1\/IBL/WH01

THE NAUTILUS

Volume 109

1995

AUTHOR INDEX

Anseeuw, P 117

CoovERT, G. A 43

CoovERT, H. K 43

GlTTENBERGER, E 38

GOFAS, S 14

Goto, Y 117

Groves. L. T 113

Kreiser, BR HI

Martins, A M. F 1

MiTTON, J. B Ill

Povel, G. D E 38

Shen, J 41

Squires, R L 113

TippETT, D. L 127

Vermeij, G. J 120

NEW TAXA PROPOSED IN VOLUME 109 (1995)

GASTROPODA

Pleurotomariidae

Perotrochus metivieri .\nseeuw and Goto, 1995, new species 117

Barleeidae

Barleeia aemilii Gofas, 1995, new species 19

Pseudodiala aequinoctialis Gofas, 1995, new species 26

Barleeia chefiae Gofas, 1995, new species 19

Barleeia cinguloides Gofas, 1995, new species 22

Pseudodiala corollaria Gofas, 1995, new species 26

Lirobarleeia elata Gofas, 1995, new species 28

Pseudodiala niso Gofas, 1995, new species 24

Barleeia pervulgata Gofas, 1995, new species 20

Barleeia picta Gofas, 1995, new species 24

Lirobarleeia pupoides Gofas, 1995, new species 29

Lirobarleeia sublaevis Gofas, 1995, new species 28

Barleeia taeniolata Gofas, 1995, new species 20

Barleeia tomensis Gofas, 1995, new species 20

Barleeia verdensis Gofas, 1995, new species 17

Cypraeidae

Proadusta goedertorum Groves and Squires, 1995, new species 114

Cystiscidae

Granulininae Coovert and Coovert, 1995, new subfamily 73

Persiculinae Coovert and Coovert, 1995, new subfamily 70

Plesiocystiscinae Coovert and Coovert, 1995, new subfamily 66

Plesiocystiscus Coovert and Coovert, 1995, new genus 66

Marginellidae

Austroginellini Coovert and Coovert, 1995. new tribe 73

Serrataginella Coovert and Coovert, 1995, new genus 82

Prunini Coovert and Coovert, 1995, new tribe 89

Turridae

Drillia fClathrodrillia) dautzenbergi Tippett, 1995, new name 129

Clalhurella eversoni Tippett, 1995, new species 129

Feniinorea kathijae Tippett, 1995, new species 132

Fenimorea petiti Tippett, 1995, new species 132

Drillia (Clathrodrillia) petuchi Tippett, 1995, new species 133

Sediliopsis riosi Tippett, 1995, new species 133

Drillia (Drillia) wolfei Tippett, 1995, new species 127

Ellobiidae

Allochroa nana Martins, 1995, new species 8

Allochroa tenuis Martins. 1995, new species _ 8

THE NAUTILUS

Volninc 109, Niimhcr 1

August 31, 1995

ISS\n()2.S-1344

A (juartcrhj devoted

la nudacologij.

EDITOR-IN-CHIEF

Dr. M. G. HarascwTcli

Division of Molliisks

National Museum of

Natural History

Smithsonian Institution

Washington, DC 20560

ASSOCIATE EDITOR

Dr. R.Tucker Ahhott

American Malacologists, Inc.

RO. Box 2255

Melbourne, FL 32902

CONSULTING EDITORS

Dr. Riidiger Bieler

Department of Invertebrates

Field Museum of

Natural History

Chicago, IL 60605

Dr. Arthur E. Bogan

Freshwater MoHuscan Research

36 Venus Wdv

Sevvell, NJ 080S0

Dr. Robert T.Dillon. Jr.

Department of Biology

College of Charleston

Charleston, SC 29424

Dr. Wilhani K.. I'hucrson

Department of Living Invertebrates

The American Museum of Natural

History

New York, NY 10024

Mr. Richard I. jolmson

Department of Mollusks

Museum of Comparative Zoology

Harvard University

Cambridge, MA 02138

Dr. Aurele La Rocque

Department of Geology

The Ohio State University

Cohiinbus. OH 43210

Dr. James H. McLean

Department of Malacology

Los Angeles County Museum of

Natural History

900 Exposition Boulevard

Los Angeles, CA 90007

Dr. Arthur S. Merrill

% Department of Mollusks

Museum of Comparative Zoology

Harvard University

Cambridge, MA 02138

Dr. Paula M. Mikkelsen

Department ol Malacology

Delaware Museum of Natural Histon

RO. Box .3937

Wilmington, DE 19807

Dr. Donald R. .Moore

Division of Marine Geologv

and Geophysics

Rosenstiel School of Marine and

Atmospheric Science

University oi Miami

4600 Rickcnbackcr f ^ansrwax

Miami. FL 33149

Dr. Gustav Paulav

Marine Laboratoiy

University of Guam

Mangilao'. Guam 96923

Mr. Richard E. Petit

RO. Box .30

North Myrtle Beach, SC 29582

Dr. EdwardJ.Petuch

Department of Geologj'

Florida Atlantic University

Boca Raton, FL. 33431

Dr. David H. Stansberv

Museum of Zoology

The Ohio State University

Columbus, OH 4.3210

Dr. Ruth D. Turner

Department of .Mollusks

Museutu of Comparative Zoology

Harvard University

Cambridge, MA 02138

Dr. Geerat J. Vermeij

Department of Geology

University of (California at Davis

Davis, CA 9.5616

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THEe^NAUTILUS

CONTENTS

Volume 109, Number 1

August SI, 1995

ISSN 0028-1344

Antonio M. de Frias Martins

Systematic Revision of Atlochroa Ancey, 1887 (Pulmonata:

Ellobiidae) with an Account of the Anatomy of Allochroa

layardi (H. & A. Adams, 1855) and the Description of Two

New Species 1

A Remarkable Species Richness of the Barleeidae

(Gastropoda: Rissoacea) in the Eastern Atlantic 14

Shell Growth and Decollation in Terrestrial Gastropods 38

Cannibalism in the Terrestrial Slug Deroceras lacve 41

Serge Cofas

Edmund Gittenberger

G. David E. Povel

Jianming Shen

THE NAUTILUS 109(1):1-13, 1995

Page 1

Systematic Revision of Allochroa Ancey, 1887

(Pulmonata: Ellobiidae), with an Account of the

Anatomy of Allochroa layardi (H. & A. Adams, 1855)

and the Description of Two New Species

Antonio M. de Frias Martins

Dt'pailaiiU'iito tie Biologia

Universidade dos A(,ores

P-9502 Ponta Delgada codex

Sao Miguel, A(;ores, Portugal

ABSTRACT

Of the nominal species of Allochroa Ancey, 1887 described to

date, only two are recognized: A. hronnii (Pliilippi, 1846), en-

demic to the Hawaiian Islands, and A. layardi (H & A Adams,

1855), distributed throughout the Indo-Pacific. Examination of

the I'SNM collection revealed two additional species, A. nana

Martins, n. sp. and A. tenuis Martins, n sp,, herein described.

The anatomy of A. laijardi is described and serves as the basis

for determining the relationships of this genus within the El-

lobiidae. Allochroa has a monaulic, entirely glandular pallial

gonoduct and a wide visceral nerve ring with a long right

parietovisceral connective, confirming placement in the Py-

thiinae Its close relationship to the Mediterranean and Eastern

North .\tlantic genus Ovatella Bivona, 1832 is indicated by

similar apertural dentition, protoconch morphology of A. ten-

uis, and b\ the internal structure of the penial complex, where

the penis appears as the development of one of the pilasters.

Key Words: Ellobiidae, Pythiinae, Allochroa, systematics,

anatomy.

INTRODUCTION

The Ellobiidae are a group of primitive, mostly halo-

philic pulmonale mollusks, commonly found near and

above the hightide limit of mangroves, salt marshes, and

boulder and cobble beaches. Morphologically very di-

verse and ranging in length from barely 1 mm (Leu-

conopsis ) to about 100 mm (Ellobium). they nevertheless

share basic shell characters and body features that allow

them to be considered within a single family. Pfeiffer

(1853) was the first to attempt a subfamilial organization,

based on shell characters alone. He assigned those species

with a thick, reflected outer lip to the Auriculea [=EI-

lobiinae], while he included the species with sharp outer

lip in the Melampea [=Melampodinae]. Utilizing radular

and also some anatomical characters, Odhner (1925) rec-

ognized a more complex subfamilial arrangement and

redistributed the ellobiid genera among four additional

subfamilies: Carychiinae, Pedipedinae, Pythiinae and

Cassidulinae. Odhner s outline was supported on con-

chological features by Zilch (1959). The first detailed use

of anatomical characters, in a review that included spe-

cies belonging to 11 genera, was by Morton (1955). His

study supported Odhner's classification, but the Cassi-

dulinae were included in the Pythiinae, on the basis of

similarities of the reproductive system. In a study of

species belonging to 17 of the 23 genera currently rec-

ognized in the Ellobiidae, Martins (in press a) clarified

Morton's (1955) findings by redefining the subfamilial

morphological patterns of the reproductive and nervous

systems.

Preserved specimens of Allochroa layardi (H. & A.

Adams, 1855) are used here as the basis for detailed

anatomical investigations of the family extending the

preliminary findings reported in Martins (in press b).

Insight on the anatomy of A. layardi provided infor-

mation needed to define the phylogenetic relationships

of Allochroa Ancey, 1887 within the Pythiinae. The pres-

ent work reviews the taxonomy of the species of Alloch-

roa and describes the anatomy of A. layardi. Examina-

tion of the Mollusk collection at the National Museum

of Natural History, Smithsonian Institution, consisting of

only dry material, revealed the existence of two new

species, herein described.

MATERIALS AND METHODS

This taxonomic revision is based primarily on dry spec-

imens of Allochroa in the USNM collection. Four spec-

imens of A. layardi, collected on Henderson Island in

1991, were provided for anatomical work. The fully re-

tracted animals were preserved in alcohol and, except

for the eyes and the tip of the tentacles, appear to have

lost all other pigmentation. Two specimens were dis-

sected under a binocular microscope and drawn using a

camera lucida.

Shells with protoconchs were mounted using double-

Page 2

THE NAUTILUS, Vol. 109, No. 1

coated tape. Radulae were stretched across a cover slip

after cleaning with KOH solution. Both structures were

mounted on stubs and examined and photographed using

a Cambridge Stereoscan 250 Mk2 Scanning Electron Mi-

croscope.

Acronyms for specimen repositories are as follows:

ANSP: The Academy of Natural Sciences, Philadel-

phia

BMNH: The Natural History Museum, London

MCZ: Museum of Comparative Zoology, Harvard

LIniversity, Cambridge

MNHN: Museum National d'Histoire Naturelle de

Paris

USNM: National Museum of Natural History, Smith-

sonian Institution, Washington, D.C.

KEY TO THE SPECIES OF ALLOCHROA ANCEY,

1887:

1. Shell thick 2

Shell thin .'3

2. Aperture oval elongated (base narrow) A. bronnii

Aperture auriculate (base expanded) A. layardi

3. Shell length at least 4 mm 4

Shell length less than 4 mm 5

4. Protoconch width less than .5 mm A. layardi

Protoconch width at least .6 mm 5

5. Protoconch without umbilicus and with apertural lip

not reflected A. nana

Protoconch with umbilicus and with apertural lip

slightly reflected A. tenuis

SYSTEMATICS

Class GASTROPODA Cuvier, 1797

Subclass PULMONATA Cuvier, 1817

Order Archaeopulmonata Morton, 1955

Family Ellobiidae H. & A. Adams in Pfeiffer, 1854

Subfamily Pythiinae Odhner, 1925

Genus Allochroa Ancey, 1887

Allochroa Aiiiey, 1887:288 Type species by original designa-

tion: Auricula bronnii Pliilippi, 18-1(1

Description: Shell lengtli to 12 mm, oval-conic, fragile

to .somewhat solid, sometimes hirsute. Umbilicus absent

Spire with up to 7.4 flat to slightly convex whorls, sculp-

tured with more or le.ss marked spiral lines; body whorl

55 to 80% of shell Icngtli, usually with three chestnut

spiral bands over a whitish to light-brown background,

the band just above the aperture consistently strongest,

the one near the suture sometimes faint, the one near

the columella largest, sometimes coalescing w ith the me-

dian band. Aperture about 50 to 60% of shell length,

oval-elongated to .semi-circular, angled posteriorly,

rounded anteriorly; inner lip slightly reflected over the

columella, with three more or less etiuidistant teeth: one

oblique columellar tooth, a more or less horizontal an-

terior parietal tooth, and a sometimes strongest, oblique

posterior parietal tooth; outer lip sharp, usually with one

prominent fold entering the aperture at about the level

of the parietal teeth, forming a somewhat marked anal

canal. Protoconch bulbous, smooth, with up to 1.3 whorls

visible.

Radula: Central tooth narrow, base deeply emargin-

ate, with two median flaring arms, unicuspid; lateral

teeth with elongated, sinuous base with two projecting

asymmetric arms, inner arm lower, crown large and long,

subquadrangular; marginal teeth with sharp endocone

gradually becoming as long as mesocone as row pro-

gresses outward, developing a sometimes conspicuous

basal ectocone.

Anatomy: Reproductive system monaulic, pallial gon-

oducts entirely glandular, pallial gland present. Central

nervous system with wide visceral ring; right parieto-

visceral connective as long as left.

Remarks: The name Allochroa was proposed by Ancey

(1887) to replace the supposedly preoccupied "Laimo-

donta des auteurs [H. & A. Adams, 1853] (non Nuttall)"

(type species Auricula Inonnii Philippi, 1846). "Lai-

modonta Nuttall' , wrongly taken as an emendation of

Laemodonta Philippi, 1846, was restricted to a different

assemblage (type species Auricu/as/ria^a Philippi, 1846),

following Nevill (1878). A more detailed account of the

synonymy of Laemodonta and Laimodonta is presented

elsewhere (Martins, in press a).

Ancey (1887) also made clear that Marinula King,

1832, very similar to Pedipes Scopoli, 1777, was not an

eligible group for his new assemblage, and that Monica

H. & A. Adams, 1855[=Oiw^e'//a Bivona, 1832], although

decidedly conchologically closely related, is restricted to

Mediterranean and West Atlantic species.

Allochroa is conchologically characterized by the pres-

ence of an outer lip internal lamella which, in conjunc-

tion with the anterior parietal tooth, divides the aperture

into two roughly equal halves: a posterior portion, where

the shell is thinner; an anterior, thickened half, some-

times with an additional internal fold on the outer lip.

The inner lip ol the aperture is typically pythiine, i.e.,

roughly equidistantly tridentate, with one small colu-

mellar tooth and two parietal teeth. The lamellar, outer

lip tooth readily separates this genus from the closely

related Laemodonta, which usually has two round tu-

bercles on the outer lip, and also from Ovatella, where

the outer lip lamellae are much shorter. The single col-

umellar tooth sets Allochroa apart from Pedipes which,

as in P. mirahilis (Miihlfeld, 1816) and P. pedipes (Bru-

guiere, 1789), has a strong, crest-like outer lip fold pen-

etrating inside the aperture. In Pedipes, the single pa-

rietal tooth is very strongly pronoimced.

The six s])ecies assigned by authors to Allochroa are

here synonymized into two: the Hawaiian Allochroa

Inonnii (Philippi, 1846) and the Indo-Pacific Allochroa

layardi (H. & A. Adams, 1855). Two additional new

species are described herein.

Kuroda and Habe (1952) listed Pedipes ajfinis F"er-

ussac, 1821, under A//of/jroa, owing prol)ably to the pres-

ence of a strong lamella inside the outer lip. Ferussac

A. M. de F"rias Martins, 1995

Page 3

(1821), however, had mentioned a bidentate cohimella

as characteristic of Pedipes, and Ancey (1887) also used

this feature of P. affinis to separate it from Laernodonta ,

where Jickeli (1874) had erroneously placed it. Following

the above reasoning, I concur with Ancey (1887) in con-

sidering Ferussac's species a Pedipes.

