan, the iccoid
from Shirahania confirms the (presumablv sporadic! oc-
currence of this species in the Kii Peninsula (the record
in Reid. 19Sfia. was baseil on a shell illustrated bv llabe,
1964). Additional collecting in northwestern .Australia
(M. Stucke\", personal communication! has jailed to (ind
tliis species hnther west than the (jjbnrg Peninsula, so
its absence from Western Australia appears to be real.

Litfdriiriii iLittdhnopsis) siihritidta Reid. 19S6a

Dislrihution: New record: Mast-arene Islands: Ro-
dngues iXMWM.

iJlttiidiiii \lAllttrhH>it\i\^i'hilij>j)uiii(i (Reeve. 1S57)

Dislribiilion: New lecords: New Sondi Wales: Wiioli
Wooli Rivei (A.\1S).

Littoriirid (LilloriiKijisis^ fill)-./: iSowerbv. 1S32)
(Figures 23-30)

Animal: Penis (Figures 2:3-30): in specimens Irom
Western .\nstralia the glandular limb is ol'ten longer and
more robust than previonsK reported (Reid. 19S6a: fig.
54): it mav be as long and up to 3 times as broad as the
penial filament.

Uistrihution: New records: Northern Territon: For-
svth Creek. F coast Joseph Bonaparte ( inlT i iUINI I >:
Western Australia: Berkeley River and Revvlv Island. \\
coast h'seph Bonaparte CnlF i B\IN 1 1 1.

Remarks: Beid il9SBa) noted that the geographical
form found from C.ipe l.evecine to I'Amouth (;ulf in

Western Australia dilli-red from the tvpical form on the
east and north i-oast of .Australia In more munerous but
less prominent prinian ribs and bv details ol color pat-
tern. Taxonomic recognition was not considered justified
since no anatomical differences were detected. New ma-
terial from throughout the rau'^e in Western .Australia
inclndiiiii the two listed above that fill in ;i liaji in the
known distribution! confirms the leporled shell diller-
ences. but adds some significant details. The tvpical car-
inate (eastern) shell tv])e extends to both localities in
Joseph Bonajiarle (inll listed above, whereas a large
sampli' (about 100, BMNIl) from (.'a[K' I,eve(|ue dis-
plavs characteristics intermediate between eastern and
western shells. The western forms olten, but not alvvavs,
have penes with an nnusuallv long and robust glandular
branih ol the bast', exceeding in si/e anv seen from
northern or eastern ,Australia (samjiles ol (50 western and
55 eastern penes*. In none ol the samples Irom Western
Australia or Noilhein rerritoi"v is there anv morpholog-
ical evid.ence lor a mixture of two discrete shell t\[)vs
with correlated penial dilleiences. The earlii'r taxonomic
conclusion therefore si'cms justified Irom a moipholog-
ical viewpoint, and the evidence appi'ars to show a
blending ol characters between two forms with a con-
tiimous distributioir rather than allopatn' or limited
overlaii between tvso disliuet lornis. However, this in-
tert'sting case should be investigated with genetic tech-
m(|U(>s (presentK being done bv M. Stuckev. Universitv
ol Western .Australia).

Till- a])|)arent trend towards <i longer glantlnlar branih
in the |)enis ol the weslern firm ol L. filosti might repav
Inrlhei- investigation. It could be significant that the east-
ern form (with rclativelv shorter glandular branch) is
broadlv svnpatric with /,. pliilijipidiiri. a congener vvitii
a much longei' glandular branch. In contrast tiie western
form (with rclativelv longer biancli is largelv sx'uipatric
with L. ciii'j^idatd. in which penial shape is more similar
to that of the eastern form oi' 1.. filosn (,see Reid. 19S6a:
figs 41. 58). If penial shape is a species recognition char-
acter, its variation within /.. filosd might suggest a case
of character dis]ilaceinent,

L'ltlonnid iLitloriiiopsis) aniodinidud (Ileude. 1S85)

Disliibiition: New records: Taiwan; Tan Shui. C'hiiia:
S:mva. Hainan \ielu:im: Sun Sot Cave. Ila Long Bav:
Fo River. Nha Trang. Singapore: Changi South Uill
BMNli .

Littordrid (Litl()ritioj)sis) (Iclicdiiild Nevill. 1885)

i Figures 31 -(5(5. (36. (37>

.Vniinal: Penis iFitjnres (51. (52 1 bilurc;ite; glandular
disc large, with thin m.ugiu; filament small. 0.2 total
length of base, separated from wrinkled base bv con-
striction: sperm groove open. Paraspermalo/.oa (Figure
.3(5:': 14-20 (iin. oval. 1 (rarelv 2) large rectangular rod
])iece filling cell: grannies large, lew. distinct (obsena-
tiou of single sperm sample piesened iu 8()9f ethanol.

Page 12fi

THE NAUTILUS. \ol. 115. \o. 4

33

20 ^m

D. G. Rckl. 2001

Faec

whicli max rausc shrinkatic nl' ahnut 20%; Hcid. 1996:
fi). N(i (lata nil Icnialc. Radula (Figures 36, 37): rclatixc
leil'^th (l.Tfi: I'achidiaii tontli: liasi- flared, wiitral I'lisp
sliield-sliaped, small cusp and one denticle on each sid<':
lateral tooth: 5 cusps, lai'j;est central iiis]i liluni: imiei-
marginal tooth: 4 cusps: ontia' marginal tooth: .") cusps.

Distribution: Habitat: loiiage ol Aiiccniiiii iilhii on
mudd\- riwr hank: water ol low salinit\' (from 2-3C{ In
wet season to 16,8% in dn season) (C. Feudred, per-
sonal coninninication). New records: Bodra Kaal. Cha\-
labogi, Sundari)ans, Bangladesh (BMMI': Krns:idai Is-
land', f:ulf of Mannar India (BMNIP.

Remarks: Beid (19S6al did not lia\e ;iccess to an\
preseni'd specimens, lint since then two males lia\e
been dissected and one radula prepared, leased on some
siniilarit\' oi shells, Reid (19'S6a) suggesteil that L, dcli-
catnla was most closeK" relateil to L (inhntiiiiund auil
that anatomical exidence w:is i'e(|\iired to confiini that
they were indeed distini't. The newK described pciiial
form is snfficientlv ditierent irom that ol L. (inhuiinknui.
in which the filament is large (up to hall total length of
penis; Reid, 1986a) to leaxe no doubt that the t^\■o are
separate species. The paraspermato/.oa are similar in
bodi. The penis of L. (Iclicatiila does not clitler siguifi-
cantK from that of L, juillc.sccus and the radnlae ol these
species are also closeK similar. Radular tooth lorm is not,
however, a reliable taxonomic charatter in Lilloniiiti
(Reid and Mak, 1999). Despite the penial similarih L.
delicatiila is beliexed to be ilistinct trom L jxillrsccns.
since the shells are \en difierent (delicate, with 1 1-14
primarv grocnes, 35-50 ribs on last whoil and n.urow
columella in L. drltciiliild: solid, with 9-10 priinan
grooves, no secondaiA' sculpture and broad columella in
L. pallcsccns: see Reid, 1986a). Paraspermato/.oa also
differ, being rounded with small rod pieces in L. pdllcs-
ceiis (although onlv a single sample has been seen in L.
delicatida). These hvo ha\e not vet been collected to-
gether, but both ha\e been recordeil trom the .\ndamau
islands (Reid, 1986a), so that the conclusion that tlie\
are distinct might be tested in future. There is a super-
ficial similarih' to L. hi'iiiirdciisis i see alxne).

The new record from the Clulf of Mannar greatly in-
creases the known lange of tills species, previously re-
corded ouK' from Port ( .'anulug (Bengal) and the ,\n-
damau Islands. It is likeK, however, that the centei- ol
distribution is the northern Bav of Bengal. Only single
shells are a\ailable from the .\ndaman Islands (BMNll:
Reid, 1986a) and the (hill of Mannar The latter was

collected in 1934 1)\ R. Winck-worth; the species was not
loimd on a recent \isit to the (hilf ol Mannar (personal

obseiAation. 2000 > and is presumabK rare in this area.

IJIIonihd I l'(ihi\toiind ' inchiiioslomd iCirav 1839)

l)istril)iiti(>ii: New records: |a[)an: Mixako Island:
V.ie\aina Isl.mds lligo et af, 1999).

Heniarks: Reid i i986a! recorded the rare occurrence
ol .1 pinkisli orange moqih in this uormalK monomor-
phie spi-cies. This li.is now been seen in two sam[)les
Irom localities in Ila Pong Ba\; Nii'tnam iBMNII): the
lii'i|ueiK\ w:islow (2% and 5% . N = 21 and 14 ' and in
I'ach case the specimens were collected among saltmaish
grass (see Remarks on L. rarinifcnO.

LitliTdrid iPdliislid-iiid^ idriiiifcrd > Menke, 1S.30)

.Shell: Two cases ol color poKinorphism ha\e been re-
corded Ml this species, hitherto thought to be rather con-
stant in coloration (Reid. lOStia'. In a samiilc ol 78 shells
fiiim Pembar Poinbok B.MNIP. most had a cream to
gre\ ground color with icd brown pattern, but in sexcn
shells l9'/fl tlie ground color was orange. The sample
was collected on tnmks ol dwarl Aiicciiuid and Soniicr-
dliti trees 1 in in height on ,i mudd\ loreshore. .\ sei'ond
poKinoiphic sample was collected among saltmarsh
grass 70 km south of Raiiong. 4'liailaiid (BMNTP. Here
the shells were of the small, sniooth form (e.g. Reid.
1986a: fig. 821); the range of colors was similar and three
(6.5'7f ) of 46 shells wt're orange.

Distriliiilioii:

bok (H.MNIP.

New recortl: ludonesia: Pembar Pom-

Kcmarks: Plsewhere in the subgenus I'dlu.sloriiid
pinkish oiange shells are \<'|-\ rarcK cnc-ountered in /-.
articuUita and /. dulditostdind. although this inorph is
wideK distributed i though alwa\s at low frequent')) in
poK nioiiihic members of the subgenus Lill(>riiH>i)si\
Lillonirid cdnuifcni is normalK found on trunks at low
le\els (up to 0.5 iiii on trees in the landward Iringes ot
mangrove forests i Reid. 1986a: 192) and shells Iroin this
habitat are of overall dark brown color and not poly-
morphic or varialile. It max be significant that both (loly-
morphic samples are from foliage in open suiinv condi-
lious. in otliei' woi'ds from visnalK varied niicrohabitats
where the animals might be exposed to visual predation
(e.g. bv birds or crabs). It is under exactlv these condi-
tions tli:il the polvmorphic species ol the subgenus lAt-

Figures 23-33. Penes anil paiusperinatnzoa of IJllnrnriri fihisti and lAllonina dclicatula. 23-30. I'l-iic's ol I., jilosd. 23-26. Jiilago
Beacli. Cape Peveque. Wc-steni .Australia iB.MNll 201)11)120: shell II = 2().S mm, 20,S mm, I9,S mm, 18.7 mm). 27, 29. Lookout
Hill. Broome, Western .Australia B.MNII 20()1()I2I; slicll II = 19.7 mm. 15. S imiil. 28. Mangrove Point. Roebuck Bay. Broome.
Western .Australia (BMNll 20010122: shell II - 21.1 niiiii 30. Witliiiell Bav. Burimp Peninsula. Western .Australia iBMNII
20010141: .slu41 H = 1S..3 nmii. 31-33. Penes anil paiaspemiatozoa of/.. dclUiitulii: Bocira Kaal, ( :Iiavi;il>o<;i. Simdadians, Bang-
ladesh (BMNH 20010123; shell H = 14,4 mm, IP') mm). Dotted line indicates extent of opaciue glandular portion of penial
glandular disc.

Paw 12S

THK NAUTILUS. Xol. 115. Xo. 4

D. C. Hrid, 2001

Puee 129

toriitiipsis occur and for wliicli \isiial sclccfiDu l)\ pred-
ators has hccn suggested as a possi!)ii' explanation for
llie maintenance of the poKinoqiliisni. wliicli is assumed
to have a genetic basis (Cook, 19.Sfia, 1), 1992; Held.
19S7). Tlie same process could possihK maintain these
rare, local cases of polvnioqjhisni in the noiinalK niono-
moii^hic L. carinifera: a similar explanation has lieen
proposed lor local poKnioqihism in L iiilcniicilia h\
Cook and Bridle (1995). Ho\\e\er, it should he noted
that like almost all others in the genus, this s]iecies has
planktotrophic dexelopment and therefore the potential
for wide gene How; selection on local populations would
therefore ha\e to be strong, or gene flow would luixc to
be restricted in some wa\, in order to achievf local gi'-
netic differentiation of pohnioiphic populations. .\ sim-
ilar case of local poKanoqihism has been lound m L
iiwlanostoma (see abo\e).

L'lttonihd {P(iltisti)n)ia) sinensis iFhilippi. IS47)
(Figures 3S-.52. 5fi-fi4, 6S)

Litoriim sinensis Pliilippi. 1S47: 16-17, Litorina pi. fi. fi'^. 2.3

(China; lectot\pe I Reid. 19S6a) MNHN -I- 2 paralectotx -

pes, seen. Fissures 42, 43). Lischke, 1S7I1); 71-72. Weiii-

kauff, 1S82: .S3-84, pi. 11, figs 9, 12.
Littorina intermedia var. sinensis — \e\ill, 1S.S.5: 147.
Littorina sinensis — Pilsbuiy, 1895; 62.
Litorina strif^ata Lischke, 1871a: 148-149 (Nagasaki. Japan:

t\pes lost: not Littorina intermedia \ar slri'j,(ifii I'hilippi.