Similarly, the New Caledonian Pedipes forestieri Sou-

verbie and Montrouzier, 1864, has been erroneously as-

signed to Allochroa (Franc, 1954; Smith, 1992). In the

original description Souverbie and Montrouzier (1864a)

mentioned the presence of two columellar teeth, stating

that their observations were based on a single, probably

juvenile specimen. In a later paper (1864b) the same

authors, reacting to the epistolary comment of L. Pfeiffer

on that subject "Spec, tiov., sed Marinida". changed the

ta.xonomic combination to "Marinula (an Pedipes?)",

cautiously justifying, however, their original generic

choice by pointing out features such as " 1 ) shell positively

striated transversely; 2) two positively columellar folds;

3) external border of the columellar lip contouring be-

hind the upper columellar fold as in Pedipes mirabilis,

Miihlf., which we possess." The accompanying figure

(Souverbie & Montrouzier, 1864a: pi. X, fig. 1) appro-

priately illustrated their reasoning. Franc (1954) claimed

to have found an adult specimen of Souverbie and Mon-

trouzier s species in the Jousseaume collection at the

MNHN, and assigned it to Allochroa. The redescription

of Pedipes forestieri provided by Franc (1954), mentions

the presence of a spiral fold inside the outer lip char-

acteristic of Allochroa, but corroborates the presence of

"two columellar folds closer to each other than to the

parietal", a feature diagnostic of some pedipedine gen-

era, namely Pedipes and Marinula. Taking into account

Souverbie and Montrouzier's (1864a, b) insistence on the

pedipedine characteristics of the apertural dentition, a

feature confirmed by Franc (1954), and in my inability

to examine the type material, I consider the assignment

of Pedipes forestieri to Allochroa to be unwarranted.

The similarity of the basic patterns of shell morphology

and color throughout the genus creates some difficulties

in separating species. Larger specimens of the highly

variable, widely distributed A. layardi overlap in shape

and color with smaller specimens of the Hawaiian A.

hronnii (compare Figs. 1 and 5). At the other extreme,

thin, less distinctly banded forms of A. layardi could be

confused with lighter specimens of A. tenuis, n. sp. (com-

pare Figs. 10 and 19) However, species can be readily

separated on the basis of other conchological characters

such as the ratio number of whorls/shell length or using

protoconch features.

Allochroa bronnii (Philippi, 1846)

Figures 1-2, 21, 31-32

Auricula Irronnii Philippi, 1846;98 [Hawaiian Islands; location

of type unknown]. Auricula sandwichiensis Souleyet, 1852:

524, pi. 29, figs. 29-32 [Hawaiian Islands; type not found,

supposedly at BMNH (Gray, 1855)]. Ophicardelus (Lai-

modonta) sandwichensis (Eydoux & Souleyet). H. & A.

Adams, 1855a: 34 [misspelling of santftuic/iiensis]. Melam-

pus bronni (Philippi). Pfeiffer 1854:147. Ellobium hrownii

(Philippi) H & A Adams, 1855b:237 [misspelling of fcron-

nii] Laimodunta sandwichensis (Eydoux & Souleyet). H.

& A Adams. 1855b:246, pi. 83, figs 1, la [misspelling of

sandwichiensis]. Mclampus sandwichiensis (Souleyet).

Pfeiffer, 1856:50. Ophicardelus sandwichensis (Eydoux).

J. E. Gray in M. E. Gray, 1859:(3), pi. 304, fig. 2; (4):1I8.

Laernodonta Inonni (Philippi). Martens & Langkavel, 1871:

57. Marinula {Laernodonta) bronni (Philippi). Nevill, 1878:

220. Allochroa Inonni (Philippi). Ancey, 1887:288. Tralia

{Laernodonta) Inonni (Philippi). Crosse, 1894:321. Me-

lampus (Laimodonta) Ifronnii (Philippi). Kobelt, 1898:

200, pi. 23, figs. 1, 2. Laimodonta bronni (PhiMppi). Dautz-

enberg & Fischer, 1905: 107

Description: Shell (Figs. 1-2, 21) length to 12 mm, ovate-

conical, somewhat solid; violet-brown to light-brown,

banded. Spire moderately high with up to 7.4 flattened

whorls, with marked spiral striae crossed by irregular

growth lines that give the surface a matte appearance;

one light-brown subsutural band, a median white band

followed by a dark-brown band. Body whorl averaging

60% of shell length, sculptured as spire; coloration pattern

showing a subsutural light-brown band followed by a

marked, white, narrow band halfway between the suture

and the aperture, a wide, dark-brown band extending

just below the posterior apertural angle, a whitish to

brownish band as wide as the previous one gradually

merging with the dark-brown, anteriormost band that

extends to the anterior tip of the aperture. Aperture oval-

elongated, rounded anteriorly, acute posteriorly; inner

lip with three subequidistant teeth: one oblique colu-

mellar tooth faintly continuous with a callus that runs

along the anterior border of the aperture and connects

to the tip of the outer lip tooth; two parietal teeth, the

anterior one half the length of the aperture, perpendic-

ular to the columellar axis and continuing inward; the

posterior one usually strongest, not continuing inward;

outer lip sharp, with a ridge-like tooth about opposite

the anterior parietal tooth, penetrating inside the aper-

ture for about half a whorl; a wide callus develops an-

terior to the outer lip tooth. Protoconch 235x155 ^m,

whitish, smooth, somewhat inflated, barely over one

whorl, with elongated, narrowing lip (Fig. 21).

Radula: (Figs. 31-32) Formula [31-h24-H-l-24-l-31]

X 100-1-. Central tooth slightly above the plane of the

lateral teeth; base narrow, subquadrangular, deeply

emarginate with medial lateral projections flaring out;

crown less than half the length of the lateral teeth, nar-

row, unicuspid: mesocone somewhat sharp. Lateral teeth

24, base elongated, slightly bent medially, with lateral

projections flaring out asymmetrically, the inner projec-

tion smaller, at about the first third of the base, the outer

one at about mid-length of the base; crown covering three

quarters of base, quadrangular, with a longitudinal de-

pression in the middle. Marginal teeth 31; basis barely

longer than crown, diminishing gradually to completely

disappear under crown, developing a basal ectocone;

crown bicuspid, endocone sharp, short on first teeth,

gradually elongating towards the end of the row, me-

socone long and narrow, blunt at tip.

Page 4

THE NAUTILUS, Vol. 109, No. 1

A. M. de Frias Martins, 199.5

Page 5

Remarks: The Hawaiian Allochroa bronnii is the largest

species of the genus, sometimes reaching 12 mm in length

(Souleyet, 1852; USNM 131288). It has been confused

with larger forms of the highly variable Allochroa lay-

ardi, an Indo-Pacific species (Dautzenberg & Fischer,

1905). They can be set apart by the narrower base of the

aperture in A. bronnii, whereas shells of identical size

of A. latjardi have a wider apertural base (see Figs. 1

and 5). In Hawaii, where both species overlap, they are

morphologically very distinct (see Figs. 1, 2, 4). Although

size alone seems to be sufficiently diagnostic to separate

these two species, A. bronnii shows also a proportionally

smaller number of whorls for the length of the shell.

However, anatomical studies are needed to ascertain bet-

ter the differences between these two closely related spe-

cies.

Habitat Notes: "Found seaward of Melampits, Pedipes,

and Laemodonta. often buried in gravel in high tide

pools and along rocky shores" (Kay, 1979).

Range: Allochroa bronnii is endemic to the Hawaiian

Islands.

Material Examined: (Sample size in parenthesis). HA-

WAIIAN ISLANDS: [USNM 20795 (5), 31582 (4), 98175

(6), 98176 (9), 98178 (3), 131288 (1), 319528 (4). 346615

(11)]; HAWAII ISLAND: Keei, near Napoopoo, Kona

Coast [USNM 767596 (1)]; OAHU ISLAND: Honolulu

[USNM 159574 (1)]; Pearl City, Honolulu [MCZ 55989

(1)]; 1935 dredgings, Hickham Air Force Base, Pearl

Harbor [USNM 484416 (3)]; Kaneohe Bay [USNM 887316

(8)].

Allochroa latjardi (H. & A. Adams, 1855)

Figures 3-14, 22-27, 30, 33-35, 39-45

Melampus layardi 'Ads.' Pfeiffer, 1854:147. Nomen nudum^

Ophicardcltis (Lainwdonta) latjardi H & A Adams, 1855a:

35 [Ceylon, lectotvpe herein selected BMNH 196980/1

(Fig. 3); two paralectotypes BMNH 196980/2-3]. Lai-

modonla layardi (H. & A Adams) H. & A. Adams, 18.55b

246. Lainwdonta pfeifferi Dunker in Pfeiffer I860: 201

[Japan; location of type unknown); Dunker, 1861:25, pi

2, fig. 19. Lainwdonta conica Pease 1863: 242 [Pacific

Islands; lectotype selected by Kay (1965) BMNH 1964292

(Fig. 4); five paralectotypes BMNH 1964293]. Laimadonta

conica Pease. Pease, 1868101, pi 12, fig 15 (misspelling

of Laimodonta). Lainwdonta anaaensis Mousson, 1869:

63 (Anaa Island, Paumotu group; location of type un-

known]. Lacmodo7ita conica (Pease) Martens & Lang-

kavel, 1871:57, Ptecotrema anaaensis (Mousson). Paetel,

1873:114. Lainwdonta hronii var. conica (Pease). Jickeli,

1874:178, pi. 7, fig, 3. Melampus pfeifferi (Dunker). Pfeif-

fer, 1876:319. Melampus co7iicus (Pease). Pfeiffer, 1876:

319. PMelampus anaaensis (Mousson). Pfeiffer, 1876320.

Marimtla (Laimodonta) conica (Pease). Nevill, 1878:220.

Marinula (Laenwdonta) conica (Pease). Martens, 1880:

208. Laimodonta conica var conicoides Tapparone Ca-

nefri, 1883:240 [Aru Island, New Guinea; location of type

unknown]. Laenwdonta (Tralia) layardi (H. & A. Adams).

Crosse, 1894:321. Melampus (Laimodonta) conicus (Pease).

Kohelt, 1898:224, pi. 26, fig. 14. Melampus (Laimodonta)

anaaensis (Mousson). Kobelt, 1898:228, pi. 27, figs. 10-11.

Laimodonta bronni var. producta Dautzenberg & H. Fi-

scher, 1905:108, pi. IV, figs 20, 21 [Anse du Crane, Bah'i'e

d'Along, Tonkin; syntypes MNHN (Fig, 5)]. Enterodonta

conica (Pease), Germain, 1921:264. Allochroa conica

(Pease), van Benthem Jutting, 1941:283. Allochroa pfeif-

feri (Dunker) Kuroda & Habe, 1952:38. Allochroa layardi

(H. & A Adams) Franc, 19.54:365.

Description: Shell (Figs. 3-14, 22-27) length to 8 mm,

ovate-conical, fragile to somewhat solid; chestnut-brown

to pale-yellow, banded. Spire moderately high, whorls

6.0 to 6.5, plano-convex, with spiral striae crossed by

irregularly marked longitudinal growth lines which give

the surface a faint granulose appearance. Body whorl

about 70% shell length, sculptured as the previous whorls;

color pattern showing a faint subsutural light band, a

marked whitish band halfway between the suture and

the aperture, also visible on the two previous whorls,

sometimes with a third, wider pale-yellow band gradu-

ally becoming dark-brown anteriorly. Aperture auricu-

late; inner lip tridentate; one oblique columellar tooth

not reaching the border of the lip; two parietal teeth,

anterior lamellar, transverse, continuing inwards, pos-

terior largest, thick and bifid in larger individuals, slight-

ly elongated but not continuing inwards; outer lip sharp,

with a ridge-like tooth about halfway along the length

of the aperture, penetrating deep inside the aperture; a

wide callus may develop anteriorly, as a continuation of

the outer lip tooth, also penetrating inwards; a second

outer lip tooth is sometimes present over the callus, lo-

cated at a level between the anterior parietal tooth and

the columellar tooth. Juveniles pilose, the hairs distrib-

uted along the spiral striae (Fig. 12). Protoconch 210-

235 X 150-175 nm whitish, smooth, somewhat inflated,

a little over one whorl, with rounded peristome (Figs.

22-27).

Figures 1-2. Allochroa bronnii (Philippi); 1. Hawaii, USNM 98175, si = 8 2 mm; 2. Pearl City, Honolulu, Hawaii, MCZ 55989,

si = 3.5 mm.

Figures 3-12. Allochroa layardi (H. & A Adams);3. Lectotype, Ceylon, BMNH 196980.a, si = 7.0 mm, 4. Lectotype of Laimodonta

conica Pease, Pacific Islands [Hawaii], BMNH 1964292, si = 4.2 mm; 5. Holotype of Laimodonta bronni var, producta Dautzenberg

& Fischer, Bah'ie d'Along, Anse du Oane, Tonkin, China, MNHN (not numbered), si = 8 0 mm; 6. Paumotu [Tuamotu] Islands,

specimen illustrated in Pease (1868, pi, 12, fig. 15), ANSP 22610 [type written in label], si = 7 0 mm; 7. Mataira Island, Raroia,

Tuamotu Islands, USNM 698771, si = 5 8 mm; 8. North Beach, Henderson Island, Pitcairn Islands, R C. Preece coll , si = 6.1 mm;

9. Lijeron Island, Jaluit Atoll, Marshall Islands, USNM 6600.35, si = 4 6 mm; 10. Lugbon Island, Philippines, USNM 201147, si =

.5.5 mm; 11. Riu Kiu Islands, Japan, USNM 345877, si = 5.0 mm; 12. Hirado, Hizen, Japan, USNM :345878, si = 4.1 mm. si =

shell length.

Page 6

THE NAUTILUS, Vol. 109, No. 1

Figures 13-14. Allorhroa layardi (H & A. Adams), Mataira Island, Raroia, Tuamotu Islands, USNM 698771, si = 3 5, and 2.5

mm.

Figures 15-18. Allochroa nana Martins, n. sp., Eninman Island, Bikini, Marshall Islands; 15, 17-18. paratypes, USNM 585963,

si = 2.2, 3.7, and 3.0 mm; 16. holotype, USNM 585963.a, si = 3.2 mm Figures 19-20. Allochroa tenuis Martins, n sp , Noumea,

New Caledonia; 19. holotype, USNM 724374.a, si = 4.1 mm; 20. paratype, USNM 724374, si = 3.5 mm. si = shell length.

Radula: (Figs. 33-35) formula [26 + 23+1+23 + 26] x

115. Centra! tooth slightly above the plane of the lateral

teeth; base deeply emarginate, with lateral projections

flaring out at mid-length, gradually tapering to neck

region; crown half-length of that of lateral teeth, narrow,

unicuspid; mesocono somewhat sharp. Lateral teeth 23;

base elongated, slightly bent medially; a rounded inner

prominence arises on the lower third, an outer projection

flares out at mid-length and tapers towards neck; crown

covering two thirds of base and as wide as the latter,

unicuspid; cusp rounded. Marginal teeth 26; bases of first

teeth barely longer than crown, gradually tliniinishing

to completely disappear underneath crown, developing

a conspicuous lateral basal cusp on the last 13 teeth;

crown bicuspid; endocone, sharp, short on first teeth,

gradually elongating to about three quarters of meso-

cone; mesocone as long as that of last lateral teeth, round-

ed at tip.

Visceral mass: Conical, not following external winding

of shell, with whorl impressions weakly insinuated, in-

dicating the extensive resorption of the iiuier whorls of

the shell.

Digestive system: Mandible (Fig. 39) with fibrous ap-

pearance, tripartite; median portion roughly quadran-

A. M. de Frias Martins, 1995

Page 7

gular with extremities abruptly tapering posteriorly, lat-

eral portions of sparse, loose fibers, tapering on both

extremities. Salivary glands long, cylindrical. Stomach

(Fig. 40) tripartite; cardiac region dilated, thin, receiving

at its base the duct of anterior diverticulum; mid-portion

comprising most of the stomach's volume, muscularized;

gastric caecum thin, pouch-like, receiving posterior di-

verticulum; pyloric region small, thin.

Reproductive system: (Fig. 30, 41-44) Ovotestis of

long, inverted claviform acini embedded in the conical

posterior lobe of the digestive gland, connecting with a

collecting duct by their thinner ends, the broader, dark-

red dotted extremities appearing at the surface of the

digestive gland as an irregular, winding band; proximal

half of the hermaphroditic duct thin, relatively straight,

distal half forming a greatly convoluted and somewhat

dilated seminal vesicle; pallial gonoduct hermaphroditic

to the vaginal aperture; albumen gland beneath the an-

terior lobe of digestive gland and intestinal convolutions;

posterior mucus gland tightly convoluted; anterior mucus

gland and prostate gland covering entire length of pallial

spermoviduct; bursa spherical, fitting under posterior

mucus gland; bursa duct nearly equal in length to sper-

moviduct, emptying near vaginal opening. Pallial gland

long, sharply bent about midlength, anterior portion more

dilated and slightly longer than posterior one, emptying

at the female genital opening. Penial complex relatively

short, proximal third dilated; vas deferens adhering to

penial sheath, entering penial structure at the distal end

of the proximal quarter; penis oval-oblong, blunt, con-

stituted of a thickened, sharply bent, compactly folded

muscular pilaster, lodged in the thin, membranous di-

lation of the penial sheath; internal wall of the penial

sheath with fine, radiating grooves at the site of the

dilation, with two irregularly grooved longitudinal folds

running forward to the aperture, one of them connecting

distally to the penis, the other originating at the level of

the tip of the penial papilla; penial retractor about as

long as the penial complex, relatively thick.

Nervous stjstem: (Fig. 45) Cerebral commissure short,

about half the width of a cerebral ganglion; left cerebro-

pleural connective twice the length of the right one; right

cerebro-pedal connective somewhat longer than the left

one; pleural ganglia small; right pleuro-parietal connec-

tive very short, about onethird of the left one; left parietal

ganglion as small as pleural ganglia, right one three times

larger, with a thick pneumostomal nerve from where a

rudimentary osphradial ganglion buds off; parieto-vis-

ceral connectives relatively long, the left one somewhat

longer.

Remarks: Allochroa layardi was described by H. & A.

Adams (1855a) from specimens from Ceylon [Sri Lanka],

in the Cuming collection. H. & A. Adams' manuscript,

although submitted for publication in 1854, was not pub-

lished until January 1855 (see Duncan, 1937; Trew, 1992).

Pfeiffer, who had access to the manuscript of the Adams

brothers, listed their new species in his Synopsis (1854),

giving credit to the original author(s) and placing the

species under the genus Melampus. Pfeiffer, however.

did not add any descriptive notes; for this reason, the

name "Melampus layardi 'Ad.' Pfeiffer' should be treat-

ed as nomen nudum. The Adams brothers' name was

apparently ignored in the various subsequent descrip-

tions of taxa here considered as synonyms of A. layardi,

and comparisons referred rather to A. hronnii. Such was

the case of Laimodonta pfeifferi. from Japan, and of

Laimodonta Inonni var. producta from China (see Fig.

5). Allochroa laijardi, however, has a more slender over-

all shape and is smaller than A. bronnii, mainly in Hawaii

where both species coexist. Pease (1863) described Lai-

modonta conica from the Hawaiian Islands (Fig. 4); five

years later, however. Pease (1868) figured his Hawaiian

species using a specimen from Paumotus [Tuamotus Is-

lands], almost twice the size and more elongated than

the type specimens (Fig. 6). Mousson (1869) described

and figured the larger Tuamotu shells from Anaa Island

as Laemodonta anaaensis. commenting on their clear

relationship with A. bronnii, but at the same time calling

attention to the smaller size of his species relative to the

latter; his measurements (8 mm), however, are the largest

known for A. layardi.

Very variable across the Indo-Pacific, A. laijardi is

usually markedly striated, although some specimens can

be extremely smooth (Fig. 8). It also varies in shell thick-

ness, the thick specimens sometimes showing an addi-

tional internal lamella on the outer lip (Figs. 7, 9, 13),

which Tapparone Canefri (1883) considered to be the

variety conicoides of Laimodonta conica [=A. layardi].

The color pattern remains basically the same throughout;

however, the banding delimitation may sometimes lack

sharpness (Fig. 10). The shape and size of the protoconch

are consistently homogeneous throughout the distribu-

tional range of the species (Figs. 22-27). Allochroa bron-

nii (Fig. 21) possesses a protoconch similar to that of A.

layardi, also stressing the close affinities of the two spe-

cies. However, as pointed out before, the size and ratio

between the number of whorls and the length of the shell

justify their taxonomic separation.

Only one radula of each species was studied. It appears

that the base of the central tooth of A. layardi is decidedly

triangular (Fig. 33), whereas that of A. bronnii is nar-

rowly quadrangular (Fig. 31) The crown of the lateral

teeth of A. bronnii seems to be narrower and longer than

that of A. layardi.

Habitat Notes: Gassies (1863) stated that this animal ap-

pears to be completely marine, and that Montrouzier

found it under stones near the shore, together with col-

umbellids, ranellids and tritons. Preece (personal com-

munication) found them in crevices of exposed rocks at

about high tide level.

Range: Allochroa layardi is widely distributed through-

out the Indo-Pacific; it has been reported from Mauritius

(Morelet, 1882), Sri Lanka, India (Nevill, 1878), China,

Japan, Philippines, New Guinea, Australia, New Cale-

donia, Marshall Islands, Tuamotu Islands, Hawaii.

Material Examined: (Sample size in parenthesis) CEY-

LON [SRI LANKA] [BMNH 196980/1 (lectotype),

Pages

THE NAUTILUS, Vol. 109, No. 1

196980/2-3 (paralectotypes of Ophicardelus (Laimo-

donta)layardi)]. CHINA; Bahie d'Along, Ansedu Crane,

Tonquin [MNHN, not numbered, holotype and paratype

of Laimodonta bronni var. producta]. JAPAN: Hirado,

Hizen [USNM 345878 (30)]; Riu Kiu Islands [USNM

345877 (56)]; Loo Choo Islands [USNM 30811 (12), 161507

(2)]; Nago, Okinawa Island, Loo Choo Islands [USNM

363720 (1)]. PHILIPPINE ISLANDS: Bohol Island

[USNM 302973 (4)]; off Point Origon, Tablas [USNM

281503 (1)]; Liigbon Island [USNM 201147 (7)]; Point

Hinayangan, Leyte [USNM 887317 ex Chicago Academy

of Sciences (3)]; Sogod Bay, Leyte [USNM 283799 (1)];

off Point Dumurug, Masbate [USNM 274834 (1)]; off

Adyagan Island, E Masbate [USNM 281808(1)]; Ragay

Gulf, off Arena Point, Luzon [USNM 284250 (1)]; off

Matacot Point, W Louzon [USNM 259939 (1)]; off Pes-

cador Island, Tanon Station [USNM 281169 (1)]; off Cap-

itancillo Island, N Cebu [USNM 287800 (1)]; off Lanis

Point, E Cebu [USNM 290562 (2)]; Pujada Bay, E Min-

danao [USNM 255647 (1), 276398 (2)]; Tilig, Lubang

Island [USNM 229914 (1)]. NEW CALEDONIA: Lifu

[USNM 422639 (1), 422640 (1), 423167 (4)]. AUSTRA-

LIA: [USNM 887318 (1)]; mouth of Bigota Creek, Barrow

Island [USNM 691788 (1)]. COOK ISLANDS: Tupapa,

E of Avarua, Rarotonga [USNM 721509 (1)]. MAR-

SHALL ISLANDS: N end of Lijeron Island, Jaluit Atoll

[USNM 660035 (8)]. TUAMOTU ISLANDS: [USNM

98177 (3), 711465 (6)]; RAROIA ISLANDS: Ngarumaoa

Island [USNM 697524 (1), 697767 (1), 698079 (2), 720349

(17)]; S end of Ngarumaoa Island [USNM 697593 (3)];

Mataira Island [USNM 698771 (8), 711673 (1)]; Kumek-

umo Island [USNM 698492 (5)]; Teteu Island [USNM

722685 (1), 722687 (1)]; S end of Teteu Island [USNM

722756 (50)]; Havana Island [USNM (1 fragment)]; Okaea

Island [USNM 722834 ( 1 )]; S end of Opakea Island [USNM

722554 (79), 723548 (1)]; Oneroa Island [USNM 720768

(2)]; Ohava Island [USNM 720503 (19)]; Otikaheru Island

[USNM 720684 (1)]; Vahi Tahi [USNM 613207 (1)]; Te-

mao Harbor, Makatea [USNM 629788 (1)]; NE side of

Tepukamaruia Island, Takumo[USNM 723724 (1)]; Motu

Taraururoa, Mangareva Island [USNM 638224 (4)]. PIT-

CAIRN ISLANDS: E end of North Beach, Henderson

Island [R. C. Preece collection (4)]. HAWAIIAN IS-

LANDS: [BMNH 1964292 (lectotype), 1964293 (5 para-

lectotypes)]; USNM 15157 (2), 20796 (3), 20797 (12),

887319 (4)].

Allochroa nana Martins, new species

Figures 15-18, 28

Description: Shell (Figs. 15-18, 28) length to 3.7 mm,

ovato-conical, fragile, spirally striated, dark-brown to

golden-brown, banded. Spire moderately high, 5 plano-

convex, dark-brown to brown whorls, sometimes with a

faint equatorial lighter band; spiral striae undulated due

to crossing by marked growth lines which arc predom-

inant on the early whorls of teleoconch. Body whorl about

70% of shell length, sculptured as previous whorls; band-

ed color pattern as follows: a light-brown, fine band from

the suture to the first spiral stria, a darker band on the

next two striae, a wider, white band covering the next

3-4 striae, a dark-brown band extending to the posterior

end of the aperture, frequently continuing to the anterior

tip of the shell, sometimes broken up by a diffuse, lighter

band extending from the posterior end of the aperture

to about the level of the anterior parietal tooth. Aperture

semicircular, oblique; inner lip equidistantly tridentate:

one small, oblique columellar tooth, two parietal teeth,

the anterior strongest and perpendicular to the colu-

mellar axis, the posterior about as strong as columellar

tooth; outer lip sharp, very rarely with an elongated

thickening opposite the anterior parietal tooth and weak-

ly projecting inside the aperture (Fig. 17). Juveniles pi-

lose (Fig. 15). Protoconch 285x210 ^m, bulbous, light-

brown, with growth lines showing as faint narrow bands,

with elongated, somewhat widening lip (Fig. 28).

Type Locality: Eninman Island, Bikini, Marshall Islands.

From drift line along tide pool at N end of island, outer

reef side.

Material Examined: (Sample size in parenthesis) MAR-

SHALL ISLANDS: Eninman Island, Bikini [USNM

585963 (holotype), 887320 (64 paratypes)]; Rojoa Island,

Enewetak [USNM 587120 (1)]; Pujiyoru Island, Enew-

etak [USNM 587101 (1)]; N end of Bock Island, Rongerik

[USNM 594920 (2)J

Etymology: nanus, i = dwarf.

Range: Known from Bikini, Enewetak and Rongerik, in

the Marshall Islands.

Remarks: Allochroa nana is the smallest species of the

genus, and size alone will easily distinguish it from the

previously described species. It is, however, best char-

acterized by its large protoconch with faint colabral bands

marking the growth lines and with a somewhat widening

lip (Fig. 28).

Allochroa tenuis Martins, new species

Figures 19-20, 29, 36-38

Description: Shell (Figs. 19-20, 29) length to 4.6 mm,

ovato-conical, fragile, spirally striated, purple-brown to

brown, banded. Spire moderately high, 5.5 plano-convex,

spirally striated, dark-brown whorls, the last one darkest,

usually purple-brown. Body whorl about 70% of shell

length, sculptured as previous \\ horls; baniled color pat-

tern as follows: a gradually darkening brown band ex-

tending from the suture to the third stria, a marked white

band over the next three striae, and a more or less ho-

mogeneous brown region extending to the tip of the shell,

sometimes broken up by a diffuse, lighter band between

the end of the aperture and the posterior parietal tooth.

Aperture ovate; inner lip equidistantly tridentate; one

columellar tooth, obli(]ue, not reaching the border of the

aperture lip, two parietal teeth, the anterior strongest

and perpendicular to the columellar axis, the posterior

about as strong as columellar tooth; outer lip sharp, rarely

with an elongated thickening opposite the anterior pa-

A. M. de Frias Martins, 1995

Page 9

Figures 21-29. Protoconchs of Allochroa species 21. Allochroa bronnii (Philippi), Hawaii, USNM 98175. 22-27. A. layardi (H.

& A. Adams); 22. Hawaii, LISNM 20797; 23. paralectotype, Ceylon, BMNH 196980; 24. Henderson Island, Pitcairn Islands (R.

C. Preece); 25. Lugbon Island, Philippines, USNM 201147; 26. Riu Kiu, Japan, USNM 345877; 27. Mataira Island, Raroia, Tuamotu

Islands, USNM 698771 (specimen shown in fig. 14). 28. A. nana Martins, n. sp., Eninman Island, Bikini, Marshall Islands, lectotype,

USNM 585963. 29. A. tenuis Martins, n. sp., Noumea, New Caledonia, lectotype, USNM 724347. Scale bar = 100 Mm.

rietal tooth and weakly projecting inside the aperture

(Figs. 19-20). Protoconch 385 x 240 ^m, bulbous, light-

brown, showing 1.3 whorls, with somewhat reflected lip

creating an umbilical slit (Fig. 29).

Radula: (Figs. 36-38) Formula [17 + 14 + 1 + 14 + 17]

X 100 + . Central tooth barely above the plane of the

lateral teeth; base deeply emarginate, with lateral pro-

jections Daring out at mid length; crown small, with nar-

row, pointed mesocone. Lateral teeth 14, base elongated

with lateral projections asymmetrical, the inner lower

and serving as "point d'apui" for the articulation with

the outer basal projection of the next tooth in the row

(see Fig. 36); crown subquadrangular, long, with rounded

mesocone. Marginal teeth 17, with one sharp endocone

becoming longer towards the end of the row; mesocone

becoming sharper towards the end of the row.

Type Locality: Lighthouse island opposite Noumea, New

Caledonia. From crevices on undersides of coral slabs,

outer reef.

Material Examined: (Sample size in parenthesis) NEW

CALEDONIA: Lighthouse island opposite Noumea

[USNM 724374 (holotype), 887321 (117 paratypes)]; N

side of N'Go Bay [USNM 725042 (1)]; Touaourou [USNM

724628 (1), 724722 (1)]. PHILIPPINES: Pujada Bay, E

Mindanao [USNM 887322 (1)].

Etymology: tenuis, e = thin.

Range: Known from New Caledonia and Mindanao,

Philippines.

Remarks: Allochroa tenuis is readily distinguished from

all other species in the genus on the basis of its thinner,

darker shell and its large protoconch with flaring lip (see

Fig. 29). The smaller, lighter but equally thin Allochroa

nana, described above, also has a large protoconch, but

without a flaring lip. The shape of the protoconch of A.

tenuis, although unique among the known Allochroa,

strongly resembles that of Ovatella aequalis (Lowe, 1832).

Page 10

THE NAUTILUS, Vol. 109, No. 1

Figure 30. Internal structure of tlie peiiial comple.x of Alluchrua layardi (H. & A. Adams), Henderson Island, Pitcairn Islands

Figures 31-38. Radulae of Allochroa species. 31-32. A. bronnii (Pfiilippi), Hawaii; 31. central tooth and first lateral teeth; 32.

last lateral teeth and marginal teeth 33-35. A. layardi (H & A Adams), Henderson Island, Pitcairn Islands; 33, central tooth and

first lateral teeth; 34. last lateral teeth and marginal teeth; 35, marginal teeth showing basal cusp (arrow ) 36-38. A tenuis Martins,

n. sp,, Noumea, New Caledonia; 36, central tooth (arrow) and lateral teeth; 37. last lateral teeth and marginal teeth in a worn

region; 38 last lateral teeth and marginal teeth in a fresh region. Scale bar = 100 fim for Fig. 30, 10 nm for all others.

The relationships of Allochroa with this European genus

are therefore strengthened.