1846 = Littoraria strii:,ata). Lischke. 1S711); 7:). pi, .5. fig.

22 Uiot Philippi, 1846).
Littorina stri'^ata ( Lischke i — Pilsbr\. 1895: 62 (not I'hilippi.

L846I.
Littorina {Meliirajilie^ strif^ata iLischket — Tnon. 1887: 245. pi.

43, fig. 33 (not Philippi, 1846 1.
Littorinopsis stri<s,ata (Lischke) — Kuroda and ll.ihe. 1952: 61

(not Philippi, 1846). Ovama and Takeniura. 1961: fit;, 10

(not Philippi, 1846).
Littoraria slri<iata (Lischke) — Kojima, 1958 (cgii capsule: not

Philippi, 1846). .\zuma, I960: 10 (not Philippi, 1846).

Higo, 1973: 46 (not Philippi, 1846).
Littoraria stri<iala CDiniker') — Yoo, 1976: 56, pi. 7. fi>j;s 18, 19

(not Philippi, 1846).
Littoraria seahra stri'^ata (Lischke) — Higo and ( ioto. 199.): 71

(not Philippi, 1846).
? Littorina adonis Yokoyama, 1927: 451, pi. 51, fig. 8 (Koyasu.

southern Musashi, Japan: Upper Musashino [Pliocene:

Rosewater, 1970]: holotxpe. Geological Institute, Univer-

sit\- of Tok-\-o: not seen). Rosewater 1970: 45.3. pi. .349,

figs 8, 9.
Littoraria adonis — Okutani, 1986: 71, fig. vninumhered.
Littorina i Littorinopsis) seahra seahra — Rosewater. 1970: 4.56-

461 (in part: not Linnaeus, 1758 = Littoraria sealira).

Littorina seahra — (^ai, 1991: 51. H'i. 43 mot Linnaeus. 1758).

Littoraria yPahistorina) ariieulata—l\cid. 1986a: 21)0-209, figs
90c, e, 92h, i (penes), 93c (egg capsule) 94 (in part; not
IJttorinii intermedia var ariieiihila Philippi, 1846 = Lit-
loraria ariienlala). Clioe, 1992: 289-290, fig. 54 (not Phi-
lippi. 1S46). Reid, 1992a: 195-197, fisJ. 4g (in part; not
Philippi. 1846). Hiiro ct al.. 1999: 92. '

Vdlislinina (sic) ariirnlata — Fnkuda el ;il.. 1992: 57, pi. 9, figs
!29ii. h (not Philippi, 1S46),

Nomenclature: The uonjenc lature ol this specii's has
snlfered numerous elianges. 'i"he name Littorina sinensis
li;is seldom been used, most recentk' b\' Filsbn (1895).
It w;is redescribed as Litorina striiifda b\ Lischke
(1871a) and this name bei'ame familiar in \arious com-
binations in the |;ip;mese literature ol tlu' mid-tx\eutieth
eeutm'x. Hosewiiter i 19701 coml)ined it with some 20
other speeiis nndei- the name Littorina seahra. The Pli-
ocene fossil Littorina (uhmls has not lieen examined, but
is probabK this species, and the name was used for Re-
cent |a])anese s]iecimens l)\ Okutani (19S6). It lias be-
come genenilK known as Littoraria artieidata following
Heid ( 19S6ai, but that taxon is lii'i-e shown to consist of
t\\() spei-ies. .-\s discussed below, tlie onK entirely diag-
nostic character is the penis. ;illhouij;li the majority ol
shells can be coufidentK identified, .\lthough tlie lecto-
t\pe is a (In shell, the form of the .i])ertui-e ;ind colu-
mell;i, anil the diliuse eoloi- pattern on die shoulder re-
'j;ion, are txpical ol the present speiies.

Material E.xamined: 'r\])es as indie:iteil: 44 lots; .35
penes: 1 sperm s;im[ile: 12 palli;il o\iducts: 5 radulae.

.Shell (Figures 42-52): Adult size nuige 6.3-20.6
unrr Shape high-turliinate to elongate (H/B = 1 .28-
1.73; SH = 1.57-2.05); whorls rounded, suture im-
pressed, periphcn of last whorl slightly or not at all an-
gled; motlerate tfiickness. .\Iatm-e lip not flared; colu-
mella pillar conca\c, excavated; inner lip of aperture
shaipK raised adj;icent to base ol colmnelia. .Sculpture
,,f (7)8-9 priman spinil giooves on spire whorls; inter-
\cning ribs usu;ilK rem;iin imdi\ided, numbering 20-23
on hist whorl; occ;ision;ilK some ribs may be divided by
nil impressed line ;iud become more numerous: ribs of
e(|nal \\idth above periphen, slightly narrower on base;
groo\es MO impressed lines on spire whorls, becoming
deeper and wider on last whorl, commonly up to hall
width of intent'uing ribs at perijilKU-N- of whori (rarely
ei|u;il to rib width); in smootliest shells grooves renuiin
;is impiessed lini's ouK. .Microsculpture of taint spiral
stii;ie over rib surface, with iLxial microstriae in grooves.
I'rotoconch 0..15 imii di;mieter. 3.5 whorls, with spiral

Figures 34-41. Railulae ol Littoraria species. 34. 35. /, hen-alensis new species: pan.txpe: Hare Island, Cull of Mannar.
Taiiiiliiadii India (BVINH 20000755: shell H = 16.3 nnii: Hal viiw and 4.5° from anterior). 36, 37. /.. deheatnia: Bodra kaal.
Chavhihoai Sundadmns, Bangladesh (BMNIl 20010123: shell II - 1 1. 1 nun: fk.t view and 4.5° ironi ;intenor>. 38-41. /-. smens,s^
.38 39 \hiu<n-ove trees Tan Shiii Taipei. Tiiwan ( BMNH 20010130; shell II = 111 iniii: flat view and 45 from antenor). 40,
41. Rocks, Kaluri, .Maelnin. Citv. Kviishn fpan B\l\ll :0()10I31: sl„ll II = 15.5 iiini: Hat view and 4.5° Ironi anterior'. Scale
bars = 100 (xni.

Page 130

THE NAUTILUS. Vol. 115, Xo. 4

D. C. Rfid, 2001

Faaf 131

rihs and siniisigera notcli. Color xarialilc: cicaiii witli
pattern of orange brown to dark hrown tlaslus on lilis.
witli whitisli dashes between; degree ol axial alignment
of dashes \aries Ironi ditfnsel\ tessellated or niarbleti
pattern with alignient into short axial stripes at sntnre
and periphen- onK (Fignres 43. 44. 51 > to the lare more
complete alignient in oblique axial stripes . I''igme Kit:
pattern often emphasizes 2-3 ribs at ptM'iphen. (Jilii-
mella puqile brown or white: aperturi' tiivmi with ex-
terior pattern sliowing through.

Animal: Headfoot. operenlum, paias[U'rmato/()a. pall-
ial ONiduct and radula do not differ signifieantK Irom
those of L. aiiiculafa as described b\- Reid il9Sfiai. .As
in L. ai-tiaikita (Reitl and Mak, 1999: fig. 4(:, D^ the
radula shows likeK phenotxpic plasticitx and differs in
tooth shape on rock and xxood substrates (compare Fig-
ures 39 and 41). Penis (Figures 56-64) not bilurcate.
small glandular disc incoiporated into distal end ol w lin-
kled base: smooth narrowlx' elongate filament 0.3-0.5 to-
tal length of penis: penial \as defererrs an open grooxe
to filament tip. Pelagic egg capsule a lens-shapi'd bicon-
\ex disc without peripheral flange, 350 |xm diameter,
containing single oxiun about 70 |jLm diameter (Kojima.
195S: fig. 2a, "b; reproduced in Reid, I9S6a: fig. 93c ».
De\elopment planktotrophic.

Distribution: Habitat in littoral fringe on rock (in-
cluding granite bouklers and concrete sea walls) in both
sheltered and moderatelx- exposed situations: .ilso on
trunks at seaward edge of mangrox'e forests (publisla-d
accounts of the ecological distribution of 'Litfohnii sea-
bra' on the rock-x- shores of Hong Kong and southern
China probabk refer to a mixture of this species and L
aiiiculata, e.g. Chambers, 1980; Ohgaki. 19S5: Yi and
Li, 1988; You. 1990). Range (Figure 68) from southern
China to Yellow Sea, South Korea, Kxiishu and Seto In-
land Sea, perhaps Rxuk-xii Islands. Records: C^hina: Ma-
cao (BMNH); Hong Kong (Shek O: Deep Ba\: Tai Po;
Three Fathoms Cove; Hoi Ha, Mirs Bay; W n Kwai Sh;i;
Aberdeen; all BMNH); Xiamen (Hai C'ang: Culayang:
both B.MNH); Spider Island, Fujian (USXM); Zhenhai.
Ningbo River (BMNH); Shawai.shan. moutii of Yangtze
Rixer (BMNH); Qingdao (Tai Phig; Huan Dao; Ilui
Quan; all B.MNH). liiiwan; Tan Shui (BMNH). Japan:
Rnikxai Islands (USNM); Kago.shinia (BMNH); .\mak-
usa '(USNM, NSMT); Matsuura, Nagasaki Pref.
(BMNH); Hirado (AMS): Maebani Cit\ (BMNH): Fh-
ime Pref. (USNM); Kasaoka, Okaxama Pref (LiSN.M):
Nagae River, Okaxama Cit>- (BMNH); Yoshino Ri\er.

'iokushima ('it\ iBMNIJi. South Korea; Sacliun Kun.
K\oni£sang \:imdo i.\NSi';.

Hi-niarks: Ri'id if986a) concluded that L. aiiiculata
was a single s])ecies extending from .Australia to India
and Japan, but with two geographical forms of more re-
stricted distrilintion. one from norlliwestern .Australia
and the other from ( :hina and |apan. The latter was
characterised b\ stronger sculpture, a more diffuse pat-
tern and it was said that the penial filament xvas 'some-
times relati\el\ longer than in specimens from South
East Asia and .Australia'. It was also noted tliat the egg
capsule ol the j.ipanese lorm di'scribed b\ Kojima
i 1958) was largei- tlian that of 7^. aiUnilata from Queens-
land and lacked the circumferential flange. Howexer,
with little inloiination on tlii' range ol intraspecific var-
iiition the significance ol the difference in capsule shape
was unclear The easteiu .\sian form was 'tentatixelv as-
signed to /,. aiUciilatti' . but it was suggested that further
work was rec|uired.

Since then, much additional material has been gath-
eri-il. It has been found that ;ill specimens from northern
China, Korea and |;ii)an shaic the longer penial filament.
.Most significant, however is the discoxen- that the t\p-
ical form of /., aiiiculata and the eastern .Asian 'form'
occur sMitopic.ilK at sc\cial localities in southern C'liina,
around Hong Kong and .Xiamen. .At these localities the
[lenial differences remain distinct (e.g. Figures 57, 58
and 65. (i6. from a localitx- in Hong Kongl and are cor-
related witli small, but consistent, differences in shell
shape, sculpture and color This leaxes no doubt that the
'forms' are separate species.

The tlifferences between /., aiiiculata sciisii stricto
;ind L s/)i((/s7s ;ne summarised in Table 2. The most
useful ami entircK diagnostic character is the shape of
the penis; in /, sinensis (F"igures 56-64) the penial fil-
.iment is 0.3 to 0.5 of the entire penial length (less than
0.2 in L aiiiculata. Figures 65-67) and the glandular
disc is nsnalK about half the size and less swollen. No
other .m;itomical differenc'es haxe been discovered, so
tluit moiphological identification of juxeniles and fe-
uKiles nnrst rel\ on leatures ol the shell.

it is possible that the differences in egg capsule siiape
mentioned ,ibo\e ma\ prove to be consistent. Capsules
in this group have been illustratetl three times. Reid
I 19S6a) showed cajisnles of L. aiiiculata s.s. from north
Oueensland with ;i circmnferenlial flange (diameter
248-268 txm, ovum about ti5 fji.m). Those illustrated bv
Mak i 19951 bom Hon<i Kong were of similar shape, but

Figures 42-55. Shells of Littoraria sinensis a.,.l Utfoiyirin aiiienlata 42-52. /., sinensis 42. 43. l,ect<.tvp<- of /.«,.,-/„« siiiciisi^
(Philip™ 1S47) Cliina iMXHX!. 44. Hoi Ha. Mirs Hav. Hon- Kon.4 > BMNH i'«)l()124). 45. Tai l^in-.; Hay. gin-.lao. (_l,ma ( li.MNH
'■'00101-''5) 46 Tii Po Mono Kon^ (B\I\I1 2()01O12(S' 47. Voshino Kivcr cstnan. Kaiuizavva. lokusliniia (.i(v. Sliil<()l<ii. Japan
"bMNH 20()1()127) 48. Shianzin. Shanij,,. China ■ B.MXII 2()()1()12S.. 49. llai Cang. Xiamen. China i BMNH 201)10129 . 50 Tan
Shui Taipei Taiwan (BMNH 200101.'30i. 51. .Ahc-rcken. llon..j Kong (BMNH 2001()132>. 52. Kalnri Maeharu tatx. Kvnslni JapaT,
(BMNH 20010131' 53-55. L. artunlala 53. T.i Po. lion- Kon-BMNll 200101:5Si. 54. 55. .M«-rdeen, Hong Kong (BMNH
20010137).