RELATIONSHIPS WITHIN THE ELLOBIIDAE

Since the studies of Odhner (1925) and Morton (1955)

the subfamilial division of the Ellobiidae has been based

on anatomical characters, mainly on those of the repro-

ductive system. Martins (in press a, b), studying the char-

acteristics of the central nervous system, concurred with

Morton s (1955) conclusions and provided a more com-

plete subfamilial characterization. The Fythiinae, the

most primitive subfamily, have a monaulic, entirely glan-

dular, pallial gonoduct and a wide visceral nerve ring

Figures 39-1-5. ,\natomy of Allochroa layardi (H, & A. Adams), Henderson Island, Pitcairn Islands, 39. mandible; 40. stomach;

41, 42. reproductive system; 43. penial complex; 44, internal structure of penial complex; 45. central nervous system. Scale bar

= 2 mm for Figs. 41-42, I mm for all others.

ad, anterior diverticulmn; agl, albumen gland; amgl, anterior mucus gland; avd, anterior vas deferens; br, bursa, brd, bursa duct;

ca, stomachal caecum; car, cardiac region of stomach; eg, cerebral ganglion; fp, fertilization pouch; hd, hermaphroditic duct, mb,

A. M. de Frias Martins, 1995

Page 11

muscular band; og, osphradial ganglion; ot, ovotestis; pd, posterior diverticulum; pdgl, posterior lobe of digestive gland; pe penis

per, penial retractor muscle; pg, pedal ganglion; pgl, pallial gland, pil, penial pilasters; pig, pleural ganglion; pmgl, posterior mucus

gland; pr, prostate gland/spermiduct; prg, parietal ganglion; psd, posterior vas deferens; pvl, pvloric region of stomach, sv, seminal

vesicle; v, vagina; vg, visceral ganglion

Page 12

THE NAUTILUS, Vol. 109, No. 1

witli a long right parieto-visceral connective. The Ello-

biinae have a diaulic reproductive system with entirely

glandular pallial gonoducts; the visceral nerve ring is

wide and has a short right parieto-viscerai connective.

The Carychiinae have a monaulic reproductive system

with the prostate concentrated distally on the gonoduct

and a central nervous system of the ellobiine type. The

Pedipedinae possess a monaulic or incipient semi-diaulic

pallia! gonoduct where only the pro.ximai half is glan-

dular and a concentrated visceral nerve ring. The Me-

lampodinae are characterized by an advanced semi-

diaulic reproductive system with non-glandular pallial

gonoducts and by concentration of the visceral nerve

ring.

Taking into account the preceding arrangement, Al-

lochroa must remain within the Pythiinae in view of its

monaulic, entirely glandular pallial gonoducts and the

wide visceral nerve ring with a long right parieto-viscerai

connective.

Within the Pythiinae, Allochroa shows strongest affin-

ities with the Mediterranean and Eastern North Atlantic

Ovatella. based primarily on the presence of an internal

lamella inside the outer lip, the protoconch of A. tenuis

which, although different from all others in the genus,

is very similar to that of Ovatella, and the configuration

of the internal structure of the penial complex, where

the penis appears as the development of one of the pi-

lasters (Martins, in press b).

ACKNOWLEDGMENTS

I wish to express my gratitude to Dr. M. G. Harasewych,

of the Division of Mollusks of the Smithsonian Institution,

for his most valuable support. I also want to thank Dr.

R. C. Preece, of Cambridge University, for providing

me with preserved specimens.

This work is integrated into the sabbatical project 1992/

1993 and was partially funded by the Junta Nacional de

Investigagao Cientifica e Tecnologica (JNICT).

LITERATURE CITED

Adams, H and A. Adams. 1855a [1854], Contributions to-

wards the natural history of the Auricuiidae, a family of

pulmoiiiferous Moliusca, with description of many new

species from the Cumiiigian collection. Proceedings of the

Zoological Society of Lontlon 22:30-37.

Adams, H and A Adams. 1855b-1858 The genera of Recent

Mollasca. J. van Voorst, London, 2:93-248, pis. 73-96[1855];

285-412, pis. 97-112|185fij; 413-540, pis. 113-128 [1857];

541-661, pis. 129-1.38(1858].

Ancey, C F. 1887 Nouveiles contributions malacologiques.

IV: Auriculacecs d'Aden (Arable) 15iilletiii de la Societe

Malacologiqiie P'ran^ai.se 4:273-299.

van Bentheni Jutting, W. S. S. 1941. Non-Marine Moliusca

from tlie satellite islands surrounding Java. Archives Neer-

laiidaises de Zoologie 5(3):251-348

Crosse, H. 1894. Faune malacologique terrestre et fluviatilede

la Nouvelle-Caledonie et de ses dependances. Journal de

Conchyliologie 42:161-332; 333-473, pis. 7-10.

Dautzenberg, P. and H. Fischer. 1905. Liste des Mollusques

recoltes par M. le Capitaine de Fregate Blaise au Tonkin,

et description d'especes nouveiles. Journal de Conchyliol-

ogie 53:85-234, pis. 3-6.

Duncan, F. M. 1937. On the dates of publication of the

Society's Proceedings, 1859-1926 Proceedings of the Zoo-

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Dunker, G. 1861. Moliusca Japonica descriptaet tabulistribus

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701 figs.

THE NAUTILUS 109(l):14-37, 1995

Page 14

A remarkable species richness of the Barleeidae

(Gastropoda: Rissoacea) in the Eastern Atlantic

Serge Gofas

Museum National dlli.stoire Naturelle

55 rue Buffoii

F-75005 Paris, France

ABSTRACT

The famiK Barleeidae, hitherto known in the Eastern Atlantic

from tuo European and Northwest African species ol Barlecia

and two species of Trupidurissuia from Sao Tome, is found to

be represented by 18 species, 14 of which (S Barleeia, 3 Pseu-

dodiala, 3 Lirobarleeia) are here described as new. Three new

species share several shell or radular characters with the type

species of Pseudodiala and not with Barleeia For this reason

Pseudodiala is considered to he a valid genus. There are four

centers of diversity; Senegal (3 species), the Cape Verde Islands

(3 species), Sao Tome and Principe ((i species) and Southern

Angola (6 species), together making the Eastern Atlantic the

second most species-rich area after Western America, and the

area with the highest diversity at the genus level.

Ki'y words: Barleeidae, western Africa, new species, species

diversity, biogeography

INTRODUCTION

The Barleeidae is a family of minute prosobranchs that

typically live on intertidal or shallow subtidal hard bot-

toms of temperate to tropical areas. Many species live

within coralline algae, or on "maerl"', deposits of living

calcareous algal concretions. The genera of the Barleei-

dae have been reviewed by Ponder (1983), who then

considered the Anabathriiiae as a subfamily of the Bar-

leeidae but later raised tlieui to full family status (Ponder,

1988).

Western America, from California to Panama and the

Galapagos Islands, is the area with most described species

of Barleeidae: 25 species are recognized as valid b\ Pon-

der (1983). I'he fainily is also represented in the (krib-

bean, eastern South America, East and South Africa,

South-east Asia, Australia and New Zealand, and Japan,

but generally with low (1-5) s()ecies richness.

The family is known hitherto in the Eastern Atlantic

by four species: the mainly European Barleeia unifas-

ciata (Montagu, 1803), a less well known species B. goug-

eti (Michaud, 1830) from NW Africa, and two species

of Tropidorissoia from the Island of Sao Tome, in the

Gulf of Guinea (Tomlin & Shackleford, 1915; Rolan &

Templado, 1994). Several species that may be assigned

to the Barleeidae were described from the island of St.

Helena (Smith, 1890), but the systematic position of most

of them remains uncertain until living animals are ex-

amined.

Recent collecting in Angola by the author, and other

collectors' material from West Africa in Museum Na-

tional d'Histoire Naturelle of Paris and Museo Nacional

de Ciencias Naturales, Madrid, has shown that there are

a number of undescribed species of Barleeidae on the

mainland coast of West Africa, the Cape Verde Islands,

Sao Tome and Principe, making this area one of the

centers of species diversity for the family. No species

assignable to the Anabathridae has been found in this

material, with the exception of the southernmost records

of the European species Nodiilus conturtus (Jeffreys,

1856) in Mauritania.

The material examined has the following origins:

— dredgings and shore collecting bv I. Marche-Mar-

chad, 1953-1956, mostly from Senegal (MNHN).

— intertidal collecting in the (>anarv Islands (1981)

and Mauritania (1983) by P. Bouchet (MNHN).

— the author's collecting in Angola and Sao Tome,

1981-1987, and in Europe (MNHN).

— intertidal and SCUBA collecting in the Cape Verde

Islands bv the 'la expedicion C^ientifica Iberica a Cabo

Verde ", August 1985 (MNCN).

— intertidal and shallow subtidal collecting by E. Ro-

lan in Sao Tome, Principe and the Cape Verde Is. (MNCN

and private collection of E. Rolan, Vigo).

Radulae have been examined using SEM for a number

oi species. Radular morphology has been found infor-

mative for classification at the family and genus level,

but not lor the disirimination of clo.seK related species.

ABBREVIATIONS

AMS = Australian Museimi, S\(lne\

CER = privati' collet'tion ol Emilio Rolan, V igo.

MNHN = Museum National d'Histoire Naturelle,

Paris.

MNCN = Museo Nacional de CUencias Naturales,

Madrid.

UAN = Universidade Agostinho Neto, Luanda,

cat. = catalogue number.

S. Gofas, 1995

Page 15

leg. = "legit ", collected by.

spec, specs. = live-collected specimen(s).

sh. = sheil(s).

jii\ . = juvenile(s).

SEM = Scanning electron micrograph(s).

* denotes specimens in the wet collection.

SYSTEMATICS

Family BARLEEIDAE Gray, 1857

The family is diagnosed by having: a pitted protoconch,

a groove on the sole of the metapodium reaching its

posterior end, and a single-layered operculum with an

inner peg. This last character differentiates the Barleei-

dae from the Anabathridae Pallial and metapodial ten-

tacles are t\picalK absent. The reader is referred to Pon-

der (1983) for a detailed account of the famii\.

Genus Barlecia Clark, 1855

Type species: Turbo ruber Adams, 1797 (= Turbo twi-

fasciatus Montagu, 1803), non Turbo ruber von Salis,

1793, by monotypy.

Shell with smooth, solid teleoconch in all species consid-

ered, non-umbilicate or very narrowly umbilicate, gen-

erally with reddish tinge. Aperture ovate, slightly pro-

socline to orthocline, with smooth lip thickened at some

distance inside, and then beveled to a thin edge. Head-

foot typically pigmented with black and yellow. Oper-

culum with an internal peg, a smooth nucleus on the

columellar side, and fine growth lines on the remainder

of the external surface; a conspicuous external ridge along

the columellar side, overrunning the nucleus.

Radula (Ponder, 1983 and herein. Fig. 71-72): central

tooth with a large rectangular median cusp and 2-3 small

lateral denticles, one sharp basal denticle on each side,

separated by a broad, hardly prominent lamella. Lateral

teeth (one pair) with a rather broad base, terminating

with a large, rather trapezoidal cusp flanked by 2-3

smaller cusps to the inner side and the outer side; with

a U-shaped projection beneath those. Marginal teeth (two

pairs) narrow, hook-shaped, with small, unipectinate cusps

towards the distal end.

Barleeia unifasciata (Montagu, 1803)

(Figures 1, 29-30)

Turhu ruber Adams, 179766, pi. 13 fig, 21-22 (iinii Tinlio niher

von Sails, 179.3). Type locality; Pembrokeshire (Ikitisii Isle.s).

Turbo unifasciatus Montagu, 1803:327-32H. T\pe locality;

Southampton, and Island of Burrow, Devon (British Isles)

Rissoa fulva Michaud, 18.30:12 (repr. 18.32:15). pi. 1. fig.l7-

18. Type locality: not specified; localities mentioned: Agde,

Sete (Mediterranean France) and Corsica.

Sabanaca hinf^hamiana Leach. 1852:1.54. Type localit\ ; Tor-

bay, Devon (British Isles).

Barlecia rubra var. clongata Bucquo) , Dautzenberg & Dollfus,

1884:316, pi 32, fig. 23. Barleeia clongata "nsp." Locard.

1886:272 and 576. Type localit) : Roussillon (Mediterra-

nean France).

Material examined: (collected by the author unless oth-

erwise stated; all in MNHN) Europe: Guernsey, Vazon

Bay, at low tide among corallines, 1000+ specs., 9.1994;

Guernsey, Rocquaine Bay, 1000-h specs., 9.1994. Plou-

manach (Finistere), at low tide on Corallina, 300 specs,

(many juv); Lampaul-Ploudalmezeau (Finistere), on

Corallina, 1000-1- specs, (mostly juv.; adults 2.3 x 1.3

to 2.8 X 1.8 mm); He de Sein (Finistere), 300 specs.;

Guethary (Pyrenees-Atlantiques), 54* spec. (2.9 x 1.7 to

3.3 X 2.0 mm); St. Jean de Luz (Pyrenees Atlantiques),

.30* -t- 150 specs. (2.4 x 1.4 to 3.2 x 1.8 mm); Hendaye

(Pyrenees-Atlantiques), 200 specs.; San Sebastian (Gui-

puzcoa), 1000-t- specs.; Ondarroa (Guipuzcoa), 1000-1-

specs.; Calahonda (Malaga), 200 specs., Los Escullos (Al-

meria), 7 specs. Strait of Gibraltar, Ceuta: Benzu, low

tide, 4 specs. Morocco: Punta Cires, intertidal in algae,

100 specs.; Tangier Ba>, El Ghandouri, 20 specs.; Rabat,

near Lahlou, 12 specs., Mohammedia (=Fedala), 30 sh.;

Near Tarfaya, .35 specs., Ortea leg. 8.1987 (2.5 x 1.5 to

2.7 X 1.7, pattern of foot checked on rehydrated spec-

imen). Mauritania: Bale de I'Etoile, 250 specs. (1.8 x

1.2 to 2.5 X 1.5 mm), Bouchet leg. 1983; Cap Blanc,

exposed rocky shore, 18 specs. (16 juv.), Bouchet leg.

1983; Pointe des Maures, 100-1- specs., Bouchet leg. 1983.

Senegal: Dakar area, 500 sh., Marche-Marchad leg.; Anse

Bernard, 5 specs., Marche-Marchad leg. Tenerife, Ca-

nary Islands: Playa Santo Domingo, 75 specs. (50 juv.),

Bouchet leg. 7.1981; Punta del Hidalgo, 200 specs. (30

juv), Bouchet leg. 7.1981; La Tejita, intertidal, .34 sh.

(10 juv), Bouchet leg. 7.1981; Palm-Mar, intertidal, 25

specs., Bouchet leg. 1981.

Description: Shell conical, solid, adults 2.3 x 1.3 to 3.3

X 2.0 mm. Protoconch dome-shaped (450-500 /u in di-

ameter) of 1 '/2 to 1% whorls, with very minute spiral rows

of pits. Teleoconch of 3'/4 to 3'/2 whorls. Spire whorls

rather flat, body whorl rounded and imperforate. Outer

lip slightly prosocline, beveled inside to a thin edge. Color

of shell reddish brown, paler towards the parietal inser-

tion of the body whorl; plain or with whitish spiral bands,

subsutural, median on body whorl and/or periumbilical;

rarely entirely of a pale, wax-like color. Operculum dark

crimson. Head-foot vsith superficial plain black pattern

on upper part of head, snout, sides of foot and opercular

lobes, upper part of propodium with black area extend-

ing on the edges, and grading to a broad axial white

area; sole entirely colorless. Yellow axial bar on tentacles,

large, triangular yellow granular masses behind each eye.

Habitat: On rocky shores, among red algae (preferably

Corallina) at low water mark or in tidal pools (Southgate,

1982; Borja, 1986a, 1986b, 1987); in shallow photophilous

algal mats in the Mediterranean.

Distribution: From the British Isles to Senegal, the Ca-

nar\ Is. and in the Mediterranean (including the Adriatic

and the Eastern Mediterranean, but not the Black Sea).

Page 16

THE NAUTILUS, Vol. 109, No. 1

Fifiurc I. litirlccia utiifasciata (Montagu, 1803), living animal from Oiita Length of sfiell 2.6 mm. Figure 2-3. Barlccia gougcti

(Midiaud, IS.'iOj. 2. Living animal from Outa. Lt-ngth of .shell -4. 3 mm. 3. Specimen from Dakar, length 5.6 mm.

Remarks: The above description is based on specimens

from Western Europe and Morocco; such typical B. uni-

fasciata are found as far south as Tarfaya, Morocco. The

populations examined from Haie de I'Etoile, Mauritania

and from Senegal are smaller, paler, with more variable

prohle, but are tentatively assigned to B unijasciala

pending further study.

Bellon-Humbert (1974) has figured Barleeia iinifas-

ciata from Morocco as "Barleeia niintiscula Montero-

sato , and the sympatric B. guugeli as "B. rubra" , Mon-

terosato's (1889) description and indication of size (1.5

mm) for B. minuscula, although laconic, fit Coriandria

ftilgida (.Adams, 1797), a common species on the entire

Atlantic coast of Morocco I consider Barleeia inirjuseiila

a syiionvin ot Coriandria julgida, a view further .sup-

ported 1)\ the fact that Monterosato lists both B. rubra

aiul B. gnugeti but no species referable to Coriandria.

Populations from the (]anarv Islands cannot be distin-

S. Gofas, 1995

Page 17

guislied concliologicall\ trom tliose of Western Europe,

and are thus treated as conspecific although it is likely

that very little, if any, genetic exchange occurs. The

head-foot of C^anarian specimens is darker, and the entire

propodiuin is black,

Barleeia gougcti (Michaud, 1830)

(Figures 2-3, 31-32)

Hissoa gougcti MichaiKl, 1830 9, pl.l. fig 7-S T\ [)e l(ic,ilit\

Senegal

Hijdrohia ciwipacta Jeffress, 1883:120, pl.9. tig.9. (non Hij-

drohia compacta Carpt-iiter, 1S64). Type lncalit\ : Taiigiers

(Morocco),

? Barleeia majuscula Montfni.sato, 1884:69, T>pe locality: not

specilied; localities mentioned: Ognina (Sicily), Alger (Al-

geria), Lanipedusa (Sicily channel),

Barleeia gnugeti var, unicnhir Montero.sato. 1889:34 (Casa-

blanca, Morocco)

Barleeia gougcti var, ccrca Monlcrosato, 1889:34 (t^asablanca,

Morocco),

Barleeia carrozzai van Aartsen & Giannuzzi-Savelli, 1991 (re-

placement name for Hydrobia compacta).

Material examined: (collected by the author unless oth-

erwise stated; all in MNHN) Spain: Tarifa, intertidal, 3

specs. (3.5 X 2,1 mm). Strait of Gibraltar, Ceuta: Punta

Almina 35-40 m, 38* -H 12 specs. (4.3 x 2.3 to 5.5 x

2.8 mm), Bouchet leg. 5.1986; Benzii, 8 sh. (white). Mo-

rocco: Temara, low tide, 10 spec; Mouth of Oued Yquem

(= Rose-Marie), S. of Rabat, 31 spec, de Lepinev leg.;

Mohammedia (= Fedala) 50 sh.; El Jadida (= Mazagan),

low tide, 400 specs.; Essaouira (= Mogador), 300 specs.

(2.7 X 1.6 to 6.1 X 3.0 mm); Near Tarfaya, 12 specs.,

Ortea leg. 8.1987. Mauritania: Bale de I'Etoile, 140 specs.

(2.9 X 1.7 to 4.3 X 2.3 mm), Bouchet leg. 1983. Senegal:

Dakar, 8 specs. (5.0 x 3.1 to 5.9 x 3.2 mm), Nickles

leg. 1946; Goree, 7 sh. (pale periumbilical zone), H. Fi-

scher collection; Goree, bale de St. Jean, 50 specs., Delais

leg. 1.1962; Goree 20 m, 10 specs.. Pin leg 1994; Bel-

Air near Dakar, 6-8 m, 59 specs. (3.1 x 1.8 to 5.0 x 2.7

mm, with attached Crepidula), Marche-Marchad leg.;

Dakar, Anse Bernard, 9 specs., Marche-Marchad leg.;

Dakar-ville, 10 specs.. Pin leg. 1994.

Description: Shell conical, solid, adults 2.7 x 1.6 to 6.1

X 3.0 mm. Protoconch dome-shaped (500 n in diameter),

of l'/2 to 1% whorls, with very minute spiral rows of pits,

Teleoconch of 4 to 5 whorls. Spire whorls rather flat,

profile of body whorl very faintly carinate and imper-

forate. Outer lip strongly prosocline near its parietal in-

sertion, then nearly orthocline. Columella protruding an-

teriorly, more pronounced in juveniles, onl\ slightK in

adults v\ here the aperture is more rounded. C^olor of shell

reddish brown, paler towards the parietal insertion of

the body whorl; plain or with periumbilical whitish spiral

zone; sometimes entirely of a pale, wax-like color. Oper-

culum dark crimson. Head-foot with superficial plain

black pattern on upper part of head, snout, and opercular

lobes; upper part of propodium and sole entirely color-

less. Yellow axial bar on tentacles, large, yellow granular

masses behind each eve and (most often) in each oper-

cular lobe anteriorly to the black markings.

Habitat: On hard bottoms with algal concretions, low

intertidal to 40 m.

Distribution: From the Strait of Gibraltar to Senegal.

Remarks; This large species occurs sympatrically with

B unifasciata all along the coast of NW Africa, from

the Strait of Gibraltar .southwards. Barleeia gougcti is

usualK subtidal and is restricted to the lowermost part

of the tidal zone, whereas B. unifasciata is essentially an

intertidal species. The relative abundance of both species

is variable, with B. gougeti the most abundant species

south of Rabat in Morocco.

The ranges of sizes in the two species overlap, and the

boundary cannot be clearly placed with onl\ concholog-

ical characters. The most obvious distinctive character is

the pigmentation pattern of the propodium, which has

lateral black areas in B. unifasciata (a non-variable fea-

ture in Brittany and the Basque country where 6. uni-

fasciata occurs alone), and is white in B. gougeti. The

columella of B. gougeti is more protruding than in B.

rubra. Populations where an entireK pale shell is the

dominant pattern are common in B. gougeti.

Barleeia verdcnsis Gofas, new species

(Figures 4, 33-34)

Type material: Holotype and 120 paratypes (MNCN cat.

15.05/20526); 120 paratypes (MNHN) collected alive

from the type locality.

Type locality: Salmanza, Sao Vicente, Cape N'erde Is-

lands, (16°54'N,24°57'W).

Material examined: (collected by E. Rolan except spec-

imens with date 8.1985 collected by "la expedicion ib-

erica , or where otherwise stated) Cape Verde Islands,

Boavista: Derrubado, 13 juv. sh. (CER); Rife de Chaves

4 m, 4 sh.+ 25* specs. 5.1988 (CER); Baia Teodora 2

m,32 + 12*specs. 4. 1988 (CER); Morro da Areia, 5.1986

(MNCN); Sal Rei, 40 sh., 5.1986 (MNCN) and 38 sh.

(CER). Brava: Furna, 8 sh. (CER). Sal: Mordeira, 3 specs.,

5.1986 (MNCN) and 20 sh. (CER); Fontona, 125 specs.

(mostly juv), 8. 1985 (MNCN); Curral do Dado, 14 specs.

(10 juv.) (MNCN); Joaquin Petinha, 157 specs., 8.1985

(MNCN); Parda, 1 spec. (MNCN); Palhona, 40 sh. + 4*

specs., 5.1987 (CER); Sal (without further precision), 44

sh. (CER); Serra Negra, 4* specs., 5. 1987 (CER); Calheta

Fonda 2-5 m, 15* specs., 5.1988 (CER); Palmeira, 2 +

65* specs, 8.1985 (CER); Rigona, 7 specs., 8.1985

(MNCN). Sao Nicolau: Tarrafal, 3 sh., 8.1985 (MNCN).

Santiago: Tarrafal, 38 sh. (CER). Sao Vicente: Salmanza,

intertidal, 10 specs., Cosel leg. 12.1978 (MNHN); the

type material (2.7 x 1.6 to 3.3 x 1.8 mm) and 100-t-

specs., 8.1985 (MNCN); 12 specs., 5.1987 (CER); Calhau,

4 sh. (CER).

Description: Shell conical with rather tall spire, very

solid, adults 2.7 x 1.6 to 3.3 x 1.8 mm (holotype 3.2 x

Page 18

THE NAUTILUS, Vol. 109, No. 1

8

Figure 4. liarlceia verdensis Gofas, new .species, liolotype from Salmanza (Sao Vicente, Cape Verde I,s.), length .3 2 mm Figure

5. Barleeia aemilii Gofas, new species, liolotype from .Saragasa (.Sao Vicente, C:ape Verde Is.), length 1.7 mm Figure 6. Barlccta

chefiae Gofas, new species, liolotype from Hife de Chaves (Sal, C:ape Verde Is.), length l.;5 mm Figures 7-8. Barleeia lumensis

Gofas, new species. 7. Holotype from I'raia das Conchas (Sao Tome), length 2.1 mm 8. Living specimen from Praia das Con-

chas (Sao Tome), length 2 1 mm. Figure 9. Barleeia laeniolata (Jofas, new species, iiolot\pe from Santa Ana (Sao Tome), length

1.6 mm.

S. Gofas, 1995

Page 19

1.8 mm). Protoconch dome-shaped (.'350-40U n in di-

ameter), ol 1 '/i whorls, with very minute pits arranged

in crowded spiral rows. Teleoconch with 4 whorls. Spire

whorls rather Hat, body whorl rounded, imperlorate.

Outer lip slightly prosocline, beveled both outside and

inside to a thin edge; parietal callus narrow. Color ot

shell dark reddish brown, rather uniform except for a

paler area towards the parietal insertion of the body

whorl and a poorly defined paler bantl around the col-

umellar area. Head-foot black except for the sole, the

sides of the snout, and the tentacles. Yellow axial bars

formed by crowded granules on tentacles; yellow gran-

ular masses on opercular lobes.

Habitat: Intertidal or shallow subtidal, among algae.

Distribution: Cape Verde Islands.

Remarks: This species resembles B. iinifasciata and has

been identified as "B. rubra" in the literature (e.g. (^osel,

1982 and references therein). It is distinguished b\ its

smaller protoconch, with more crowded rows ot pits,

taller spire, and peculiar profile of the bod\ whorl, due

to the fact that the aperture is beveled not only inside

as in other Barleeia, but also outside. It is distinguished

from B. goiigeti by its smaller size and less protruding

columella.

Barleeia aemilii Gofas, new species

(Figures 5, 35-36)

Type material: Holotvpe and 40 paratypes (MNCN cat.

15.05/20527); 40 paratypes (MNHN) collected alive from

the type locality.

Type locality: Saragasa, Sao Vicente, Cape Verde Islands

(I6°49'N, 24°52'W).

Material examined: (collected by E. Rolan, except spec-

imens with date 8.1985 collected by "la expedicion ib-

erica ) Cape Verde Islands, Boavisla: Sal Rei, 2 sh.

(MNCN); Rife de Chaves, 4 m, 1 sh., 5. 1988 (CER).

Santa Luzia: Praia Francisa, 3 sh. (CiER). Sal: (without

precision) 3 sh. (CER); Joaquin Petinha, intertidal, 9

specs., 8.1985 (MNCN) and 1 spec. (CER); Mordeira, 1

sh., 8.1985 (CER ); Serra Negra, 1* spec, 5.1987 (CER);

Palhona, 4 sh. (CER); Palmeira, on wharf, 8.1985, 2 specs.

(MNCN) + 1* spec. (CER); Rigona, 1 spec, 8.1985

(MNCN); Rabo de Junco, 20 sh., (CER). Santiago: S of

Tarrafal, 7 specs. + 16 sh. (CER); Prainha, Praia, 6 specs.

(CER). Sao Vicente: Calhau, 11 sh. (CER); Saragasa, the

type material, 8.1985 (1.4 x 0.85 to 1.75 x 1.0 mm);

Salmanza, intertidal, 8.1985, 1 spec (MNCN) + 30 specs.,

5.1987 (CER) + 4* specs. (CER); Mindelo, Baia Matiota,

1 spec, Cosel leg. 12.1978 (MNHN).

Description: Shell conical, solid, adults 1.4 x 0.85 to 1.8

X 1.0 mm (holotype 1.7 x 0.95 mm). Protoconch dome-

shaped with flattened top (280-300 ^ in diameter), l'/4

whorls, with very minute spiral rows of pits. Teleoconch

of 3 to 3'/4 whorls. Spire whorls rather flat, body whorl

very slightly angulated. Outer lip slightly prosocline, bev-

eled inside to a thin edge; parietal callus narrow, de-

tached over a tiny umbilical chink. Color of shell pale

orange brown, paler towards the parietal insertion of the

body whorl; with a sharply defined paler band along the

angle of the body whorl, and a less clear pale band around

the umbilical chink. Parietal callus tinged with brown.

Shell of some individuals entireK of a pale, orange-brown

color. Periostracum dull. Operculum reddish brown, more

intense along the ridge and edges. Head-foot pale, with

yellow axial bars on tentacles; greenish, iridescent buccal

mass visible in the head b\ transparency.

Habitat: Intertidal or shallow subtidal, among algae.

Distribution: C-ape Verde Islands.

Remarks: This species is readily distinguished from the

previous one by its smaller size, banded color pattern

and more pupoid profile. It is most closely related to

B.chefiae n.sp , \\ hich is still smaller, has a more rounded

periphery of the body whorl, and has a glossy periostra-

cum rather than the dull, opaque texture found in B.

aemilii. This species is dedicated to Dr. Emilio Rolan,

of Vigo (Spain), a distinguished amateur who has been

involved in most of the recent collecting in the Cape

Verde Islands.

Barleeia chefiae Gofas, new species

(Figures 6, 37-38)

Type material: Holotype (live collected) and 2 paratypes

(sh.) from the type locality (MNCN cat. 15.05/20528),

12 parat\ pes (sh.) from Porto P>rreiro (MNCN cat. 15.05/

20529), 24 paratypes (sh.) from Sal Rei (12 MNCN cat.

15.05/20530, 12 MNHN).

Type locality: Rife de Chaves, Boavista, Cape Verde

Islands, in 4 m.

Material examined: (collected by E. Rolan except spec-

imens with date 8.1985 collected by "la expedicion ib-

erica ) Cape Verde Islands, Boavista: Derrubado, 2 sh.

(CER); Baia Teodora, 1 sh. (CER); Porto Ferreiro, 4 m,

12 sh. (paratypes); Rife de Chaves, 4 m, the holotype, 2

paratypes (sh) and 6 sh., 5.1988; Sal Rei, 24 sh. (para-

types), 8.1985. Brava: (no further details), 39 sh. (CER);

Furna, 8 specs. (CER). Santa Luzia: Praia Francisa 1 m,

4 sh. (CER ). Santiago: Tarrafal, 2 sh. (CER). Sal: Pal-

hona, 1* spec (CER). Sao Vicente: Saragasa, 2 specs.,

8.1985 (MNCN); Pedrinha 2 sh. (CER).

Description: Shell pupoid, moderately solid, translucent,

adults 1.1 X 0.7 to 1.3 x 0.75 mm (holotype 1.3 x 0.75

mm). Protoconch dome-shaped with flattened top (260-

270 n in diameter), I'A whorls, with very minute spiral

rows of pits. Teleoconch of 3 to 3Vi whorls. Spire whorls

rather flat, body whorl rounded. Outer lip very slightly

prosocline, beveled inside to a thin edge; parietal callus

narrow, detached over a tiny umbilical chink. Color of

shell brownish, with two sharply defined paler bands

along the periphery of the body whorl, and around the

Page 20

THE NAUTILUS, Vol. 109, No. 1

umbilical chink, and a broad, ill-defined pale area run-

ning along tlie v\ liorls almost from suture to suture; a

dark subsuturai line ("false suture ) corresponds to the

inner surface of the whorl seen by transparency. Parietal

callus and periumbilical area dark. Operculum colorless,

translucent. Animal not observed.