Paw 132

THE NAUTILUS. Vol. 115, Xo. 4

D. G. Reid, 2001

Wm- 133

Table 2. Sniiiirian of dittfreiifi-s l)ct\\c(ii IJtlnnind sido/.s/s and /, iirliiiiliil/i isiirll ciiarac-tcrs appiv to spci-iinciis from iiortli-
wfstt'ni Pacific Ocean ()nl\ I,

Cli.iradn

l.illiii'diiii simiisi'.

Litttimha arlii-iildid

Shell sliapc Hclati\c spire lieiij;lil can !«■ taller to 2.05!:

whorls sli'^htK more roniideil

Shell Miilplnre (lroo\es up to hall to one times ril) widlh; small

sliells can I le almost smooth

Columella Slioiter ,md more concaxi', ^i\inij more o\al ap-

erture: Uiirrowcr, nmer lip of :iperlure more
siKU'pK raised

Sliell color I'alcr. often indisliuilK ilouded p.ittern: align-

ment of dashes less marked

Penis Penial filament ()..3-().5 total lensith of ])i-iiis

Distribution (!liiiia, Korea, [apaii

Does not attain such tall spire irelati\e spire

heinht to 1.S9I: whods Halter
(u'oovfs up to one qnarl<'r ril) widtli: small

shells can he almost smooth
f.oiiijer. straiiihter. jii\ini; ear-shaped aperture:

iiiiiei lip III aperture less sliarjiK raised

Darker, distinct dashes, well ali<4ned into axial

stripes at suture and periphen
Peiiiai hlameut less than 0.2 total leii'^th of penis
Southern (Miina to India and Australia

larCTei" (diameter 300-340 |xni, oMim 69-71 |j.iii). The
capsule ot L. sinensis troiii Ainaknsa, KxTisliii, siiowii h\
Kojima (I95S) was a simple lens shape witlimit ;i |)e-
ripheral flange (tliameter 350 [xin, cnxim alxiiil 70 fxiu).
Mak (1995) showed that in three Nodilitti'iiud species
egii capsule diameter \aried l)\ 39 to 53% and sn<j;i.^ested
that shape and sculpture are more consistentK diai^nos-
tic of species tlian size. It is possible that the capsules
he illustrated as I. (iiiinihilii' were indi'etl ol tliat spe-
cies rather than the sMiipalrii- L. sinensis, despite their
lars^e size, so that shape mi^lit he relatist'K constant
within tills species across its <j;i'ot;raphical ran<j;e. but this
remains to be iinestigated. (Caution is necessan, how-
ever, since capsules swell and the flany;e becomes less
noticeable bet\\'een spawning; and hatching (Berr\, 19S6,
in L. strifiafa: Ho. 19S7>. The capsules sliowni by Reid
(1986a) and Kojima (195S) are veiA' similar respecti\ei\
to the early and lute (24 hours after spawnintj;) capsules
of L. stri^ala (Berrw 19Sfi: fig. 1) anil might not lepre-
sent genuine interspecific tlilferences.

The characteristic features ol the shells ol /.. sinensis
and L. aiiienlata are diffic'ult to describe or ijuantit).
owing to considerable intraspecific \ariafion. \'e\"erthe-
less. with experience sxinjiatric samples can usually be
.successfullv identified (as confirmed In penial shape ol
males) from tluir shells. The size range is similar (max-
imum 20.6 mm in L sinensis. 19.0 mm in /,. nrlienlnla'
and both attain larger si/e in niangroxc habitats than on
rocky shores. Oxerall shape is also similar i)ut cxaiuples
of L. sinensis can attain a sliglitl\- taller s|iirt' (ndatixc
spire height up to 2.05. cf. I.S9 in /.. niiienldlni and the
whorls are a little more rounded. In L. nrlienlatd the
columellar pillar is longer and straighten so that the ap-

eitnre appears more ear-shaped, in contrast the colu-
iiK'lla oi /, sinensis is sliglitK shorter and a little con-
cave and llie aperture therefore more nearK' oxal. The
ciiiiiiiirlla III /,, sinensis is usiiallv narrower, but still
deijiK extaxaled, and the inner lip ol the aperture (ad-
jaceiil to the conca\it\ ot the columella I is more sharply
raised, sometimes so nmch so that a slight umbilical
crack is present i^oth species share a similar coimt ol
priman grooMs and ol ribs on the bod\ whorl, and both
sometimes ilispki\ dw;ui lorms with smooth shells in
which grooves are present ouK as impressed lines. How-
evei'. in large)' and more strongK scnlptin-ed examples
the grooves of /. sinensis are alwavs more pronounced
(up to half of, or even eijual to. width of intei-vcning
ribs. compcUed v\ith maximnm ol one ijuarter of rib
width in L. (irlienhitiii and such shells are mmiistakable
(Figures 46. 47).

Shell C'ojor and pattern are likewise dilficult to char-
acterise, but often assist iilentification. Ijttomria sinen-
sis is nsnallv paler. txpicalK with a more finely mottled
or tessellated or inciistinct ■elondeil' pattern, in which
whitish lii'cks are jiresent between the darker dashes. In
liiilli speiies the dashes are more or less aligned into
olilii|ne axial series, although this is nsnallv only pro-
nounced at the suture and periiihei-\ ol each whorl, giv-
ing rise to short axial stripes at these points. This align-
ment is often more developed in /,. ailienlafn. in which.
al the pi riplieiv. the axial stripes extend across four of
the spiral ribs (Figures 53. 54): in /.. sinensis this align-
ment more often covers onK two or three peripheral ribs
( {'"ignres 50, 51 1.

These contrasting sliell charaiters are emphasized by
tlie figures of svnipatric samples of the txvo species (Fig-

Fieures 56-67 Penes of IMIordria siuensis and lAtUvariii dr1i,uldla. 56-(>4. /,. snieiMs. 56. Tan Shui. Tiipei. Taiwan i li.MMl

20llll)f,3(l shell H = fl 0 I ' 57, 5S. .Shek (). lion;; Koriii di.MMl 20010133: .shell H = S.6 mm. 9.4 mm\ 59. Ilai Cang,

.Xiamen China (B\1NH 2(ll)IOf29- shell H = 9.0 num. 60. Ilium Dan. Oiu^cfio. China ilUIMI 20010134'. 61. Okavama Port,
Kojnna Bav lapan (B\l\ll 20010135: shell II = 7.2 nuu ' 62-64. Naijac Hiver .'sluai-x. Otogo. Okavama Citv. Japan dlMNH
2()0101.36- shell II = 13S mm 13.(1 mm. 1411 mm . 6.5-67. /. artienhila 6.5, 66. Sh.'k O. Hong Kong (BMMI 20010139: shell
H = 8.7 mm, 10.2 mm). 67. Aheixken floiK.; Kong .V,\\\\\ 200f0137). Dotted line indicates extent of opaipie glandular portion
of penial glandular disc.

Pii-'p 134

THE NAUTILUS. \ol. 115, No. 4

Figure 68. Geoijrapl ileal dislrilmlidii (il Ijllininiii sinensis with, toi' (.■ciiiiparisoii, tin- reeurds ot Lilt(n-cinii (iiiiculala Iroin the
same area (see text U)Y lists ol records ).

ures 46 and 53: Kiij;vii'es 51, 54, 55). Ncx'ertlieless, sliells
of tlie two can sometimes appear \irtuall\ identical (e.Q,.
Fisiure.s 44. 55) and onK- penial .siiape i.s reliabK dia'j;-
nostic. Cleo^rapliical localit\ can also aid identification.
since the distributions are known to overlap onl\ lie-
hveen Hong Kong and .Xianu'ii (l''iii;nre fiS). In lionij;
Kong tlie sites of recorded s\ntop\ are Sliek O, Tai Po,
Tliree F'atlionis Cow. \\u Kwai ,Slia and Aberdeen, ami
Hal Gang near Xiamen (all B.MNIU. These sites range
from mangro\es to moderately e\i)osed gi-,iiiile boulders
and both species are common, so that tlieic is no evi-
dence lor dillerent habitat preferences

Litlomria sinensis is also closeK' similar to /. s7r/gr;/f;,
distributed from the Philippines and soiilhern \ ielnain
to Indonesia, Malaysia and India. The leadncs nl slicll
shape and coloration described above also distinguish /..
sinensis from L. stri<i(iUi; the latter often has a more
strongly aligned pattern ol obli<|ue axial stripes, and
sculpture is \yeaker tlian in L. sinensis. Most importautK.
the penial filament of L. slri'^ala is long, tapering and

vermiloiiu. U.H to ().. of the total penial length iKeid.
1986a).

Uflamria (Puhtstorimi) aiiiculata ii'liilippi. 1S46)
(Figures 53-.55, 65-6S)

Svnonviiiv as in iveid i 19S6ai, excluding the entries now
listed under /,. s/dc/is/s above.

Distribution: bceords listed in Heid (19S6a) from
China and |a|iau all either a[)[)lv to L. sinensis or the
identiheation is doubtful. The following new ri'coids i.ill
BMNIJi have all been verified bv anatomical examina-
tion: ( :hin:i: llai Gang, Xiamen; Hong Kong (Tai Po: .Ab-
erdeen: Sliek O: Three Fadionis Goye; Beigang Iskuid);
Hainan. Xii'tuam: Sung Sot C'ave. Ha Long Bay: Do Son,
near Haiphong. Gambodia: Koh Pource, oil Sihanouk-
ville. The sjieeies is abundant at localities all aroimd Sin-
gapore (St |ohu's Island: Gluingi Point: Seutosa: Fast
Coast Park: Sarimbun: all B\1\1L. whereas the siiuilar

D. G. Rekl. 2001

Pasie 135

speck's L. striij^dta is lan' (sec lirlow '. ()iil\ a siiiij;li' pir-
sened animal of tlic /,. (ii-ticiiliitd/slri^nla <j;n)iip has
been seen ironi tlie liuliaii snlicontinent; this sjiceinieTi,
from Bomha\. was a male <il I. \liiiuil(i (sci- hclow '. so
the occurrence cit /.. nrliiiilntu in the ict^ion remains In
be N-erified. A single shell has been seen Irom Banilar-
e-Khamir, west ol Bnntlar Abbas, Iran (BMNIli. with a
color pattern resemblint; L. aiUciihitd. This wonld rep-
resent a rantic extensicm ol either /, itiiiciiliilii or /.
sthoatd. but positive identiHeatioii i'e(|niri's aii.iUimical
examination.

Littoniria (Pf;///s^)r//K/'i .s/;7<,v//^/ (I'hilijipi. 1S46)

Littdi-iiri/i stri^cilii — l^ern. IVJSfi: 144-149, fii;. 1 le'w capsule i.

Distribution: additional material b'om .South Kast
.\sia supports the suggestion that this specie's predomi-
nates in localities with clearer water, wheii-as the similar
L. articulata is present in more turbid and estnarine sites
in the region. The oecurn^nce of this species in the ,\ra-
bian Sea lias been confirmed b\ a single male specimen
from Bomba\'. The species is imconunon in Singapore.
where L. iiiiiciihitn is abundant. Xew records lall
BMNH): Singapore: Sarinibnu: St lohns Island: Changi
Point. Mahi\sia: Kudat, Sabah. Thailand: K.rabi. India:
Colaba. Boiiiba\.

Remarks: Nine additional ethanol-presened sampk's
lia\c been seen, supporting the earlier conclusion that
this is indeed a species distinct from /. iii1U-id<it<i The
onl\" diagnostic moiiihological character remains the
shape of the penis, in which the filament is 0.6 to 0.7 ol
die total length.

.'\ckno\\i.ed(;mfats

This work was stinuilated b\ the conjunction of a nnm-
lier of events: the imexpected iliscoxen' of an undescri-
lied Littoraria species in northern .\ustralia b\ M. Stuck-
e\- in 1998 (Stucke\- and Reid, in picp.'. i|uestions from
If. Fukuda about the identification ol the |;ipanese L.
aiiiculdta' and mv own obsenation of assortati\'e mating
and dilferential zonation between forms' of L. iiifcr-
iiuihti' in south indi.r In :iddition. there was a growing
realisation that \en small but consistent dillei"enc(.'s in
penial form could be of t,i\onomic significance i iieid. in
press, a). Since 19S5 I have exarnineil much additional
material of Litlorar'ui. both persoualK colk'cted and
kindK sent to the b.MMl b\ colleagues. My grateful
thanks to those wlio ha\e provided specimens lor this
stud\-: G. Feulner (I^ubai). H. Fukuda lOka\:ima I'ui-
versit\), E. Gloxer (BMNTI), J). Kadolskx (Londoni, V-
M, \iak (Hong Kong), .\. Matsukuma i Kyushu L'niver-
sit\), C;. Pendred (Portsmouth^. M. Stnckc\ (University
of'W'estern Australia), j. H, Tavloi iBMNJi). \'. Heros
(.\INHN) kindK sent <i t\|)e on loan. I thank the Tropical
Marine .Mollusc Prograiunie of Dauida iDeuTuarki lor
the opportnuitv to earn' out fieldwork in Inilia.

l,rii:H\Tl 1{K CXVV.D

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THE NAUTILUS 115(4): 140-146, 2001

Wv'c 140

Phylogeny of pneumostomal area moiphologx^ in terrestrial
Piilnionata ( Gastropoda)

Ban-) Rolh

ncpartiiu-nt ot Iincili'hratc /.Dolot",-
Santa Barbara Miisciiiii ol \atiiral

HistoiT
Santa Barbara. CA 93105 USA'

ABSTRACT

Mappini^ pneumostomal area morphologieal eharacter states on
the ph\logenetic tree generated b\' a recent ribosom;il RNA
sequence stutK allows an independent estimate of the course
of pneumostomal area e\ olution and shows to wliat extent the
RNA sequence data supports evolutionary' events pre\iouslv
hvpothesized for pneumostomal area characters. A breathing
channel without a valve, an open rectal termination, a closed
secondare' ureter, and a simple pneumostomal exeretoiv route
are plesiomoqihic within St\lommatopliora. Multiple homopla-
sv is e\ident.