Habitat: Shallow subtidal, among algae.

Distribution: Cape Verde Islands.

Remarks: This species is closely allied to the previous

one, but has a distinctive, Coriandria-like appearance

with its small size, glossy, pupoid shell and distinct band-

ing pattern. It is also distinguished from all other species

treated here by the colorless, translucent operculum. The

correct placement in Barleeia is ascertained by the char-

acters of the protoconch, operculum and radula.

This species is deilicated to Maria Josefa "C.hefi Al-

varez Aza, Dr. Rolan s wife and a companion of the Cape

Verde expeditions.

Barleeia tomensis Gofas, new species

(Figures 7-8, 39-40)

Type material: Holotype (MNHN) and 24 paratvpes (12

MNHN, 12 MNCN cat. 15.05/20531) collected alive from

the type locality.

Type locality: Praia das Conchas, Sao Tome (00°24.7'N,

06°38.0"E), at lou tide.

Material examined: Sao Tome: Esprainha, 50* + 58

specs., Gofas and Fernandes leg. 11.1985 (MNHN); Praia

das Conchas, the type material, Gofas and Fernandes

leg. 11.1985; Praia das Conchas, 13 sh., Rolan leg. (CER);

Mutamba, 28 specs., Fernandes leg. 12.198b (MNHN);

Mutamba, 53 sh., Rolan leg. (C^ER); Lagoa Azul, 5 sh.,

Rolan leg. (CER); Praia Morro Peixe, 2 specs., Gofas leg.

11.1983 (MNHN); Santa Ana, 2 m, 11 specs., Rolan leg.

(CER).

Description: Shell conical, solid, atlults 1.8 x 1.1 to 2.4

X 1.3 mm (holotvpe 2. 1 x 1.2 mm). Protoconch high

dome-shaped (350 fx in diameter) of l'/2 to 1% whorls,

with very minute spiral rows of pits. Teleoconch with

3'/4 to 3'/2 whorls. Spire whorls moderately convex, body

whorl rounded and imperlorate. Outer lip slightly pro-

socline, beveled inside to a thin edge; parietal callus

rather thin. Color of shell reddish brown, paler towards

the parietal insertion of the body whorl; typically with

two periumbilical spiral bauds on body whorl. Opercu-

lum dark crimson. Head-loot w ith superficial black pat-

tern covering the snout, fading behind the head; on oper-

cular lobes and anteriorly on the sides of the epipodial

folds; whitish/yellow axial bar on tentacles, whitish/yel-

low graiuilar masses behind each eye, on the tip of the

snout, on the sole oi the foot and the propodinm, and

on the opercular lobes anteriorly to the black markings.

tiahltal: Intertidal to shallow subtidal on rock\ shores

witli clear water; in algal growth oi coralline and en-

crusting red algae.

Distribution: Only known from the Islands of Sao Tome

and Principe.

Remarks: This species was cited by Tonilin and Shac-

klefortl (1914) and Fernandes and Rolan (1993) as Bar-

leeia rubra The spotted sole of the foot distinguishes it

from B. gougeti and B. unifaseiata, which always have

a colorless sole. The high spire recalls that of B. verdensis

n. sp., but B. tomensis has a constant, characteristic dou-

ble pale band around the umbilicus, and does not have

the aperture beveled outside.

Barleeia taeniolata Gofas, new species

(Figures 9, 41-42)

Type material: Holotype and 5 paratypes (MNCN cat.

15.05/20532), empty shells from the type locality. 5 para-

types (sh.) from Lagoa Azul (MNHN)

Type locality: Santa Ana, Sao Tome (00°15.5'N, 06°45"W),

2 m.

Material examined: Sao Tome: Santa Ana, 2 m, 6 sh.,

Rolan leg. (holotype and paratypes MNCN); Baia de Ana

Chaves, 2 m, 5 sh., Rolan leg. (CER); Praia das Conchas,

5 m, 1 sh., Rolin leg. (MNHN); Praia das Conchas, 2

sh., Gofas leg. (MNHN); Lagoa Azul, 4 m, 14 sh., Rolan

leg. (paratypes MNHN; CER); Mutamba, 4 m, 21 sh.,

Rolh leg. (C;ER). Principe: Santo Antonio, 6-10 m, 1 sh.,

Rolan leg. (CER).

Description: Shell conical, rather translucent, holotype

1.6 X 1.05 mm. Protoconch high dome-shaped (300 /x

in diameter) of I'A whorls, with very minute spiral rows

of pits. Teleoconch with 3 whorls. Spire whorls convex,

body whorl rounded and imperforate. Outer lip slightK

prosocline, thin; parietal callus rather thin. Color of shell

of a pale horny color, with two brown bands on the spire

whorls and two more on the bocK whorl, one continuing

the suture and one [jeriumbilical; the two central bands

on the body whorl commonly fused to form one broad

band. Operculum and animal unknown.

Habitat: I'nknown, [)resuinabl\ sublittoral in .shallow

water.

Distribution: Only known from the islamls of Sao Tome

and Principe.

Remarks: 'ilie pitted protoconch of this species is char-

acteristic ot Barleeia although the shell resembles a Cor-

iandria at first glance. It is easily separated irom jinenile

B. tomensis by the shorter spire, more convex whorls,

and different color pattern. Barleeia taeniolata always

has a pale area separating the suprasutural and subsuturai

browti bands on the whorls, whereas B tomensis only

has periumbilical pale bands.

Barleeia pervulgata Gofas, new species

(Figures 10-15, 47-52, 71-72)

Type material: ilolotspe (MNHN) and over 2000 para-

types (500, ot which 100 are juveniles, in each of the

S. Gofas, 1995

Page 21

Figures 10-12. Barlecia pervtilgata Gofas, new species. 10. Holotype from Chapeu Armado (Angola), length 2.2 mm. 11. Living

animal from Chapeu Armado (Angola), length of shell 2,7 mm. 12. Egg capsule containing a larva ready to hatch, from Chapeu

Armado (Angola). Scale bar 1 mm. Figures 13-15. Barleeia cf. pervulgata Gofas, new species. 13. Specimen of a "pale" morph

from Santa Maria (Angola), length 2.1 mm. 14. Specimen of a subtidal morph from Santa Maria (Angola), length 2.4 mm. 15.

Living animal of a subtidal morph from Porto Alexandre (Angola), length of shell 1.7 mm.

Page 22

THE NAUTILUS, Vol. 109, No. 1

following: AMS cat. C.305091, MNCN cat. 15.05/20533,

MNHN, UAN) collected alive from the type locality.

Type locality: Chapeu Armado, province of Namibe,

Angola {14°27'S, 12°21'E), intertidal.

Material examined: (collected by the author; all MNHN

e.xcept paratvpes specified above) Angola: (typical form)

Caotinha, 2800+ specs. (1.8 x 1.1 to 2.4 x 1.4 mm);

Limagens, 800 specs., 12.1981 (1.7 x 1.0 to 2.2 x 1.3

mm); Santa Maria, intertidal, 1000 specs., 12.1981 (1.8

X 1.1 to 2.2 X 1.3 mm, man\ pale specs); Santa Maria,

dredged 8-10 m, 200 sh.; Lucira (Bissonga), 37 specs.

(1.7 X 1.0 to 2.1 X 1.3 mm); Baia das Pipas, 300* specs.,

2.1983 ( 2.4 X 1.5 to 2.9 x 1.7 mm); Sao Nicolau, 22*

+ 100* specs., 2.1983 (2.1 x 1.3 to 2.9 x 1.7 mm);

Chapeu Armado, the type material (1.9 x 1.2 to 2.7 x

1.6 mm); Praia das Conchas, 200 specs. (2.2 x 1.4, to

2.8 x 1.7 mm); Praia Amelia, 400 + 18* specs., 2.1983

(1.6 X 1,0 to 2.5 X 1.4 mm, some white specs.), {pale,

banded form) Caotinha, 2 specs. + 8 sh.; Limagens, 130

specs., 12.1981 (1.6 x 1.1 to 2.3 x 1.4 mm); Santa Maria,

0-2 m, 1 sh.; Sta Maria, 8-10 m, 12.1981 on calcareous

algae: 50 specs. + 8 sh. (2.4 x 1.4 to 1.8 x 1.2 mm);

Lucira (Bissonga), 2 specs.; Sao Nicolau, 7* + 25 specs.

(2.1 X 1.2 to 2.2 X 13 mm); Chapeu Armado, 9 specs.

+ 24 sh. (2.0 X 1.3 to 2.7 x 1.6 mm); Praia Amelia, 19

specs. + 1 sh.; Baia dos Tigres, in mussel bed, 12*+ 3000

specs., 8.1985. (more delicate form from deeper water):

Santa Maria, 8-10 m, 12.1981 on calcareous algae, 600

specs.; Baia da Lucira (Osar), 10 m, 10 sh.; Santa Marta,

40 m in shell sand, 100 sh.; Praia Amelia, 40-60 m, 25

sh.; off Porto Alexandre, 5* + 2 specs. (1.7 x 1.1 to 2.2

x 1.3 mm).

Description: Shell conical, very solid, adults 1.7 x 1.0

to 2.9 X 1,7 mm (holotype 2.2 x 1.3 mm). Protoconch

dome-shaped (350 n in diameter) of I'/s to 1% whorls,

with very minute spiral rows of pits, sometimes also with

ill-defined spiral sculpture. Teleoconch with 3'/4 to SVz

whorls. Spire whorls rather flat, body whorl rounded and

hardly unibilicate. Outer lip slightK prosocline, beveled

inside to a thin edge; parietal callus tiiickened in adults,

tending to form a ridge. Color of shell reddish brown,

paler towards the parietal in.sertion of the body whorl;

plain or with whitish spiral bands, subsutural, median

on body whorl and/or periumbilical; rarely entirely oi

a pale, wax-like color. Operculum dark crimson. Head-

foot with superficial black pattern on upper part of head

generally forming a V-shaped |)atlern pointing forwards,

then extending over the snout to the sides and axiulK;

two small black spots on the sides of the propodium,

sometimes also a pair of larger, blurred, black to grayish

spots on the axial part of the propodium; opercular lobes

black. Yellow axial bar on tentacles; yellow granular

masses behind each eye, on each lobe of the tip of the

snout and on opercular lobes anteriorly to the black areas;

sole of foot circled by a line of yellow granules.

Habitat: Intertidal to shallow subtidal on rocky shores

with clear water; in algal mats, mainly oi coralline algae.

Distribution: Known only from the provinces of Ben-

guela and Namibe, Southern Angola.

Remarks: This is by far the commonest Barleeia of

Southern Angola. It somewhat resembles the European

species B iinifasciata but is readily diagnosed by the

yellow patterns on the foot, absent in the former.

The egg capsule (Fig. 12) has been observed in Chapeu

Armado, and is very similar to that of B. unifasciata

(Lebour, 1934; Southgate, 1982). It is spherical, contains

a single developing larva, and is attached to the algal

mat and embedded sand grains.

The above description (and Figs 10-11) is based on

the populations commonly found in the intertidal zone

from Caotinha to Praia Amelia. Besides this, there are

two forms which may be distinguished morphologically

and are sympatric in most localities:

(1) One form has a paler, commonly banded shell,

with slightly more convex whorls and more distinct um-

bilical chink (Fig. 13). Such specimens also commonK

have a taller protoconch, with more conspicuous spiral

pattern. They are found on the shore at slightly lower

levels than typical B. pervtilgala, which is restricted to

the shore. These forms look quite different at first glance,

but many individuals cannot be conviiicingK separated

within microsympatric populations; the sample from

Limagens (intertidal) contains many banded forms of

ambiguous morphology (e.g. with banded pattern but

not with convex whorls). Also, the color pattern of the

animal, usualK a useful character at species level, varies

within the same limits in both forms.

(2) A second form with a more delicate, taller, plain

brown shell, occurring always subtidally on soft bottoms

with calcareous algal concretions (Figs 14-15). This form

never occurs microsympatrically with typical B. per-

vulgata, but is found in Santa Maria together with the

above "paler" form in 5-10 m, where both lorms cannot

be convincingK separated. The color pattern of the an-

imal is not very different from that of palest intertidal

morphs.

More study (e.g. using allozyme electrophoresis) may

be necessary to determine whether the differences are

merely ecophenotypic variation, or if there are more than

one species with overlapping variation and/or limited

introgression.

Barleeia ciugiilnidcs CJolas, new species

(Figures 16-17, 43-44)

Type material: Holotype (MNHN) and M paratypes (16

of which 4 juv., in each of the following: AMS cat.

C.305092, MNCN cat. 15.05/20534, MNHN, UAN)col-

leited alive Irom the t\pf locality.

Typ<' locality: Praia .Vmelia, province of Namibe, .Vngola

(r5°13'S, 12°07'E), intertidal.

Material examined: (collected by the author; all MNHN

except paratypes specified above) Angola: Baia de Santa

Maria 8-10 m, 5 sh.; Santa Marta, 40 m, 2 sh.; Baia das

Pipas, 53* specs., 2,1983 (2.2 x 1.2 to 2.4 x 1.4 mm);

S. Gofas, 1995

Page 23

Figures 16-17. Barlecia cinguluidc.s Gofas, new species. 16. Holotype from Praia Amelia (Angola), length 2.4 mm. 17. Living

specimen from Praia Amelia (Angola), length of shell 2.4 mm. Figure 18. Barleeia picta Gofas, new species, holotype from Santa

Maria (Angola), and operculum seen froiii outer side; length of shell 1.8 mm.

Praia Amelia, intertidal, the type material (adults 2 2 x

1.3 to 2.5 X 1,4 mm) and 50 specs, (jiiv.).

Description: Shell high conical, quite solid, translucent,

adults 2.2 X 1.2 to 2.5 x 1.4 mm (holotype 2.4 x 1.3

mm). Protoconch high dome-shaped (350 n in diameter)

of l'/2 to 1% whorls, with spiral rows of relatively coarse

pits, some of them irregularly fused along one row. Te-

leoconch with 3'/4 to 3'/2 whorls. Spire whorls rather flat,

body whorl with a faint but sharp keel continuing the

suture, imperforate. Outer lip nearly orthocline except

for the parietal insertion where it is pro.socline, beveled

inside to a thin edge; parietal callus moderately thick-

ened. Color of teleoconch pale buff, with narrow brown

lines, one subsutural, one suprasutural on the spire and

running just above the keel on body whorl, and one

periumbilical. Operculum pale brown, external surface

shagreened with hardly visible growth lines. Head-foot

with superficial black pattern forming two patches an-

teriorly on each side of propodium, two longitudinal

streaks on the snout and patches on opercular lobes. A

solid yellow bar on tentacles, yellow granular masses

behind eyes, small yellow spots on each lobe of the snout,

and on the sides of propodium; sole colorless.

Habitat: Intertidal on rock\ platforms, among rocks cov-

ered with coralline algae and lying on coarse, clean sand.

Distribution: Known only from the provinces of Ben-

guela and Namibe, Southern Angola.

Remarks: This species recalls the European rissoid Cin-

giila cingilliis (Montagu, 1803) in shape, color pattern,

and even the intertidal habitat in sand under stones. It

is readily distinguished from other Barleeidae by its color

pattern; banded forms of 6. pcrvitlgata never have such

narrow lines and lack the peripheral keel. The most close-

ly related species is B. picta n. sp., similar in shell shape

and in having coarse punctures on the protoconch, but

colored with flames instead of spiral lines.

Page 24

THE NAUTILUS, Vol. 109, No. 1

Barleeia picta Gofas, new species

(Figures 18, 45-46)

Type material: Holotvpe (MNHN) and 37 paratypes (9

AMS cat. C.305093,'9 MNCN cat. 15.05/20535, 10

MNHN, 9 LI AN) collected alive from the type locality.

Type locality: Baia de Santa Maria, province of Ben-

guela, Angola (13°35'S, 12°33'E), on calcareous algal bot-

tom in 8-10 m.

Material examined: (collected by the author; all MNHN

except paratypes specified above) Angola: off Ambrizete,

45 m, 7 sh. (2.2 x 1.4 to 2.6 x 1.6 mm, subfossil?); off

Ilha de Luanda, 120 m: 1 sh (subfossil); Santa Maria, the

type material (1.8 x 1.1 to 2.3 x 1.3 mm); Lucira (Santa

Marta),12sh. (1.5 x 0.9tol.7 x 1.0 mm); Lucira (Caesar),

1 sh.; Sao Nicolau, 15 sh (1.8 x 1,0 to 1.9 x 1.1 mm);

Ponta Albina/Baia dos Tigres, 40 m, 12 sh.

Description: Shell conical, translucent, adults 1.5 x 0.9

to 2.3 X 1.3 mm (holotype 1.8 x l.Q mm). Protoconch

dome-shaped (300 n in diameter) of l'/2 to 114 whorls,

with ill-defined, Dat spirals and covered with spiral rows

of rather coarse, pits. Teleoconch with 3'/4 to 4 whorls.

Spire whorls flat, body whorl faintly angulated along a

line continuing the suture, non-umbilicate. Outer lip or-

thocline, with very thin edge, thickened in the adult at

some distance from its edge. Color of shell very pale buft

to pinkish on apical whorls; two last whorls with a series

of very faint subsutural flames, a colorless zone around

the periphery of the body whorl; another series ot taint

flames on periumbilical area, matching the subsutural

ones or merged into a continuous darker periumbilical

band. Operculum dark crimson, along the ridge and

edges, pale yellowish on distal surface; external surface

shagreened with hardly distinct growth lines. Patterns

of animal not observed in life, seen on rehydrated spec-

imens to have black markings on opercular lobes and on

the snout.

Habitat: On bottoms of calcareous red algae ("maerl")

in a few meters depth and clear water.

Distribution: Known only from Angola; live-taken onl\'

in the [irovince of Benguela, Southern Angola.

Remarks: This species shares with B. cinguloidcs the

sculpture of very coarse pits in the protoconch. Shells

found on the continental shelf of Northern Angola are

similar, but larger. They are presumably derived from

Pleistocene deposits which occur in 50-120 m; the ma-

terial in such ,sam|)lcs contains many littoral species, in-

cluding some now restricted to the south

Genus Pseudodiala Ponder, 1967

Type species: Diala acuta (Carpenter, 1864, by original

designation.

Shell with smooth, rather thin teleoconch, very narrov\ ly

umbilicate, generalK whitish; spire whorls rather flat,

bodv whorl with a faint peripheral keel. Aperture ovate.

with a thin, opisthocline outer lip. Operculum as in Bar-

leeia .

Radula (Ponder, 1983 and herein. Fig. 73-74): central

tooth with a large median cusp with blunt tip, and 2-3

small lateral denticles; one sharp basal denticle on each

side, separated by a broad U-shaped projection. Lateral

and marginal teeth as in Barleeia.

Ponder (1983) synonymized Pseudodiala with Bar-

leeia, but P. niso n. sp. shares with the type species of

Pseudodiala, P. acuta, the opisthocline aperture (see Fig.

19) and the narrow central cusp of the radula. The West

African species assigned to Pseudodiala also stand apart

from Barleeia by their habitat on soft bottoms of the

shelf rather than in intertidal or shallow subtidal hard

bottom biota. These are retained as a separate group on

these grounds.

Pseudodiala niso Gofas, new species

(Figures 19, 53-54, 73-74)

Type material: Holotvpe collected alive (MNHN) and

55 paratvpes (15 AMS cat. C.305094, 20 MNCN cat.

15.05/20536, 20 MNHN), empty shells from the type

locality.

Type locality: Senegal, off Goree, 50 m (14°32'N,

17°25.5'W).

Material examined: (all MNHN except paratypes speci-

fied above) Senegal: off Goree, 50 m, the holotype and

55 paratypes (sh.), Marche-Marchad leg.; Cap de Naze,

25 m, from fish stomach, 3 specs. (3.2 x 1.8 to 3.5 x

2.0 mm), Marche-Marchad leg.; Off Saloum, 50 m, 28

sh., Marche-Marchad leg. ;'"Petite cote (i.e. coast be-

tween Dakar and the Saloum estuar\ ), 32 m, 1 spec,

Leung-Tack leg.

Description: Shell conical, translucent, adults 2.7 x 1.5

to 3.5 X 2.0 mm (holotype 2.7 x 1.7 mm). Protoconch

stub-shaped with rounded top (400 ^ in diameter) of l'/2

to 1% whorls, with imperforate surface, ill-defined, flat

spirals and a very definite suprasutural keel. Teleoconch

with 3'/2 to 4'/2 whorls. Spire whorls hardly convex, body

whorl carinate along a line continuing the suture, dis-

tinctk umbilicate Outer lip strongK opisthocline, bev-

eled to a (juite thin edge, parietal callus separating from

the body whorl in adults. Color of shell uniformly whitish

to gravish. Operculum dark crimson, w ith a faint exter-

nal ridge along the cohunellar siile. .Vniinal not observed.

Habitat: On the continental shelf, 30-100 m

Distribution: Known only from Senegal.

Remarks: The imperforate protoconch of this species is

at\pical in the Barleeidae, but this character state is

bridged to the more usual pittetl protoconch by the tiny

punctures of the next species P. aequiiinctialis n. sp. The

iaint suprasutural keel in the protoconch ol P. niso is

liiiinologous to the stronger suprasutural spiral ridge in

/' acquinoctialis. and the overall siniilarit\ in proto-

conchs indicates a close relationship between these spe-

S. Gofas, 1995

Page 25

Figure 19. Pseudududa luso Ciofas, new spccii'S. li(ilot>pe from off Goree (50 m). Senegal, leiigtli 2 7 iiiiii. Figure 20. PscudodUila

aequinoctialis Gofas, new species, holotspe from off Equatorial Cknnea (1.50 ni). lengtli 1.6 nun Figure 21. Pseudodiala corollaria

Gofas, new species, liolot> pe from Santa Ana, Sao Tome, lengtfi 1.7 mm. Figure 22. Tropidormuia taphrudcs Tomlin & Sfiackleford,

1915, specimen from Praia das Conctias, Sao Tome, length 1.7 mm Figure 23. Twpidorissoia secttnda Rolan & Templado, 199-4,

paratype (MNHN) from Praia Emilia, Sao Tome, length 1.1 mm.

Page 26

THE NAUTILUS, Vol. 109, No. 1

cies. Tlie name is allusive to the resemblance with the

eulimid genus Niso.

Pseudodiala acquiiwctialis Gofas, new species

(Figures 20, 55-56)

Type material: The holotype (MNHN).

Type locality: Equatorial Guinea (0r40'N, 09''25'W),

150 m.

Material examined: Equatorial Guinea: the holotype.

Principe: Santo .Antonio, 6-10 m, 21 sh., Rolan leg. (CIER);

Baia das Agulhas, 8 m, 21 sh., Rolan leg. (CER).

Description: Shell conical, translucent, adults 1.6 x 1.0

to 2.4 X l.,3 mm (holotype 1.6 x 0.9 mm). Protoconch

globular with rounded top (300 fi in diameter) of 1 '/2

whorl, with broad spiral ridges, the lowermost of which

forms a suprasutural keel, and tiny punctures evenly

distributed without lineations. Teleoconch with 3'/2

whorls, externally smooth, sometimes with irregular in-

ternal punctures seen by transparency. Early spire whorls

somewhat convex, body w horl carinate along a line con-

tinuing the suture, distinctly umbilicate. Outer lip strong-

ly opisthocline, thin. Color of shell uniformly whitish to

grayish. Animal and opercuhun not observed.

Habitat: Unknown, holotype collected as an empty shell

on the continental shelf (150 m depth), from a geotechni-

cal core for petroleum exploration, additional material

dredged in (i-10 m on soft bottoms around the island of

Principe.

Distribution: Known from Equatorial Guinea and the

island ot Principe.

Remarks: This species is allied to P. niso. with which it

shares the general shape, opisthocline lip (broken on all

specimens but inferred from the growth lines), the keeled

and umbilicate body whorl. It is readily distinguished

from P. niso by its protoconch with distinct spirals and

tiny perforations. It is distinguished from the next species

P. corollaria n. sp. by lacking the brown color on the

parietal side of the aperture, by its larger size, higher

spire, flatter whorls and by the profile of its last whorl,

which is slightly angulose (more conspicuously in im-

mature specimens).

The holotype, collected in a box core, is slightly im-

mature and possibly derived from a Pleistocene deposit,

but is the best preserved specimen. On some specimens

from Principe, some tiny and evenly distributed jjunc-

tures can be seen by transparency inside the spire w horls

This feature may be restricted to some specimens and

could not be .seen on the holotype.

Pseudodiala corollaria (iofas, new species

(Figures 21, 57-.58)

Type material: Holotype (MNCN cat. 15.05/20.536) and

7 paratypes collected alive from the type locality. 3 para-

types (MNHN) collected alive from Praia Mutamba.

Type locality: Santa Ana, SSo Tome (00°15.5'N, 06°45'W),

2 m.

Material examined: (all collected by E. Rolan) Sao Tome:

Santa Ana, 2 m, 8 specs., (holotype and paratypes

MNCN)and 3 sh.(CER); Praia Mutamba, 4 m, 3 specs.,

(paratypes MNHN); Lagoa Azul, 4 m, 5 sh. (CER); Praia

das Conchas, 5 m, 6 sh. (C>ER).

Description: Shell conical, translucent, adults 1.4 x 0.8

to 1.8 X 1.0 mm (holotype 1.7 x 1.0 mm). Protoconch

globular with rounded top (300 n in diameter) of l'/2

whorl, with broad spiral ridges, and tin\ irregular pits,

very loosely arranged along spiral rows. Teleoconch with

3 to 3'/2 whorls, externally smooth, sometimes with ir-

regular internal punctures seen by transparency. Spire

whorls quite convex, body whorl rounded, distinctK um-

bilicate. Outer lip strongly opisthocline, thin. C^olor of

shell whitish to grayish, translucent, with the parietal

edge of the aperture strongly tinged with brown. Pro-

toconch generally more opaque, tinged with \ellowish

or reddish. Operculum dark crimson. Animal not ob-

served; remains of a black mantle seen attached to the

interior of the shell.

Habitat: On soft bottoms of muddy sand in shallow sub-

tidal sites.

Distribution: Known from the Island of Sao Tome.

Remarks: This species differs from the closely related

P. aequinoctialis by not being carinate at any stage, by

its smaller size and more convex whorls, b\ the more

distinctly pitted protoconch, and by the dark columellar

edge of the aperture.

Genus Tropidorissoia Tomlin and Shackleford, 1915

Type species: Tropidorissoia taphrodes Tomlin and

Shackleford, 1915, by monotypy.

Shell with stout, solid teleoconch and strong spiral keels,

very narrowly umbilicate, generally tinged w ith brown.

Aperture ovate, slightly prosocline to orthocline, with

smooth imier lip, thickened at some distance inside and

then beveled to a thin edge. Protoconch with spiral series

of pits, with or without superimposed spiral cords. Head-

foot pigmented with black and/or yellow. Operculum

as in Rarleeia excejit lor pale brown color. Radula (Fig.

75-76) as in Barlecia.

All the species from St. Helena (Rissoa acqua, R. per-

fccta, R. plotia, R. varicifera, R. vaga, R. ivallichi, R.

simulaiis. all of Smith, 1890) classified or tentatively

classilied in Tropidorissoia by Tomlin and Shackleford

(1915) were referred or tentatively referred by Ponder

(1985) to his new ris.soid subgenus Lirocingula. on the

basis of conchological siinilaril\ w ith the South African

t\[)e species /.. winslowae (Bartsch, 1928). Data on the

soft parts, radulae and opercula are at present wanting

for a definitive placement.

S. Gofas, 1995

Page 27

Tropidorissoia taphrodes Tonilin ami Sliacklt'tord, 1915

(Figures 22, 59-60, 75-76)

Trupidurissiiiu laplixnlcs roiiillii and Sliacklt'tord, 191.5 pi. 5,

tie. 5

Type material: llolotype (live collected) in British Mu-

seum (Natural History); 15 paratypes in National Mu-

seum of Wales, Cardiff, cat. 1955.158.1121.

Type localily:Isiand of Sao Tome.

Material examined: Sao Tome: Esprainha, 19* -t- '30

specs., Gofas and Fernandes leg. 11.1985; Praia Mutam-

ba, 9 specs., Fernandes leg. 12.1986; Praia Mutamba, 6

sfi., Rolan leg. (CER); Lagoa Azul, 5 m, 5 sh., Rolan leg.

(CER); Praia das Conchas (Guadalupe), 19* + 40 specs.,

Gofas & Fernandes leg. 11.1985; Praia das Conchas, 11

sh., Rolan leg. (CER); Baia de Ana Chaves, 4 sh., Rolan

leg. (CER); Santa Ana, 3 specs. Rolan leg. (CER). Prin-

cipe: Santo Antonio, 6-10 m, 41 specs. Rolan leg. (C!ER);

Baia das Agulhas, 8 m, 19 specs,, Rolan leg. (C]ER).

Description: Shell conical, solid, adults 0.8 x 0.5 to 1.2

X 0.7 mm. Protoconch dome-shaped (350 p in diameter)

of l'/2 whorl, with strong spiral cords and superimposed

minute spiral rows of pits. Teleoconch of 2 to 2'/i; whorls,

with two strong, rounded keels on the spire whorls and

with a third similar keel on the body whorl; a spiral

thickening around the small umbilical chink. Outer lip

slightly prosocline, thickened deep inside, then beveled

to a thin edge. Parietal callus moderately developed,

detached from the umbilical chink. Color of shell reddish

brown, sometimes pale yellowish. Operculum yellov\,

translucent, with an internal peg. Head-foot with su-

perficial gra\ pattern on the snout and opercular lobes.

Opaque vellowish white flecks a.xially on tentacles, tri-

angular opaque yellowish-white granular masses behind

each eye. Reddish buccal mass clearly seen by transpar-

ency. Foot white, with tiny opaque white flecks on the

sole; metapodium not conspicuously cleft as in Barlccio.

Habitat: Intertidal to shallow subtidal on rocky shores

with clear water; in algal mat, mainly of coralline algae.

Distribution: Known only from the islands of Sao Tome

and Principe

Remarks: Fernandes ami Rolan (1993) noted a differ-

ence in the populations from Principe, these having fre-

quently a pale shell whereas all specimens from Sao

Tome are uniformly reddish-brown. In the material ex-

amined, 36 out of 41 specimens from Santo Antonio and

all 19 specimens from Baia das .Agulhas are pale.

The protoconch has a peculiar sculpture, cjuite differ-

ent from that of most Barleeia, but similar to that of

" Barleeia" congenita Smith, 1890 from St. Helena Island

(Fig. 63-64).

Tropidorissoia secunda Rolan and Templado, 1994

(Figures 23, 61-62)

Tritpulfjrissoiu sccuruhi Holan ami 'rcinplado, 1994: 2.37-242,

liU 1-1

Type material: llolotype (MNCN cat. 15.05/6975);

Paratypes (3 MNHN, 3 British Museum (Natural His-

tory), 3 American Museum of Natural History, 10 CER,

10 private collection of F". Fernandes, Luanda).

Type locality: Praia Emilia, Sao Tome (0O°21.7'N-

06°43.5'E),

Material examined: (collected by E, Rolan) Sao Tome:

The MNHN paratypes; Praia das Conchas, 6 sh. (CER);

Lagoa Azul, 4 m, 25 sh., (CER); Baia de Ana Chaves, 1

sh. (C;ER). Principe: Santo Antonio, 6-10 m: 17 sh. (CER);

Baia das Agulhas, 8 m, 25 sh. (CER).