Additional kcij words: Mollusca, land snail, anatomx'. ph\lo-
genetic svstematics. parsimon\-.

INTRODUCTION

The molecular pli\l()genetic aiiuKsi.s ol Wade et al.
(2001) examined nucleotide sequences of the ribosonia!
RNA gene cluster of over 100 species of Stvlomniato-
phora. representing 50 families plus out-groups. That
stud\ supported the monoplivlv of Stvlommatophora
loiind 1)\' previous auahses (e.g.. Endierton et al.. 1990;
Wade and Mordan. 2000). Its other significant findings
include (1) a tuudaincntal dichotomv between an "acha-
tinoid clade" (including Achatiniilae. S\d)nlinidae. and
Streptaxidae) and all remaining ("uon-achatinoid") stx-
lonnnatophorans; (2) the monoplnk of nian\' traditional
family groups is su|)portcd; and (3) the orthurethran
condition, long presumi'd to be plesiomoiphic in Stv-
lonunatophora. is most likeK a deri\('d state. Wade et al.
(2001) onlv briefl\' considered morphologieal cliaracters
other than those tvpil\ing the traditional groups Or-
tliurethra, Mesurcthra, and Sigmurethra. Moqihological
correlates, if th(>\- exist. Iiaxc vet to be (lisco\ered for
many of the deep branches of their |)h\logenetic tree
(Wade et al., 2001: fig. I). For exam]ile, there appears
to be no extant, single moi-phological character state that
is autaponioiphic for eitlier the "achatinoid clade" or the

' Address for correspondence: 7'1.5 Cole Street, San p'raneisco.
C;A 94117 USA.

"non-achatinoid clade (P. Morilau. personal conmumi-
cation, 2001). It is intrinsicalK interesting, for under-
standing the histon and mode of pulmonate e\olutiou
and to increase the liaison bet\veen molecular and mor-
phological data sets, to plot the cfistribution of moipho-
logical character states on the plnlogenetic tree gener-
ated b\' molecidar analysis.

Goodfriend (1986), examining the lower coni'se of the
secondaiA' ureter in .Sr/gr/c/ Beck. hS.'BT. obser\ed that the
ureteric region of the pneumostome might pro\e to be
taxonomicalK usefid. Emberton (1991) included several
pneumostomal area characters in his ph\logenetic aual-
\sis ol 17 subfamilies of Stvlonnnatophora. Su\"oro\
(2000) studied the pneumostomal area of terrestrial pul-
monati' gastropods and pnnided an analwsis ot four prin-
cipal characters, each ha\ing from h\o to sexen mutualK'
exclusixe states. He described tlie distribution of these
character states across 34 families and Inpothesized
transformation series, based on a process of functional
anaKsis. There is moderateK good overlap betxveen the
array of taxa examined bv Suvorov (2000) anti that ana-
l\y,ed bv Wade et al. (20()1).

In this paper, the pneumostonud character states de-
scribed l)\ Su\oro\" (2000) are mapped on the |)h\!oge-
netii- lianicwork generated b\' the Watle et al. (2001)
stuck. This allows an independent estimate of the course
ol |int'uinostoiiial area exolution and shows to what ex-
tent the RN.\ se(|uence data supports the e\olutionar\'
iiderences ol Sn\iiid\, EssentialK the same method was
emploNctl b\ Roth iUJOO) to estimate the liiston nuder-
King the absence of dart sac and other reproducti\e or-
gans in several genera ol I lelmiiitlicigKplidae.

M.VIERI.VES .\M) .METHODS

Taxa: Table 1 lists the st\lonunatophoran species stud-
ied b\ Su\oro\ (2000) and their famiK assignments. Su-
\oro\'s results aiul iliscussion are presented in tei'ms ol
lamilies ratlu'r than species; the implication — ;iltliough
it is not explieilK stated — is tluit he lound pni'niiiostoni;il
;ue;i eluuMcters to be constant within a lamiK.

Nearly all oi these families were inclndi'tl in the mo-

B. Roth. 2001

Pane 141

Table 1. St\l(iiiiiiiatcipli(iiaii species stiulieil li\ Sii\()i"(i\
(2000), and taxon codes useti m this stud\, TiixononiN and oiik'r
of ta\a as in original, except as noted. Families williout indi-
cated codes are not eladistiealK anaK/.ed lieiriii

Succineidae (SUC)

SuiTiufii jmtris (Linnaens, 17.5S)
Oxiiloiiiii Mirsi {Esniark. LS86)
Siiaiiii'lld (ihlou'^d ( Drapaniand. ISOl)
Partnlidae (PAR)

Partula otaltcitdiui iBniguiere. 1702
Clansiliidae (CL.\)
Macroiifistnt icnihcosa (Drapaniand, LSOl)
Mviitissii '^nicilicostd (Rossniiissler. 1836)
Scrohiffi'/i tiitiricd (KiMiicki. 1837)
Scrmlina scrnilalii ( L. Pfeiffer. 1847)
Subnlinidae (SUB)
Sul>uUiui (K-tonci (Brngniere. 1789)
RunuiKi (Iccollatd (Linnaens. 1758)
Gh'ssuld ccijhmira (L. Pfeiffer. 1845)
Fenissaciidae

Auijihoivllti inchnnjioUlcs (Lowe. 18341
.\cliutinidae i.-VCHi

Achdtiiid ftilicd (Bowdich. 1822)
Orthidicidae (ORT)

Liffius fascidtiis (Miiller. 17741
.^cavldae (ACA)

Acdviis jiluH-nix (L. Pleitter, 1854)
Sagdidae (SAG)

S«;i(/r/ ciHikidiid (Cinielni, 1791)
Ceriidae (CER)

Ccrion iduiuid (Bnigniere, 1792)
Cochlicopidae (COO
CticidUopd lubriai (Miiller, 1774)
Cocltlicopa niteiis (Gallenstein. 1852)
Orcnlidue (ORC-1, ORC-2)*
Sjthiiniiliuiii doliohnn iBniguii're. 1792)
Enxinoldiirid zonifcrd (Pilsbn, 1934)
Ldurid ci/liiulrdccd (Da Costa, 1778)
Chondrinidae (CHO)
Clumdriiid Hiodid tdurica (Kessler, 1860)
Choiulhiid clicntd ccdicd\ii(i Klirniann. 1931
PvTaniidulidae (PIT?)

Pi/rdinkhda nipcsths iDraparnaud. 18(11)
Eni'dae** (END
Rdchis ttdcarcnsis Fischer-Pii'tte. 1964
Mcrdip^cra olKcuni (Miilk-r 1774)
Eiid inoiitdud (Drapaniand, 1801)
Bn'j)hdl(ipsis ci/liiidncd (Meiike, 1828)
Cidicdsicola I'dddi'i Kohelt. 1880
Brad\baenidae (BRA)

Bnidt/lxii'iid fniticiiid (Miiller, 1774)
Hvgroniiidae (If\'G)
Xcnipictd dcrbcntUid (Knnicki. 1836)
Xcropicta kninickii (Knnicki. 1833)
MtiDdcliiiidcs viciiKi ( Kossiniissler, 1842)
Helicodontidae

Ocstophiiru l)dHtdlii { Hossniiissler, 18.38)
Helicidae (HEL)
Ci'pdcd lindiihotwnsis (Fenissac, 1821)
Helix dUiisccns Rossniiissler. 18.39
HelniintliogKptidae

Hclmiutlio^hiptd armsd (W. O Binnew 1858)
Hninholdtianiilae
Hnmlxildtidiid sn.

Tahlc I . ( )ontnined.

1„ Pleiffer. 1842)
185()i

SpllilK teroillllul.ie

SpliiiK ti'i'Dcliild idiididissiina Dr.iparn.iud, 1801 )
Lpiphr.ignioplioridai'

Epiplird'^moplwra dihilii
l'",ndodontitlae lEND'

Hiillncnid fldlliidlidii 1. I'leiller
Disci(hie (DISi

Disiiis nidcnihis (Stiuli'r. 1820)
PoKg\ridae (POLi

Vdlijllipd -iiptidii till d 8a\, 1818
Zonitidai' (DAll. in p;n't)

Oxi/cliilw- iniiiiS^rrliciis (Monsson, 186.3)

Oxi/clnlus ddhiiisi (Moiis.son, 1863)

Ac'fiipi'i iciiicillus (Lamarck, 1822)
Dandeliardiidae (D.'VU. in part)

Iiif^urid tiiii^iiiii iKosen. 1911'
P;n'maeellidae

Vanndcvlld ihcrti Licliw.iid. 1S41
(Jastrodontidae

'/Aiiiiliiidcs nitidus (Miiller 1774)
.\nopluintidae (ARPi

Hcuscuifs lu'palt'itsis i Blanlord. 1904)
Ilaplotrematidae (IIAP'

Hdplotrcdid idiiiimuui i.\nce\. 1888 i***
Limacidae (LIM)

Liiiidx iiidciddtns I Kali'niczenko. 1851)

Biclzid idcnddiis iBiel/. 18511
.\rioiiidae (.ART

Arioii hisildiiivus .Mabille, 1868

Arioii fusridtiis (Nilsson, 182.5)

* Orcnlidae siiisu Su\oro\ (2000i is polypli\k-tic: ()R(:-2 rep-
resents a clade comprising Lduria species, assigned to Pnpil-
lidae b\ Wadeet al. i2()01 >.

** Enidae is called Biiliminidae In Wade et al. 2001 .
*** Sn\oro\ (2000: 91) witliont jnstification introduced tlie
couibin.itiiin Aninimitd iiuniiniiin lor this t.Lxon.

Iciiilar .iiiaKsis cl Wade et al. i20()l; 'lahle 1 :. ahhoilgh
ill iiian\ cases witli (lilTcrent species as exemplars. Based
OH the Hndiiig tiiat most traditional family groups hold
togetlier as monoph\letii' units in molecular aiialy.sis
(Wade et a!.. 2001; 4is). 1 iiere proceed on the a.ssump-
tion that — with a few exceptions noted below — tiie hun-
ilies of the Sn\oro\ and Wade et al. analyses can be
treated as e(]iii\ ;deiit, and that eharaeter states ob.serxed
to he true for a famiK l)v Sinorov can be regarded as
tnii' lor the same nominal family represented on the
nlnlogenetic tree of Wade et al. This assumption can. of
course, be tested 1)\ dissection ot tlie Wade et al. ex-
emplar species that were not examined by Su\()ro\.

[■"igure 1 ri'plicates tlie branching structure ol Wade
et al.'s (2001: fig. D neighbor- joining pli\logenetic tree,
with respi'ct to the taxa studied by Su\()ro\. The tree is
Imther simidified b\ being drawn as a cladogram rather
than a pli\lognnn; in other words, branch lengtlis, whicli
in the origin;d reflect genetic distances among taxa. have
no significiuice in this figure. On the oiigin;d tree, some
lietails of tlie first- and second-order branching direct!)-

Page 142

THE NAUTILUS, \ ol.

15. Xo. 4

i

1 —

1 ORC-1

^ PYR

one ''

Orthurethra

1 L!M

^ ARP

Limacoidea
sensu lato

'

1 — DAU

—

Al-ll

1 CLA

"non-
achatinoid

clade"

i END

1 HEL

1 POL

Helicoidea
sensu lato

' HYG

"""

—

—

1 — sue

"Clade B"

1 ORT

1 ACH

l"achatinoid

SUB

out-group

clade"

Figure 1. ( lladniirain liased on tlic iiiolcfulai' pli\liit;i-iictic
lupothcsis III Watli' ft al. (2001 ). simplified to iiR-lude onI\ t;i\a
studied In Siuoiciv l2(X)0). ;uk1 sliowiiia; tlie extent ot liii^lier
clades refenvd to in text: ■aeliatinoid clade" and "non-aehati-
noid elade" of Wade et al. (2001): Oithnrethra Pilsbn, 1900:
Limacoidea .st'iisti lato ol Hausdoit I 199.S); Helicoidea sens\i
latti and "(Made 15" ol' Roth and Sadeghian (in press). Taxon
codes as in 'I'alilc 1 .

aboxe the "acluiliiuiid - noii-aciiafliinid (licli(itniii\ aic
.supported li\ rclaliscK lew cliaiaclcrs ami iiia\ not he
corrolxirati'd lis lutiire studies.

Tlie Idildwiiitr ta.xa coiikl not lie [ilaecd on the cla(l<i-
grani: Epiphraginophoridae. Feinssaeiidae, (iastrodon-
tidae. Helieodoiitidae. HeliiiintliogKplidae. Hmiiliold-
tiaiiidae. Pannaeellidae. and S|iliiiKter-oeliilidae. Wade et
al. (2()()i) ineiuded "HelmiiilluigKptidae. ' liiit their ex-
emplar, Mouadciiia fidcHs (Cyax. is.'34). is a s|ieeies ol'
Bradybaenidae. Both Su\oro\- (2()()()) and Wade et al.
(2001) treated Oxijriiiliis Fit/.inger. 1S33. as a genns ol
Zonititlae. In liis phylogenetie anaKsis (if Limacoidea
scu.sii Idlo. ilansdorl' (199S) placed ()xi/tliihi\ in Daii-
dehardiiclae, and the cla(l(> exeiiqilified In il is so laliek'd
herein.

Wade et al. (2001) included no Sagdidae. I'lmlierton's
(1991) eladistic analysis rexcaied a elade comprising
Tlnj.s<iii(>j)li(>ra .Strehel and Pi'effer, ISSO (Thxsanophor-
idae): the New World 'camaenid " Plcunxloiitr Fiselier
von Waldlieim, ISOT; and the sagdid genera S/i<i(lii and

Aijuaclnnui Pilslii-\, I92fi. These ta\a are united in the
synapomoiiihies ol rigiit parietal and \isceral ganglia
fused and right parietal and right pleural ganglia fused.
Fleurodoitic and the sagilid genera are further united h\-
the sxTiapomoiphy of left parietal and left pleural gangUa
fused (Emberton, 199L 218). Based on tlu'se findings,
Sagdidae is tentatively placed on tin- cladogram in the
position of New World "Camaenidae," exemplified in the
Wade et al. anaKsis In Polydontes .Montfort, ISIO.-
Henvever, Goodfrieml ( 1986) obserxed that the distinc-
tive features of the sagdid pneuinostomal area are not
shared by Pleurodontidae such as Lahiihullin^ Beck,
1S.37, and Plciinxliuitf.

Wade et al. !20()1) included no Endodontidae; tiieir
:inaKsis resoKed an "endodontoid clade" consisting of
mi'inbers of Punctidae, (,'haropidae. and Otoconchidae,
lint another traditionalK' '"endodontoitl" (aniilw Discidae,
fell into a distinct clade of its own. Endodontidae is ten-
tatixelv placed on the cladogram in the position of the
"endodontoid clade" ol Wade et al.

Waile et al. (2001) included no Orthalicidae: their sole
member of the dixerse "bulinniloid' clade was a New
Zealand species of Phicostiihis. Orthaiicidae. exemplified
b\- Liiiims fa.sciatus. is tentatixeK' placed on the clado-
gram in the position of Watle et al.'s Bulimuiidae.

The correctness of the tentative placements herein of
Endodontidae, Orthalieitlae and Sagchdae could be test-
ed In additional molecular stuilies.

Characters: Table 2 shows the distribution of char-
acter states through the taxa anal\y.ed herein, based on
the obsenations of SuMirox' l200(J). The characters and
their alternative states are as follows; all are described
in more detail and illustratetl b\- Su\oro\ i2()00: 90-99):

1. Breathing channel: (.\) with pectinate folds forming
a \aKe: (B) without \al\e.

2. Rectal termination: (.\) open ("unclosed"), witliout
acklitional structm"es: (B) open, with lobe; (C) open, with
papilla: (D' open, with one rectal pilaster; (E) open, with
two rectal pilasters; (F) partialK closed; i(;i full\- closed.

■ >. Seccnidarx uretiu": (A) open ( '"uncloseir"); (B)
closed.

4. Pneumostomal exeri'toiA routes; (A) sim[ili'; (\i) Y-
shaped with Ll-shaped pilaster; (C) Y-shaped, closed; (D)
r-slia[ied with anal pihister; (E) Y-shaped with anal pi-
laster.

Polarit\ ol tnuislonnalion series was inferred In iter-
;Ui\e out-gi'oup comparison of secjuentialK more iuclu-
si\i' clades and application ol the "Reiatixc .\pomoiphv
iiule": Homologous characters lound in the members of
a monophvlelie gidiip and in its sister gl'oiip ;ii"e ple-
siomorphic. while homologous charactei's lound ouK' in
the in-group :ii'e ;ipom(irpliic (Brooks and McLennan,
1991. (■specialK pp. I(i (i2).

-The \alid tainiK -group n;nne lor llie New World "(.'aniaeni-
dae" is Pleurodontidae \on Ihenn^. 1912; ;i suprafaniilial group
containing Pleurodontidae and Saiidid.ie l'ilslir\. 1S9.'5. wxiiild
lie eoirectK kuowu as Sawloidea.

B. Roth. 2001

Pasje 143

Table 2. Distribution ordiaractcr slates aiiiiiii<r |a\a auaK/ril
luMciiL Family codes as in Talile 1. See te\t lor dediiitioii ol
eliaracter state' sxiiihols lA-CJl.

Piieuiiio-

lleetal

stoiual

Breatliini;

tenniiia-

SeeomlaA

exireton.

FaiiiiK

clianiiel

tioii

ureter

routt-

sue

B

G

B

A

PAR

B

E

A

D

CLA

B

E

A

D

SUB

B

D

B

A

ACH

B

G

B

E

ORT

B

F

B

1)

ACA

B

G

A

E

SAG

A

(;

B

c

CER

B

E

A

B

COG

B

E

A

1)

ORG

B

E

A

D

CHO

B

E

A

D

PYR

B

E

A

D

EM

B

E

A

D

BRA

B

G

B

B

HYG

B

G

A

B

HEL

B

c:

A

B

END

B

.\

\

\

DIS

B

B

B

B

POL

B

c;

B

B

DAU

B

D

B

A

ARP

B

D

B

A

HAP

B

D

B

B

LIM

B

D

B

A

ARl

B

I)

B

B

RESULTS

In Fiu;nre 2 tlic eliaraetci- states (il tlit- hreatlilii'j; eliaiiiiel
are mapped on tlie elado^raiii Ironi Fi(j;iiri' 1. The ar-
rangement is simple and re(jnires onl\ one tiansloiina-
tion, from the alisenee to the presence <if a \al\e. in
Sagdidae.

In Figure 3 the character slates ol the rectal termi-
nation are mapped on tlie same chulogram. The arrange-
ment is complex. The pattern is more intelligihie if the
character i.s fir.st parsed into a three-state charai'ter with
rectal termination either (.\) open. (Yi paitialK closed.
or (Z) full\- closed (Figure 4). Out-gronp compaiison es-
tablishes an open termination as plesiomorphic for St\-
lommatophora. witli three homoplastic instances of clo-
sure: once in the clade comprising iiehcidae. Pol\g\ri-
dae, Bradvbaenidae. Ihgromiidae. and Sagdidae. and
t\\ice in the clatle comprising Succineidae and Orthali-
cidae. In the latter case it is not clear that partial closure
(Orthalicidae) preceded full closm-e (SuccineidaiO or
Nice \ersa; at the present lexel of resolution, liotli stales
appear to deri\e directK Irom the open con<lition.

The presence of some internal structure [one oi' more
pilasters, a lobe, or a papilla) is plesiomoiphic: an open
termination \\ithout additional stnictures occurs onK as
an autapomoiphic state in Endodontidae. The presence

ol .1 p.ipilla is homoplastic, occurring in .\ca\idai' and
\cliatiiiid.ie, .\ single rectal pilaster is homoplastic, oc-
curring in Ihe clade coiisisliii'4 of Limacoidea .scd.sv/ lato
and Arionidae and in I laplolrematitlae. Presence of two
rectal pilastt'is is homoplastic, occurring in all ta\a of the
orthinethian clade. in (,'lausiliidae. and in Ceriidae. .A
rectal lobe is an aiitapomorphx of I^iscidae.

in Figure ■") the character slates of the secondan ure-
ter aic mapped. Out-group comparison eslai)lishes a
ilosed ureter as plesiomoiphic lor St\lomiiiatophora.
with lour Iraiisformations to an ojieii iirc'ler ami one re-
\ersal back to a closed ureter In the clade consisting of
Limacoidea vciis7/ Into and Arionidae.

in Figure (i the character states of the pueiiiiiostomal
excreton route are mapped. Out-group I'omparison es-
tablishes a simple route icharacter state .\i as tlie prob-
able plesiomoii)hic condition, from which the four more
complex states \] through K arose. Character .states B,
D. and E show homoplasy: character state (; arises aut-
apomoiphically in Sagdidae from character state B.
There are no rexersals from complex to simple.

niscrssiox

It is not here assumed that a molecular pli\l()gen\' is ipso
facto more triithlul than aii\ other, but the|-e are several
reasons \\li\ ihe pli\ logeiiellc h\polliesis of Wade et al.
(2001 I is a useliil slanclard lor com])arisons of tlie hpe
picseuted liert-. it rejiresents the most coinprehensixe
molecular anahsis ol St\loiiiiiiatopliora to date, includ-
ing almost 100 genera and S4.3 iinaiiibignousl\- aligned
nucleotide sites. Manx of its clades lia\e strong bootstrap
support. .Although not iiecessariK determinati\e. man\-
ol its conclusions ha\i' intnitixc a])peal. sue!) as the ex-
tent to which traditionalK recogni/ed taxonomic grou])s
plot ,is monopln letic. and the geographicalK coherent
di\ision ol olhers. One limitation, as pointed out b\ the
aiilhors lliemseKcs. is that main ol the branches iiii-
medialcK aboxe the ■'achatinoid"-"iioii-ai-lialiiioid" di-
cliolom\ are rallier weakK supported. This i-ould reflect
either a limitation in the resoKing power ol the aiiaK~/ed
se(|iieiice or a phase ol explosixc clado^enesis or extinc-
tion in the liiston ol Stxlommatophora.

Inlerring exoliitionan histon b\ mapping character
states on this liranching iliagram requires no ii priori
assumplioiis other tlian that cNolntion has occurred.
(The bel.i(i\e \pomoi"ph\ l^ile is nol an assumption
about the course ol e\()lution. but a method to eiilorce
parsimoin. I iiecause the imderKiiig pli\logenetic h\-
polhesis is based on a \(M-\ tlilTerent (lata set. the pos-
sibiiilN ol circular reasoning is |iracticall\ eliminated
iHotli. I99(ii.

In contrast, the schema ol e\()liilionan transtorma-
lions ol piieiiinostomal inoiphologv proposed b\ Su\'o-
rov (2(M)0: fig. 10) is dri\en b\- so-called limctioiial an;i]-
\sis rather than h\ out-group comparison (W'alrous and
Wheeler. 19S1; Brooks and McLennan. 1991). It is
based strongK on a priori assiimjitions about tlie course
ol e\()lutioii (■moipho-plnsiological progress. " "increiis-

Past' 144

THE NAUTILUS. \ol. 115. No. 4

^

-C

-c

ENI

ORC-1

PYR

ORC-2

COC

PAR

CHO

LIM

ARP

OAU

ARI

CLA

END

AC A

HEL

POL

BRA

HYG

SAG

HAP

CER

DIS

sue
ORr

ACH
SUB

rl-

r

-L_.

ENI

A

ORC-1

A

PYR

A

ORC.2

A

COC

A

PAR

A

CHO

A

LIM

B

ARP

B

OAU

B

ARI

B

CLA

A

END

A

ACA

A

HEL

A

POL

B

BRA

B

HYG

A

SAG

B

HAP

B

CER

A

DIS

B

sue

B

ORT

B

ACH

a

SUB

B

7

ENI

BEAD

"pupilloid'

p— ORC-1

BEAD

"pupiUoid"

' PYR

BEAD

"pupilloid"

"pupilloid"

BEAD
BEAD
BEAD

CHO

"pupiDoid"

1 LIM

BDBA

"zonilOid"

1 1 ARP

BDBA

"zonitoid"

1 DAU

ARI

BDBA
BDBB

"zonitotd"

'haplotrematoid"

1 CLA

BEAD
BAAA

"pupilloid-

1 END

"endodontoid"

BCAE
BGAB

1 HEL

"helicoid"

1 POL

BGBB
BGBB

"bradybaenoid"

1 BRA

"bradybaenoid*

1 1 SAG

AGCB

"sagdoid"

1 HAP

BDBB

"haplotrematoid"

1 CER

SEAB

"cenoid"

1 ■ DIS

BBBB

"discoid"

1 — sue

BGBA
BFBD

"succinoid"

1 ORT

"orthalicoid"

, ACH

BCBE
BDBA

"achatinoid"

1 SUB

"zonitoid"

Figures 2-7. Character-state distril)iili>>iis, 2. Distriliiitioii ul lucilliiii'j; clianiiel willi \al\c (A) and witlioiit valve (B) and location
of traiislbrniation Irom B to \. mapped on cladoiirain Ironi Kiuiiiv I, -i. 1 )islnl)ntion of rectal termination states: (.\) opim ("nn-
closed"). without additional structures; ( IV* open, with lobe: (Ci open, with piipilla; (D) open, uilli oiu' leelal |iilasler: (K) open,
widi two rectal pilasters; (F) partiallv closed; iC;) hillv closed: ;md loiMtion of li^mslnrmations where lhe\ lan he inlirred from out-
j^roup comparison. 4. Distrilmlion' of open {\1, partialK closed A', and hill\ closed iZl reet.il lcriiiiii:ilioii and loc'ation of

B. Rotii, 2001

ViWf 145

int; lc\cl nl (lill<'iciitiati(in. ' iiili'iisilicalldii iil riiiic-
tions"), mostly articulated hi llic (irsi li.iHHr iIh' Twen-
tieth Ontiin- (SiivoroN. 2000: 901.

Tlif aiialxsis liciciii i|-"i^iiic 2) iiidifalcs llial a liii-alli-
ing clianncl without a \al\c is pIcsidiiKiiphu ami the
presfiK'i' (il a \al\c is ajioiniirpliit'. 'i'liis is (■(insislciil w illi
the tninslonnation scries inlcrrcd 1)\ Sinnrox i2000: 00).
The presence ol a \aKc is aiitapoiiiorpliic in and diag-
nostic of Sagtlitlac. The alisciicc of a \aKc is mil aiihi-
poniorphic for an\ iiioiiopli\ Iclic group.

This analysis (Figure 4' imiicatt's thai an epen ledal
termination is plcsiomoiphic and partial or full closure
is apomoipliic'. This is consistent with the ti-ansroniiatioii
troin open to closed iiilened In .Sii\(iro\. ,Sii\oro\ re-
ferred to the condition of an open terniinalion with \ar-
ioiis aecesson strnctm-es as intermediate hetwceii the
simple open anil closed stales i2000: 00! Iliiwe\ei. this
analysis (Fignre 3) shows no such transtoimation series
within St\lommatopliora: the oiiK nislanee of a simple
open state is that ol Fmlodontidae. which ont-groiip
comparison establishes as a translorinalion Irom a state
ha\ing two rectal pilasters.

There are hvo hoinojilastic instanci's of closure of the
rectal termination: in the elade comprising Helicidae.
Pol\g\ritlae. BraiKliaenitlae, I l\gromiidac. and Sagdidae
(essentialK', the claile Tlelicoidea sciisii Itilo" of Both
and Sadeghian. in pri'ss). and in the ilade comprising
Succineidae and Ortiialiciilae (essentialK. (.'lade iV of
Roth and Sadeghian. in pressi. ('Insure is dia'^nostic' of
each of these clades. iTiere are mi reversals Inmi :i
closed to an open stati'. .\ii open lect.ii teiinination is
not antajioinoqihic tor an\ inoiioplnletic i^roiip.

This aiiaksis (Figure 5) indicates that a closed scc-
omlan ureter is plesiomorphie for St\ lomuiatoplioia.
with tour lr:iiisloiiiiatiiiiis In an open iilelei :iiid iiiie re-
\ersal hack to a cliisei.1 ureter in the elade consisting ol
Limacoidea s()/,v(/ lafa of liaiisdorf ilOOSi plus .\rioui-
dae. This is contrar\ to the direction ol liansformalinii
argued by Snvonn (2000: 99).

This analysis (Figure 6) indicates thai a simple e\ere-
toiA route is plesiomoiphic' and that the tr:iiisloriii:itioii
to all othi'r apiiimiiphic states excejil ( ,' (Y-slKipid,
closed,) occurred homoplasticalK, \ll ol the complex
states are deii\ed as direct translorinations Ikhii the
simple state, except state (,'. which arises liniii slate H
(Y-shaped with U-shapt'd pilasteri. \'o reversals Irom a
compk'x to a simple state occur (,'haraeler slate I) i| -
shaped with anal pilaster) is uniform throughout the Oi-
thurethra. It also occurs in Clansiliidae and ( )rthalicidae.
but Su\()ro\ (2000: 92) cast doiilil on the liiinuilogv ol
the condition in ( ilaiisiliidae and that m Oiilimclhraii
taxa. Su\-oro\'s (2000: hg. 10) sc'enario ol evoliilion el

the piieiiiiioslomal area includes the lianslormations A
lo H lour times). .-\ to I). B to ( :. and I) to K. All of
these translormalions except I) to F, are found in the
[ireseiil .inaKsis. The uiimlier of transformations (S> is
onK slightly greater than that (7) suggested In Suvorov'.
Much ol Suvonn's aiiaKsis is based on what he called
■patterns." naiiiek. f) observed combinations of the
above chaiaiter states, lie named the patterns for taxa
showing tlieiii (e.g., "endodontoid." "haiilotrematoid").
but this practice is not vi'i'v hel])lul when rather reiiiotel\-
related taxa show the same pattern (e.g.. Haplolremati-
dae and \iioiiidae . His I'ignre 10 sli(ms a sciiema of

probable' ev olutionaiv relationships among the pat-
terns. A plot ol the patterns on the cladogram (Figure
. ' gives little sii|)])ort lor the schema and suggests in-
stead that, lor most ol stvlommatophoran histon. the
lour ehaiacteis ol llie piieiimostomal area have not
evolvetl in close eoneeii. Bather than being basal as in
.Suvoidvs scenario, the 'Cndodonloiil" pattern ( B.A.-\.\,
Inr characters 1 through I' is hiiiliK derived. The "pnp-
illoid" pattein (i5lv\l)i is uniform through Orthurelhra.

\ similar but probablv not honiologous pattern occurs
in ( )l;insiliidai'. 4"lie ' /onitoid " |)alt('rn (BDBA) is nni-
loiiii in l.imaeoidea s(ii.\ii lulu and, witli one transfor-
iii.ition ito i5l)l5B> in its sister group, .\rionidae: it also
occurs in the phvlogeneticallv remote Snbulinidac. The
clade llelieoidea \cii\ii liilo exelnsivi' of Sagdidae is di-
vided among the "helicoid " i iU ;.\B) and "bradxbaeuoid"
(JUiBJ-ii conhgurations. Other |)atterus are either aiita-

p< pliic 111 lioiiio[)lastic. and il is clear that tluir der-

ivalion iiiiisl involve separate traiislonnations of the
eoiiipiiiieiil eharacteis. ['or this reason. 1 believi' that the
])livlogenv ol till' [inenmostomal area is better visnali/ed
in terms ol individual characters, as done in the present
sliidv, than in lerms ol composite 'patterns'. For future
stiidv. the pneiuiioslonial area eharacteis examined b\
{■.mbeiton il99Fl max jirovide additional data, il thev
laii be coordinated with those obsei'ved bv Suvorov.

A( K\()\\I4'.I)(;MF\TS

I'etei Mordaii :iiid David i.iiidberg real! ilrafts ol till'
mamiseiipl and ollered valuable eoiiimeiits.

141 i:i;vi'ii;f cifKi)

liidoks I) i; iikI I) A. .McLennan. 1991. Plivk)gcnv. Eiologx',
.111(1 Beliaviiii: a Kcsi'arcli I'rognuii in Comparative Biol-
ii'j;v, I iiivcisilv ol OhiiMiio Press: (.'kic-ago. 434 pp.

I'.iiiheitiiii, K, (). 1991. Pokgvrid relations: a plivlogeiieticaiial-
vsis III 17 siililainilii's ol land snails i Molliisca: (Jaslropoila:
Slvloiiiiiialopkoral. '/oolc laical joiinuil nl the l.iiineaii So-
cietv l().'5: 2(17-224.

transforiiiations. 5. Distrihiitioii of open (A) and closed (IVi sccoiidan- ureter and location of transroniialioiis. 6. Distiihiitioii of
excreton- routes: (A) simple: IB) \'-sliape(l with ll-slia])e(l pilaster: ((,'' '^'-sliapc'd. closed: (Di 1" -shaped vvilli anal pilaster: .K \-
shaped with anal pilaster: and lucatioii ol' tiaiistuiiiiatieiis 7. Dislnlnilioii nl ciniihiiialioiis ol chaiacter states lor ckaractcrs 1—4
("patterns" of Suvorov. 2()()l)i.

Pase 146

THE NAUTILUS. \'ol. 115. \o. 4

Enibt'rton. K. (,'., C^. S. kuutiu. <:, \1. I)a\i.s. .S. .\1. I'liilllps.
K. M. Moiulert'wic'z. and ^ II (iiio. 1990. (.■omparisou
of rcc'i'iit fla.s.sifications ol stNlDiiiiiiatoplioran laiid-siiail
families, and csalnation ol iargc-rihosoniai liNA sequciic-
iniT for their iiliNlosicnetics. Malatolosiia 31: 327-352.

GoodfritMni. (;. .\. U)S(i. Radiation ol the land snail genn.s S(i<!,-
(hi (Pnhnonata: Sagdidat'): comparative morj5holosi\', iiio-
geograpliN' and ocoiog\' of the species in north-central Ja-
nuiica. Zoolo'^ical |onnial ol the l.innean .Society ST: .367-
398.

llansdorf B. H. 199S. Ph\logen\ of the Limacoiilea .sci/.s/f lain
iGastropoda: .St\lormnatoplioral. |onnial ol Mollnscan
Stndies 64: 35-66.

Roth, B. 1996. Homoplastic loss of dart apparatus, plixlogenv
of the genera, and a piivlogenetic tavonomy of the Hel-
mintliogivptidae (Gastropoda: I'nimonata). The Wliger
39: 18-42.

Roth. H. anil 1'. Saileghian. In press, (.liecklist ol the land snails
and slugs of Gahfomia. Santa Barbara Musmnii of Natural
lliston' Omtrihntions in Science.

Su\oro\, .\. N. 2000. F"nnctional niorph()log\ ol jineuniostomal
area in terrestrial I'uhiionata ((iastropoda). Rutlienica 10:
89-104.

Wade. C. M. and P. B. Mordan. 2000. E\-oliition within the
gastropod molluscs: using the rihosomal RNA gene cluster
as an indicator of plnlogenetic relationships. Journal of
Mollnscan Studies 66: 56.5-570.

Wade, C. M., P. B. .Mordan, and B. Clarke. 2001. .\ ph\logeny
of the land snails (C.astropoda: Pulmonata!. Proceedings
of the Roxal Society of London. iB) 268: 41.3-122.

W'atrons, L. Iv and Q. D. Wheeler. 1981. The out-group com-
parison method of character anak'sis. S\stematic Zoolog\'
30: 1-1 1.

THE NAUTILUS 115(4):147-149, 2001

Page 147

Cypmccassis chipohuuL a new species ((;astr()p()(la: Cassidae)
from the Miocene Chipola Formation of nortliwx^stcM-n Florida

Richard Duerr

Post Office Box 105.5
Okeechobee. FL .3497:3 US.\
pdiegel@,strato.iiet

AiiS'lKACT

\ new fossil gastropod of the faiiiiK Cassidae, Cijpmccassis
chipolana, is desciihed from the lower Miocene Chipola For-
mation of northwestern Florida. Ci/pmccas.sis chipohinii ap-
pears to be the only known Miocene Ci/pnurassis seiisii sfrkio
from the Americas. The new species most closely resembles
the Recent Cijprticrassis uiliiuif Kreipl and Alf, 2()()(), a species
endemic to tropical west America.

Adilitiniitil kill uonis: Tertian. Neogene. fossil.

LNTRODUCTION

Tlie tamilv Cas.sidae lias its earlie.st known record in tiie
lower Eocene. Dnring the Miocene tiie faniil\ liad be-
come established tliroughoiit the warm and temperate
seas of the world (Abbott. lyOS). Wliile memlx-rs of die
Cassidae are not nnconnnon in botii the Recent antl the
fossil record. oni\ one species of tiie genus most closelv
related to Cypraccas.sis. Cassis delta Parker, 1948, has
previously been identified from the lower Miocene Chi-
pola Formation of northwestern Florida. Complete or
iragmentan^ specimens oi' Cassis delta are tbund in main
portions of the (^Iiipola F^ormatiim botli at Temnile
Creek (the t\pe localit\ ) and along the Cliipola Rixcr.
The new species is known tiom a single specimen, col-
lected on the Chipola River.

The Miocene (Jhipola Fori nation correlates in age
with the Rurdigalian of Europe (\okes. 196.5). Three \al-
itl species of the genus Ci/pracca^sis seiisii stricto \\d\v
been described from the .Mioceiu' of Europe: C ci/-
pracifoniiis (Borson. 1820); C. suhcniinciia (d'()rl)ign\.
1852); C. snhtcstictdtis (d'Orbigm. 1852); ;in(l one Ironi
the Miocene of East Africa, C piisliilala iC.n\. 19271
(.Abbott, 1968).

Three Recent Ci/praccassis \ciisii stricto. C, tenuis
(Wood, 1928) and C. uiliiKir Irom the eastern Pacific
Ocean, and C. tcsticulus (Linnaeus. 1758) from (he .At-
lantic Ocean, are known to cxisl in tlie Wi'stern Hemi-
sphere. Ci/praccassis tcsticidiis also occurs in the Pleis-
tocene of Barbatlos and Cuba (Weisbord, 1962) and in
the Miocene of the Dominican Republic ((labb, 187.5;
Pilsbrv', 1922). |nng (1971) reported three incomplete

specimens oi an niideseril)e(l C^i/jjraccassis Iroui tlie
Miocene Crand Bay Formation of' Carriacon, West In-
dies. 'Hjis latter ta\on is morphologicailv similar to aiul
ma\ lie tiie preiiecessor oi Ci/pracrassis trsliniliis se-
iicfS.alica ((inielin, 1791) from West .MVica. Ci/pracrassis
nifa (Linnaeus, 17.58) occurs in tlie indo-i'acific Region
irom East .Africa to eastern Poixuesia lAlibolt, 1968).

Tlie hoiotvpe oi Ct/praccassis rhijutlaiui is deposited
in the collection oi tlie Department of Paleobioiogx. Xa-
(ional .Museum of Natural Ilistoa' iUS.Wl), Smithson-
ian institiitiiin. Washington. !)( '. US.-\.

SYSTEMATIC S

1797

Class Gastropoda Cjivier.
SuperiamiK' Tonnacea Peiie. 1926
FamiK Cassidai' Swainson. i8.'52
(lenus Ci/jiraccassis Stntclibun. 18.37
Snligeiins Cijpraccassis Stutchl)nn, 18:57

Cijj)raccassis iCiipraccassis) chipuhnui new species
I Figure 1 1

Description: Sliell thick, biconiiai. length diolotvpc)
57.4 mm, widtli :)6.2 mm. Whorls about 7 inciucfing
aliout :5 isomi'wiiat damaged' protoconcli whorls. Spire
sculpture ol excnk spaced rounded pustules on inner
edge oi suture and a raised cord witli narrow, elongate
knobs. Sntnrc sliglitK impn-ssed. Dorsal sculpture con-
sisting ol 1 rows ol spiral bands oi 10-1 I. iairlv rounded
nocinlrs. I;ngest al slionlder and decreasing in size an-
teiioiK. disappearing entirek on anterior tiiird ol dor-
sum. i5ands ol nodules inti-rspersed witli one row oi
greatk reduced nodules. a[)pro\imalek :>6 on posterior
row. decreasing in number and iiecoming :> rows ol elon-
gate riiis coxering toreniost third oi dorsum. Parietal
sliield broaii. thick, elevated posteriork at commissure
witli labrnm; |)osterior canal a narrow, shallow depres-
sion. Apertnrai |ioi1ion of jiarielal wall containing aliout
:50 narrow lirations oi \ar\ing lengtlis, crossing an itxiai
swelling or ridge I Figure 1) on columellar wall and ex-
tending into a|)ertnre. .\nterior third oi parietal wall and
shield slightk raised, with enlarged lirations. .\nterior

Page 14S

THE NAUTILUS, X'ol. 115. No, 4

I'imii'i's 1-.'}. Spccicsiit ( .(/yi/v/ivv/vsvs in apcrtiiial. latrral, and aliapcrluial \ic'\\s, 1. Cijjx'dccdssis thijiohnui. new spcrics, liolohpc.
USNM 51TS92, Icimtli 57.4 mm, uidlli :>(i.2 iimi. Irom Cliipola lii\cr. .>(f2S.l:).o' N, .S5°()9.558' \\ . 2. CiipnuTOssis uihmic Kreipl
and .Air. 2()()(). Kmilio Carcia (.'ollci-tioii IS.^fi'V Imi^lli ."i.l.T mm. width 24. (i mm, IrDm Islas Sccas. (ailld dc (:liiri(|ni. dri'dijcd,
120-240 111. ,saiKl/,sliC'Il bottom. 3. Q'.ijprm'Cd.ssis nija ! I .imiaciis. 175S1, lciit;th S2..5 mm. width " .1 iiiiii. hido I'aiifif Kcnion, Ph\lli,s
Dicsjcl (>)!lcctioTi. hir i-omparisoii with ('i/j>riii'<iissis rliiixiliiiin

R. DueiT. 2001

Puirc 149

siplioiial I'aiial opi'iiiiiij; on iiij;lil side (il slicll (in dorsal
\ie\v). rellexetl, deep, with eliipped edt;e. 'IVnc and lalse
umbilicus (see Kreipl. 1997: 9) present; true unihilicns
open and deep, false uniliiliiiis elosed. Onlci lip tliick.
extending slightK into apeitnic and ictni-xed npward
o\ei" dorsum. Iimer poition ol lalirnm witli ap[)ro\i-
mateK 17 single or paiicd stont lirations.

Type locality: Chipola l^ivei'. 30°8.135' N. S5°09.55S'
\V (= Tnlane Universitx' loealitv TU 950. Chipola For-
mation. Chipola Ri\er. west hank about (iOO in above
Farlex C;reek (SW 1/4 See. 20. TIN, U9\\). t:alhoun
Countx. Florida).

Type material: lIolol\pe. HSX.M ( I'aleobiologx )
517892. length 57.4 nnii, width 3fS.2 nnn.

EbmiologA': Named for the Chipola River, on which
the t\~pv loealitN is situated.

Discussion: The Miocene European (■i/j)niccassis
may be easil\- separated from C. chipohiiid l)\ the pres-
ence in the former of rib like longitudinal plications on
the dorsal surfaces of the shells or, in one species, !)\ a
smooth dorsum. The Miocene C. jiiistiilata of Fast .\f-
rica, as the name implies, has a ilorsal sculpture of rmich
larger pustules, arranged in a different pattern than
those of C. chipolana. Ci/praccassis rliipohina is sepa-
rated from the Recent eastern Pacific C /ciniis hv the
larger, thinner shell, diaplianous parietal shield, and
deeper posterior canal of C, Iruiiis The .Miocene (,', chi-
polana ma\' be easih' distinguished from the .Miocene to
Recent C. testicnliis tc.'^ticiilus of tlie Caribbean Region
b\' the reticulate sculptiu'e on the dorsum, more round-
ed shoulder, and thinner parietal shield of the lattiT The
undescribed Miocene C!i))iyicc(issis from Carriacon
(Jung, 1971), which somewhat resembles the Recent C.
tcsticnlus seiic^alica. is dillerentiated from the new spe-
cies h\ the prominent orthocline axial plicae and lack ol
nodules on the undescribed species. The Recent C;/-
praecassiH ntfa (Figure 3), ranging from East Africa to
eastern PoKnesia, resembles C. chipohiua. but differs
from it In- the much larger shell and more ronudetl pa-
rietal shield of C. nifa. Ci/jinici-iissis nifii also lacks the
raised portion, or swelling, on the anti'rior third of the
parietal wall and shield of C. cliipohnui

The species most similar to C. chipolaiui is the liecent
C. wiliitac (F'igure 2). but that species has one more
band of spiral knobs on the bod\ whorl: has S or 9 knobs
per band as opposed to 10 oi' 1 1 on C. cliipolaiia: has a
more rounded parietal shield: and lacks the axial swelling
or ridge on the columellar wall of C. cliipuliiiui. On C.
wilmac. the bands of large knobs are separated In two
rows of smaller knobs or beads compared to one low on
C. chipolana.

Features tliat are common to ('. chipolana and ('. nil-
mac include similar oxerall shapes, small ailult si/es. :ind
the unusual diagonal separation on the anterior third of

ihc colnmellai callns, ( )ii the basis of moqihological sim-
ilarities, the possibility exists (hat C, chipolana \\\d\ be
the ancestor ol the Recent C. uilniac. whiih became
isolated in the tro|)ic'al eastern Pacific with closing of the
Islhniiis ol Panama (hiring the middle I'liocene iCroniii
el af. f9S4: 43),

ACK\()\\M:nc\ii:\Ts

\l\ deepest appreciation to jose II. Leal wlio critiijiied
the original manuscrijit. proxided incentive, and pre-
paix-d the digital images ami plate. .Mv thanks and ap-
preciation also go to: Richard Petit lor inloriiiiiig me that
Abbott's Indo-Pacific Mollnsca contained inlonnation on
lossil European (^assidae: Jean (Claude .Six lor informa-
tion on lossil I'jiropean Ci/praccassis: C.nw Rosenberg
and Mark kilsoii lor relerence material: Warren Rlow
and William C, JAons lor rev iewing the maimscripl: two
anoiivmoiis reviewers tor suggestions and pertini'iit in-
lonnation: Burke and Brooks Haves for permission to
collect on their projiertv: Kniilio Ciarcia for loan of spec-
imens ol C'|//)/v/((v/vv/v nilinac: Pamela Dii'gel for plac-
ing the manuscript on a computer file: and Phvllis Die-
gel lor relevant relerence mateiial and inspiration.

1JTER.\TURF C:iTED

.\hhott. R. T 19(iS. The Helmet Shells of the WVidd (Cassidae)
Part 1. Indo-Pacific .\lolliis.a 2i9': 7-201.

Ck.iiiii.T M.. L. BvIk-11. H. Pooiv. B. Blackvvvklen J, I.idiiicoal
and |. Hazel 19S4. .Age and correlation ol eiiicrt;e(l Plio-
cene and Pleistocene deposits, U,S. .Mlaiitic Coastal Plain.
Palaeogeoi^rapliv. Palaeo< liiiiatolot^v. I'alacoecoiogx 47:
21-51.

C;ilih, W. E. 1S7.>. On the toponraphv and <ieolo<.^ of Santo
Doininsjo. Transactions ol the .\iiicrican I'hilosophical So-
cietx. new series. 1.5: 49-259. 2 ina])s.

|nii'4. P, 1971, i-'ossil Molliisks Ironi ('arriacoii. West Inilies.
Bulletins otAiii.-rieaii P;ik'oii((ilo'j>. 61(269): 147-262. pis.
1-21.

kreipl K 1997, Becent Cassidae. Wriaii Clirista HeiiMiien.
\\ lesh;lllell 151 pp.

Kiiipl K and \ All. 2009. \ new species ol (..tjprai'cassis
Stiitclihiin. 1S.37 i .Mollu.si-a: (iastropodal frnin Pacific
P:iiiaina. Pa Coiicliiglia .32(297); 43-15.

I';irker |. I). 194S. .\ new Cassis and other iiiolliisks Ironi the
( ^liipoki iMniiiation. The Nautilus 61 : 90-95. pi. 6. fi<;s. 1 -
21).

I'llshi-v. II. .\. 1922. A revision ol W. \\. t.ahh's Tertiai-v .Mol-
lusc;! ol S.iiito Doiniiigo. Proceedings of the .Acadeiiiv ol
N.iliir.il Sciences of Fliikulelpliia 73(192P: 305-1.35. pis.
16 47,

\nkes. P. 11, I9(i5. Notes on the ;ige ol the Chipoki I'"onn;ition
I Miocene' ol .Nortlivvesteni Florid;!. Tnkiiie Studies in Ce-
ologv 3; 20.5-20S,

Weishord. N. I'7 1962. l,;ile Ceuo/oic ( iastroiiods Ironi North-
ern \cne/.n(4;i. Bulletins ol .\iueric;ui P;ileoutolo<;v
12(193): 1-672, pis. 1-lS.

THE NAUTILUS 115(41:150. 2001

Faize 150

Note

Correction of a mistake introduced in tlie description of an
aeolid nndibranch of tlie genns Favohmis Gray, 1850

Francisco J. Garcia

Di-pai'tamciito tk- Fisi()iciy;i'a v IJiologi'a

Animal, Facultad de BioJogi'a
Universidad de .Se\i]la
Avenida Reiiia Mercedes. 6
41(1S() Sc\illa
.SP.MX
fjgarcia@cica.es

Jesus S. Troncoso

Area de Biolcigia Animal l'a< ullad de

Cieiicia.s del Mai
Uni\ersidad de N'igo
Lagoa.s-Marcosende. N'igo
.SPAIN
tioTicosofeuvii'o.es

RecentK', a new species ot Favorinus V,n\\\ 1850, troiii
tlie eastern Pacific Ocean was described bv onrselv es in
tills jonrnal (Garcia and Troncoso, 2001). The iiaiiie ul
this species was designated clciuilrxiac as "a juxtaposition
ot the names oi the vonnm'st daniiiitiTs" of the authors
Garcia and Troncoso, tliat is, Kli-iia ant! Ale.xia, respec-
tively. Ne\ertheless, a t\poii;raphic error related to the
name of the species was produced in this paper. Al-
thougli the species was cited as Fmorintis clciuik'xiae on
22 occasions in the paper, tlic name also appeared twice
(on pages 55 and 57) as h'iin>iiiins fli'iuilcixir

According to the International (>ode of Zoological No-
menclature' (ICZ\. 1999). articles 32.2.1 and 24.2.3
should lie applii'd in this sitnalion. 'i'licse articles refer
to tlie correct original spelling of a scientific name. \r-
ticle 32.2.1 states: "If a name is spelled in more tlian
one way in the work in which il was established, tlien,
except as pro\ided otlici-\\isc in (his Article, the correct
original .spelling is thai chdsrn li\ ihc i-'irst Kexiser" and
Article 24.2.3 states: "Selection ol coiiiTt oiiginal spell-
ing. If a name is spelled in mnrc than one wa\ in tin'
original work, the liist anllicn to lia\c' cited tlicni to-
gether and to have selected one spelling as cori'et'l is the
First He\iser. The selected spelling (if not incorrect un-
der Articles 32.4. or .32.5) is therein fixed as the correct
original spelling; an\- other s|)elling is incorrect (and
therefore unavailable)." .\ct-ording to these .Articles, act-
ing as First Rexisers we consider elenalexiae as the cor-
rect original spelling. The name represents a nonn in
apposition and so is indeclinable.

Since our description was published, Hans Bertseh
emended the name of the species on Michael I). Miller's
site on the World Wide Web. Bert.sch held the \iew that
the termination -tic of the name was ineoncet and that
it should 1k' corrected to -iiniiii i hence clfiuilcxidiiini)
because -armii is the projier Latin suffix for a patronxin
established in honor of more than one female iierson.
However, we belie\e tliat the spelling cIciKilcxiiiniin \\
incorrect because the plural form should oiiK be applied
when a specific name is formed Ironi a personal name

that icirresponds to hvo or more people who share iden-
tical names. For example, the aeolid Flalicllina luarcii-
\oniin was named bv GosliiUM" and Kuzirian il990t in
honor of Ernst Marcus and his wile E\('line du Bois-
Ke\nioiul Marcus, who first recorded this species. Ac-
cordiugK, a patromin di'dicated to mori' than one per-
son with nnn-icU'iitical nami's. with the specific name
loiined li\ juxta[)osition of the names ol each person,
sliiinld lie considered in the singular form (i.e., Plii/llidin
rtirUonhojji. as proposed b\' Brnnckhorst (1993), is the
I'orrect name in liniior of Mr. Gla\' Garlson and Dr Pattx-
jo Ilofl'l. Besides this. Article 9.8 of the Gode stipulates
that material distributed b\" electronic signals (such as
on the World \\ ide Web) is, for pni"|ioses ol loinial no-
nienelatnre. unpublished llf^ZN, 1999). Thus, the
eniended n.une ol 15eii:sch cannot be eonsideri'd as a
s\ noiuin ol cIciKilcxiai' because it has not been included
in a imbiished work.

.\(:K\(mLEIX;MENTS

We wisli Id expi'ess our gratitude to Richard (.'. Willan,
|os(' II. I.imI flans Bertsi-h, Miguel .\. Zara/aga. and
|os('' Ti'uiplado ln|- their eonniieiils during the |irepara-
lion 111 this note.

LITF.R.VrUHE CITED

Bnuickluirst. ]). J. 1993. The swstematics and pli\l(igeii\ (if
I'livllidiid N'udiliraiielis (l^iiridoidea). Heeerds (il the .\iis-
lialiaii Miisi'imi. Supplement Ifi: |-|()7.

Oaieia, I', \ and j. S. Triiiico.sd. 20(11. Fdvoriiiiis clcniilexidc. a
new species (()pistli<il)i'aneliia: Aeolidiidae) from tlie east-
irii Pacific Ocean. Tlie Nautilus 115: 55-fil.

( iiisliiier. T. M. ami .\. M. Kuzirian. 1990. Two new species (il
l''laliellinidac i Ojiistlieiiraneliia: .Aeolidaea) Inim hajaC^al-
ildinia, Prdceediiigs 111 the ( !alil(iriiia .\eadem\ (if Scii'iices
47l]i: 1 - l.'x

hilernaliciiial (^mimissien on /iidldgical Xdmeiiclatmc
il(,'/.\', 190!), Intern. ilidii.il Code of Zdological Nomcii-
elalme. pdiiiih fklitidn. Internatidiial Trust Idi' Zodldslical
Noiiirnclalure. I^ondon.

THE NAUTILUS 115(4):151-152, 2001

Paac 151

Book Review

Bahamian Scasliells: a Thousand Species
from Abaco, Bahamas.

Colin licdfcrn. 2001. HaliaiiiiaiiM'.islu'lls.cniii. iiif.. |-4(ii'.i
Raton, i\ + 2S0 pp.. 121 plates (IS in colori. So h\ 11
in., papi-rhack. $114.00

Till' picscntalidn dI a regional nioilu.sfan lanna is eon-
straiiR-tl In its own special set oi problems, inehuliiiti,
completenes.s, cnrreiic\ ol nonienelature and s\steniatii's
of the wide.st ili\ersit\ ol taxa. anil rigor ol ilata drawn
from resources l)e\ond the author's empiric stud\, such
as the literature' and iiinscinii iccoids. These daunting
caveats notwithstaniling, Colin Hedlein has iiearK sin-
gle-handedly chronicled a 37-\t'ar stud\ ot a maiine nioi-
luscau faunule inhabiting a relatiwh' small chain ot is-
lands, niosth' on its shallow Hanks, at the northern ram-
part of Bahamas Bank. B\ appKing his owti photograph-
ic and computer skills. Hedlein has cicated an illustrated
ta\onom\' that will seixe woikeis in xiiinalK e\er\
branch ol marine malacolog\' as well as eonsenation,
ecology. e\olution, zoogeograpin — not tn mention seri-
ous hobb\ists. who, like the anthoi', aie impelletl toward
a better appicciation ot the natural world and its order
The work begins with a short introduction, which in-
dicates the geographic and ecological st-opr ol the re-
port, the magnitude of the matc^rial stn<hrd UOS,()00
i well-chosen specimens, essentialK all personally collect-
! ed o\er 30 xears with Ifi xears in residence); in the In-
' troduction, Redferri also describes his collection tecli-
niqnes, which were di\<-'rse but emphasized drift and
sediment material, as well ;is his coinputer and photo-
graphic techniques. He cites the neark e\clnsi\e use <if
personaltx' collected Abaco specinu'us in liis discussions
and illustrations — tlie ouK important exception lieing
primaiT tvpe mateiial not in his possession.

There follows a 24tt-page sxstematic treatment of
1000 mollusks (777 gastro|iods. 10 scaphopotls. 21 po-
Ivplacophorans, 211 bixaKis, and 2 eephalopods). Ex-
cept for the latter group, this is cii'arK' an exhaustive
treatment for such a small geographical area, and the
gastropoil element is stmmingly di\erse. Each taxon is
presenteil with full scientiRc name (under the respecti\e
class, famik-, and subfamiK ), in modern-day .systematic
order, along with citation ol illnstration(s) in e\en,- in-
stance. Exceptional are about 147 taxa, which, because
of their taxonomic noxeltv, are iilentifietl only to the gi'-
neric, rareK" famiiial le\el, cununf il faiit. Then' follows
a generalK- substantial paragiaph that inclntles a detailed
description, maxinnun size, antl frecjnently more specific
reference(s) to illustration(s) of special moqihologieal
features, and random conunents on sMionymy, system-
atics, and zoogeographx at the conclusion. A shorter par-

agraph entitled "Occnrrenee in Abaco" gi\es a sense of
lic'(|n(nc\. bath\ iiielric r,in'j;e. h.ibilat. habits, and pics-
cnic ni beach drift,

.\ lonr-page glossan and ,ni 1 1 -pa'.ic- bibli(iL;i,i|)li\ con-
ilnde the text sectioir

Till' lOd black-and-whiti' and IS cnlor plates are com-
prised ol iiM'i 2700 photographs antl 7.5 Sl'l.Ms — these
latter ixernlid b\ I'jnilio {{okin. The selection and ex-
I'cnlion ail' M'IA good to I'Mellrnt — rixaled oiiK b\ the
finest woiks ol tins sort, I'artuularK striking landr/r /•/-
lliifur^ are the enln} ini.i^es ol all the several dozen shell-
less ',iastiopiids ti'eati'd in the work.

rlie lS-pa'j,i'. thii'i'-colunm index features reversed
species names (specific epithet loliiiwed In m-nns name),
comnio;! nami', and all supraspecific taxa. Enll listings of
genus-species and revcrseil common names are not in-
cluded.

The strength ol this stuiK lies in three principal areas:
the undeqiiiming ol priiilueti\i' and well-documeTited
field wiirk. seholarK cinalion and taxonomic research,
which .ii'cnrateK exploits xirtnalK' all modern (and some
essentialK merlooked ilassicali works, and the organi-
zation and qualitv of the illustrations, 'i'lie sviilhesis of
these strengths is t\pihed in the tieatment ol the taxo-
nomicalK' nettlesome groups (e.i;.. \itrinellidae. (ieri-
thiopsidae. Triphoridae, Eulimidae. .Maiginellidae. Tnr-
lidae. and ( ialeoinmatidae >, whieli ,ire dejMcted as thoi-
ougliK and clearly as in an\ modern work and nineh
cleari'r than the vast majoritv.

The editorial execution is as close to jieilei't as an\
work in I'xistence. .-Mtei' sexcral horns ot rexiew. oiiK a
lew trivial problems wi're encountered, 'i'heie aie Odo.s-
tmnid species ( .' and Iv but no D is treated. Species 602
Sliiliiji\i\ sp. and sjiecies 02S CV/rr/zo/n/.v dctoiiti appear
to be congeneric and bettei- assigned to Bdclcridiliiii
Thiele, 1929. Noinenelatorial minutiae inclutle (correc-
tion): Eiilitliidiiiiii tlialdssicolnin [ihaltissirola). I'ctalo-
coiiclius floiidaiKi {lloriddiiiis). Miiivxiclla mac^iiitiji
(ntc^iiiti/i). Anfiilis ccniluin i('crafa). Ctciioidis srahm
isodivr). The bibliographx is uearK' exha\isti\e. but I was
stMuied b\ the absence ol at least three rclerences. Ba-
\a\ (1922). .\Iorch (lS7Hi. :nid Nowell-Usticke (1969),
whii'h are essential because thev originalK describe
man\ ■forgotten" but relevant taxa that are defined for
the first time in the moik'rn literature in Hedlern's work,
Tliese references wci'e easiK lonnd in the compilation
by Mikkeksen, P. M., H. Bieler and W. K. Petit i 1993), a
reference that was also missing.

I'Ins wnrk is the finest regional fauna! treatment to
enter llie annals nl western .Atlantic malacologX' in (ner
a ileiaile. It will prove essential to any serious student
in this and lelateil disciiilines lor man\ times that inter-
\al of time. The book is at present availabli' only from

Pime 152

THE NAUTILUS. \ol. 115, No. 4

(Ijaliamanianscaslu'lls.c'oin) ,iik1 TIk' Haili'\-Mattlu"
Shell Museum.

LITER.\TURE CMTED

Ba\a\, .\. 1922. Sables littoraiix tic la mer de las Antilles prin-
enant des ahords de CdIoii et de Cuba. Bulletin dn Mu-
seum National d'Histoire Naturelle 28: 42.'3— 12S.

Mikkelsen. P. M.. R. Bieler and R. E. Petit. 199.3. A liililioji-
rapln of Caribbean nialacolocT\- 1826—199.3. American
Malacological Bulletin 10: 267-290.

Mfirch, O. A. L. 1876. S\iiopsis Mollusconnn Marinoruni In-

diarnni Occidentaliuni nupnmis Insul.uinn Danicannn.
Malakozoologische Blatter 23: 4.5-58. 87-143.
Nowell-Usticke, C. W. 1969. A supplcmcntan' Listing of New
Shells (illustrated). To be added to the check list of the
niariue shells of St. Croix. PrnatcK published. 32 pp.. 6
pis.

Haii^ G. Lee, MD

Suitc'.500

1801 Barrs Streeet

Jacksonville. FL 32204 USA

shells@hglee.com

P/fTS

Tliis puhlicatiiiii is spunsorei! in jiart 1)\

tile State of Florida, Department of State.

DKi.sion of Cultural Affairs, aiul

the Florida Art.s Coiuieil

FHE NAUTILUS

\()lume 115
2001

AUTHOR I\i:)FA

BoNFiTTO, A -S4 M 1 1 .( isi , A\ i( II, p. 39

Cledon, M 15 Mninssi. M 84

Coles, B. F 1 05 N i- k. )i , \. | ( : I (15

COONERT. G. A I OwiA, B 77

DE-Souz.^, P. J. S 1 l'i-\( iiAs/\i)i-ii, P K 15. 39

DuERR, R 147 PiTiT, R. E 35. Ill

G,\RciA. F. J 55. 15(1 Hi II), D. G 115

Geiger, D. L 77 Ivijii, B 140

Ituarte, C. F. 50 .S\ni:i,i,i, B ,S4

Kantor, Yu. 1 99 .Si iiMKi,'/, G. W 22, 45

Kii.BURN, R. N 99 Stom:, |. R 90

Leal. J. H 37 Tkom < ib.i ). J . .S 55, 1 5(1

Lee, H 151 \ M.des. A 29

Macia. S 62 Wise. J. B 68

Marshall. B. A 36

NEWT.AX.^ PROPOSED IN \()LU.\IK 115 (2()(»1)

GASTROPODA

Arrhiti'ctonica ^cinincsd Stiiiiielz. lu'w sp(>cies (Architectoiiiridac) 23

AiistrodaphncUd i/ciiu-iu-iisis Bonfitto. .Morassi and Sabelli. 2001. iirw species (Turridae) 85

BuUata gucniiiii de-Soiiza and Coovert. 200 L new species ( Marsjincllidae) 2

Bullata anahiiiric de-Sniiza and Coo\ci1. 2001. iii'w species (Martjiiicllidae) 5

Ctilliosliiiiiii kiiiKiluiniiii .MarsJiall, 2001, new iiaiiic (CalliostoiiialHhic! 36

Ci/praccii.ssis clupoliina Duerr, 2001, new species { C^assidae ) ] 47

Faiorintis clcnalcxiac Garcia and Troncnso. 2001, new species (Aeolidiidae) 55

Gastrocoptti ( Gastroatpla ) ro'^crscnsifi Nekola and Coles, 2001 , new species ( Pnpillidae) 109

Gmnosdlariiiin fioiiddinim Schmelz, 2001, new species (Arcliitectonicidae) 26

Heliacus iToriitistii) calliouncnsis Schmelz. 2001. new species (Arcliitectonicidae) 24

Heliacus ( Torinista I coinpiicttis Schmelz. 200 1 , new species ( Architectonicidae ) 25

Littoraria {Littoriii(iji\i^ > henaalcnsis Reid. 200 1 . new species ( Littoiinidae ) 120

Mitm (Fu.simitr(i< i-iilliniiiicii'iis Schmelz. 2001. new species (Mitridae) 46

Scdbrifolii iSiiiiiiisitiiiii I chijiohiiui Schmelz. 2001 . new species (Mitridae) 47

BI\AL\IA

Pisiiliinii ihiijuiiditinn Ituarte. 2001. new species (Sphaeiiidae) 50

HEN'IEW KHS FOR XOlAi.ME 115

Hans Bertsdi William G. Lvons Richard E. Petit

Riidii^er Biclcr Marta J. de Maiiitenon Harold G. Pierce

liacliel C'ollin Bruce A. Marshall Rohert Robertson

Robert T. Dillon, |r. Glaus Meier-Brook Gar\' Rosenberg

Garlos S. Gallardo Paula M. Mikkelsen Gar\' Schmelz

Terrence M. Gosliner Peter B. Mordan [olin Slapcinsk-s'

M. G. HarasewNch Diarinaid O Foighil |olni D. Ta\lor

Da\id G. Herbert Shun-ichi Ohgald Richard G. Willan

Yuri I. Kaiitor Guido Pastorino |ohn B. \\'ise

Richard N. Kilburn Tiniothv Pearce

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