Description: Shell conical, solid, adults 0.8 x 0.5 to 1.2

X 0 7 mm. Protoconch dome-shaped (220 yu in diameter)

of 1 '/2 whorl, with minute spiral rows of pits. Teleoconch

of 2 to 2'/2 w horls, with a strong median keel on the spire

whorls and with a suprasutural cord which extends on

the bnd\ whorl to form another keel. Outer lip slightly

prosocline, beveled inside to a thin edge. Parietal callus

moderately developed, detached to form an umbilical

chink. Color of shell white to pinkish. Operculum yellow,

translucent. Head-foot with superficial plain black pat-

tern on the snout and opercular lobes. Opaque white

axial bar on tentacles, large, triangular opaque white

granular masses behind each eye. Pink buccal mass vis-

ible by transparency.

Habitat: Subtidal in a few meters depth.

Distribution: Only know n from the islands of Sao Tome

and Principe,

Genus Liroharleeia Ponder, 1985

Type species: Alvania nigrcscens Bartsch & Rehder,

1939 (Alvania galapagensis Bartsch, 1911 sensu Ponder,

1983), pending decision of ICZN (see Hertz, 1994 for a

discussion of the type species).

Shell with elongate, solid teleoconch and spiral sculpture

in all species considered, non-umbilicate or very narrow-

ly umbilicate, generally tinged with brown. Protoconch

with minute pits loosely arranged in a spiral direction in

the species considered herein; with strong spiral ridges

and no pits in the type species. Outer lip opisthocline

like in most Rissoina species. Head-foot pigmented with

black and/or yellow. Operculum as in Barleeia.

Radula (Ponder, 1983 and herein. Fig. 77-78): central

tooth with a large rectangular median cusp, one large

triangular denticle on each side of it, and sometimes one

or two very small denticles flanking these on the sides of

the tooth; one sharp basal denticle on each side, separated

by a broad, prominent lamella. Lateral teeth (one pair)

with a rather broad base, terminating with a large, rather

trapezoidal cusp flanked by slightly smaller cusps to ei-

ther side; with a U-shaped projection beneath those. Mar-

Page 28

THE NAUTILUS, Vol. 109, No. 1

ginal teeth (two pairs) narrow, hook-shaped, with uni-

pectiiiate small cusps towarils the distal end.

I have included the three species described here in

Liroharleeia despite important differences in both pro-

toconch and teleoconch scupture. They are congeneric

with L. albolirata (Carpenter, 1864) and L. kelscyi (Dall

& Bartsch, 1902), two West American species that Ponder

(1983) included in Lirubarleeia notwithstanding the same

differences. The radula of the two species described here-

in agrees fairly well with those illustrated by Ponder

(1983), although lateral outermost denticles are wanting

in the central tooth of L. siiblaevis n. sp. The specimen

figured by Ponder (1983) to illustrate the designated type

species Alvania galapagensis Bartsch, 1911 has been

shown by Hertz (1994) to belong to Alvania nigreacens

Bartsch & Rehder, 1939, and this case of misidentification

should be settled by a decision of ICZN. The actual A.

galapagensis is a rare species from deeper water and

ma\ be a pickworthiid.

The type species of Fictonoha Ponder, 1983 also re-

sembles the species dealt with here, but differs in having

a very different central tooth on the radula.

Liroharleeia data Gofas, new species

(Figures 24, 65-66)

Type material: Holotvpe (MNHN) and 200 paratypes

(50 AMS cat. C.305095, 50 MNCN cat. 15.05/20538, 50

VINHN, 50 UAN) collected alive from the type locality.

Type locality: Baia de Santa Maria, province of Ben-

guela, Angola (13°35'S, 12°33'E), on calcareous algal bot-

tom in 8-10 m.

IVIaterial examined: (collected by the author; all MNHN

except paratypes specified above) Angola: Caotinha, 1

sh.; Baia de Santa Maria, the type material; Baia da

Lucira (Cesar) on calcareous algal bottom 10 m, 1 spec,

and 12 sh.; Santa Marta, dredged 40 m on shell sand, 25

sh.; Chapeu Armado, 1 sh.; Praia Amelia, 40-60 m, 3 sh.

Description: Shell elongate, solid, adults 3.2 x 1.6 to 6.2

X 2.6 nun (holotype 5.1 x 2.3 mm). Protoconcli dome-

shaped (500 n in diameter) with 1 to l'/-i whorl, a[)par-

ently smooth at low magnification, with very minute,

irregular pits, arranged in spiral bands so as to suggest

a faint spiral sculpture. Teleoconch with 5'/4 to 6 whorls,

with spiral sculpture of very uneejual, sharp cords (15-

20 on penultimate whorl). Early spire whorls rather flat,

the later ones moderately convex, body whorl rounded,

hardly umbilicate, with its latest part wider and salient

from the profile. Outer lip opisthocline in adults, thick-

ened at some distance in the last 'A whorl and beveled

inside to the edge; parietal callus moderatcK developed,

detached anteriorly in adults. Color of shell pale brown

to whitish, with very faint, delicate axial flames starting

from the suture; paler towards the termination of the

body whorl; inside ol a()erture whitish to brown, often

with two brown streaks on the lip. Operculum dark crim-

son. Patterns of animal not observed in liie, seen on

rehydrated specimens to have black markings on oper-

cular lobes and on the snout.

Habitat: On bottoms of calcareous red algae ("maerl")

or coarse, clean shell gravel in a few meters depth in

sheltered bays with transparent waters.

Distribution: Only known from Angola.

Remarks: There is some similarity with Rissoa glypta

Smith, 1890 from St. Helena, which is smaller, has weak

axial folds and somewhat carinate whorls. Liroharleeia

data is the largest of the Angolan Liroharleeia, it is

distinguished from both L. pupoides n. sp. and L. siib-

laevis n. sp. by its more acute apical spire angle, and

coarser sculpture; from L. siiblaevis also by the black

markings on the animal and by the protruding lip of the

aperture in the adults.

Liroharleeia sublaevis Gofas, new species

(Figures 25-26, 67-68, 77-78)

Type material: Holotype (MNHN) and 480 paratypes

(120 of which 20 juv. in each of the following: AMS cat.

C.305091, MNCN cat. 15.05/20540, MNHN, UAN) from

the type locality, live collected.

Type locality: Praia das Conchas, province of Namibe,

Angola (15°07.0"S, 12°06.7'E), in large tidal pools.

IVIaterial examined: (collected by the author; all MNHN

except paratypes specified above) Angola: Caotinha, 8

specs.; Limagens, 4 sh.; Baia de Santa Maria, 8-10 m,

500 sh.; Lucira (Cesar) 8-10 m, 150 sh. (2.6 x 1.4 to 4.3

X 2.2 mm); Lucira (Bissonga) intertidal 12* -I- 18 specs.;

Santa Marta, 3 sh.; Sao Nicolau 14* specs. -I- 30 sh.,

2.1983. Clhapeu Armado, 100 sjiecs. (many juv.); Praia

das Conchas, the type material and 8* specs. (2.8 x 1.4

to 4.6 X 2.3 mm); Praia Amelia, 52 specs. (2.5 x 1.4 to

3.6 X 1.6 mm).

Description: Shell elongate, very solid, adults 2.5 x 1.4

to 4.6 X 2.3 mm (holotype 3.0 x 1.5 mm). Protoconch

dome-shaped (400^1 in diameter) of I'j whorls, appar-

entlv smooth at low magnification, with extremely min-

ute, irregular pits, loosely arranged along a spiral direc-

tion. Teleoconch of 3V2 to 4'/2 whorls, with faint, flat spiral

threads, the intervals of which show spiral series ot punc-

tures (only visible under SEM). Whorls moderateK con-

vex, body whorl rounded, slightK constricted, hardly

umbilicate. Outer lip orthocline, thickened at some dis-

tance in the last '4 w horl and beveled inside to the edge;

parietal callus rather thin, (.letached anteriorly. C-olor of

shell whitish or pale orange brown with subsutural area

paler; occasionally w ith two blurred spiral orange brown

bands (jn a whitish background. Operculum dark crim-

son. Head-loot with \ellow axial bars on tentacles, yellow

granular masses behind each eye, on the tip oi the snout,

on each side of the anterior part of the propodium. Pink

buccal mass clearK \ isible by transparency.

Habitat: intertitlal, in pools or flats with rocks embedtled

in clean, coarse sand and algal overgrowth.

S. Gofas, 1995

Page 29

Figure 24. Lirobarleeia data Gofas, new species, holotype from Santa Maria (Angola), length 3.1 mm Figures 25-26. Lirobarleeia

sublaevis Gofas, new species. 25. Holotype from Praia das Conchas (Angola), length 3.0 mm. 26. Living specimen from Caotinha

(Angola), length 3.1 mm

Distribution: Only known from Angola.

Remarks: Lirobarleeia siihlacvis iTia> resemble some very

smooth specimens in the population from Corimba, re-

ferred to L. piipoides n. sp., but may be distinguished

by its lack of black patterns on the head-foot, the absence

of any pigmentation on its opercular lobes, and the reg-

ularly fusiform profile of its shell, w Inch lacks the pro-

truding body w horl.

Lirobarleeia pupoides Gofas, new species

(Figures 27-28, 69-70)

Type material: Holotype (MNHN) and 200 paratypes

(50 AMS cat. C.305096, 50 MNCN cat. 15.05/20539, 50

MNHN, 50 UAN) collected alive from the type locality.

Type locality: Corimba, province of Luanda, Angola

(8°50'S, 13°09'E), on shell gravel bar in 20 m depth.

Material examined: (collected by the author; all MNHN

except paratypes specified above) Angola: Corimba, the

type material (2.2 x 1.1 to 3.8 x 1.5 mm); Corimba,

Praia Etambar in shell sand: 34 shells (2.5 x 1.1 to 3.4

X 1.5). (Lirobarleeia cf. pupoides) Baia dos Tigres, 8*

-I- 31 specs., 8.1985. (2.4 x 1.1 to 2.8 x 1.2 mm).

Page 30

THE NAUTILUS, Vol. 109, No. 1

Figures 27-28. Lirobarleeia pupoides Gofas, new species. 27. Holotype from Oirimba (Angola), length 2.6 mm 28. Living

specimen from C^oriniba (Angola), lengtfi 3.0 mm.

Description: Shell pupoid, elongate, moderately solid,

adults 2.2 X LI to 3.8 x L5 mm (holotype 2.6 x L2

mm). Protoconch dome-shaped (400 n in diameter) of

I'/i whorl, apparently smooth at low magnification, with

extremely minute, crowded, irregular pits, loosely ar-

ranged along a spiral direction. Teleoconch of 3'/2 to AVi

whorls, with flat spiral threads, the intervals of which

show spiral series of punctures (only visible under SEM),

and with very faint axial wrinkles. Whorls flat to mod-

erately convex; body whorl rounded, hardly umbilicate.

Outer lip orthocline, beveled inside to the edge which is

quite rounded in adults; parietal callus moderately de-

veloped, somewhat detached anteriorly and posteriorly.

Color of shell orange brown to pale yellowish, occasion-

ally with two blurred spiral bands inside the aperture.

Operculum dark crimson. Head-foot with superficial

blaik blotches on each side of the snout, occasionally

confluent and extending over the head; black on oper-

cular lobes and anteriorly on the sides of the propodium;

yellow axial bar on tentacles, yellow granular masses

behind each eye, on the tip of the snout, on the anterior

part of the propodium, and on the opercular lobes an-

teriorly to the black markings.

Habitat: In coarse shell sand, shallow subtiilal.

Distribution: Only known from Angola.

Remarks: Very smooth specimens from C-'orimba may

resemble the previous species, L. sublaevis in having a

similar protoconch and teleoconch microsculpture. Li-

robarleeia siihlacvis generally has a thicker and larger

shell, a more fusiform profile without a protruding lip

and lacks any black pattern on the head-foot and the

opercular lobes. Old shells, possibly subfossil, collected

on the beach at C^orimba, are larger (up to 4.3 x IS

mm) and have a coarser spiral sculpture on the teleo-

conch, approaching that of L. elata n. sp. They never-

theless retain the pupoid outline that is typical of this

species. In the extreme South of Angola, in Baia dos

Tigres, there is a local population of Lirobarleeia resem-

bling that of Clorimba both in shell morphology and color

pattern of the head-foot, and also living in ca. 10 m

depth on a coarse shell sand. It is not clear, from the

material available, if it is the same species with a disjunct

range, occurring where adequate substrate exists, or if it

is a sister species derived independently from an ances-

tral stock.

DISCUSSION

At the local level, sympatric species of Barleeidae may

show differentiation in size, or in habitat. The NW Af-

rican species Barleeia unifasciata and B. gougeti may

be microsympatric and have distinct, although overlap-

ping size ranges. The three Cape Verde Islands species

described herein (B. verdensis n. sp., B. aemilii n. sp.,

B chefiae n. sp.) and the two Barleeia from Sao Tome

(B. tomensis n. sp., B. taeniolata n. sp.) also exhibit size

differentiation among microsympatric species.

The svnipatric occurrence of several species (up to

four in Baia de Santa Maria, Angola) is accompanied by

a .segregation in habitat. Among the .Angolan species,

Barleeia pervulgata n. sp. (the typical form) is found

intertidally in algal tufts, B. cinguloides u. sp intertiilally

among stones lying on coarse sand, while 6. pieta n. sp.

occurs subtidally among algal concretions together with

Barleeia cf. pervulgata and Lirobarleeia elata n. sp. The

two Southern species of Lirobarleeia are also .segregated,

L. sublaevis n. sp. occurring together with B. einguloides

or in tidal pools, while L. elata is always subtidal. There

is a preference for soft bottoms among Lirobarleeia and

Fseudoitiala, and for hard bottoms among Barleeia.

Thus lar, the coast of Western Ameriia was known as

S. Gofas, 1995

Page 31

Figures 29-40. Protoconchs of Barleeia general views (scale bars 100 n) and enlarged portions of the surface (scale bars fO m),

the latter all to scale, 29-30. Barleeia umjaseiata (Montagu, 1803), Ploumanac'h (France). 31-32. Barleeia gougeti (Michaud,

1830), Essaouira (Morocco). 33-34. Barleeia verdensis Gofas, new species, paratype, Salmanza (Sao Vicente, Cape Verde Is.). 35-

36. Barleeia aemilii Gofas, new species, Salmanza (Sao Vicente, Cape Verde Is.). 37-38. Barleeia chefiae Gofas, new species,

paratype, Sai-Rei (Boavista, Cape Verde Is.). 39-40. Barleeia tomensis Gofas, new species, paratype, Praia das Conchas (Sao Tome).

Page 32

THE NAUTILUS, Vol. 109, No. 1

Figures 41-52. Protoconchs of Barleeia: general views (scale bars 100 fi) and enlarged portions of the surface (scale bars 10 m).

the latter all to scale. 41-42. Barleeia taeniolata Gofas, new species, paratype, Santa Ana (Sao Tome). 43-44. Barleeia cinguloides

Gofas. new species, paratvpe. Praia Amelia (Angola). 4.5-46. Barleeia picta (lofas, new species, paratype, Santa Maria (Angola).

47-48. Barleeia pcrvnlgata Gofas, new species, (;:aotiiiha (Angola). 49-50. Barleeia pervulgata Gofas, new species, paratype,

Chapeu Armado (Angola). 51-52. Barleeia cf. pervulgata Gofas, new species, Porto Alexandre (Angola).

S. Gofas, 1995

Page 33

Figures 53-64. Protoconchs of Pseudodiaia, Tropidorissoia and "Barleeia". general views (scale bars 100 n) and enlarged portions

of the surface (scale bars 10 n), the latter all to scale. 53-54. Pseudodiaia niso Gofas, new species, off Saloum (Senegal). 55-56.

Pseudodiaia aequinoctialis Gofas, new species, holotype. Equatorial Guinea. 57-58. Pseudodiaia coroUaria Gofas, new species,

paratype, Santa Ana (Sao Tome). 59-60. Tropidorissoia taphrodes Tomlin & Shackleford, 1915, Praia das Conchas (Sao Tome).

61-62. Tropidorissoia secunda Rolan & Templado, 1994, Praia das Conchas (Sao Tome). 63-64. "Barleeia" congenita Smith,

1890, St, Helena.

Page 34

THE NAUTILUS, Vol. 109, No. 1

Figure§ 65-70. Protoconchs of Lirobarleeia general views (scale bars 100 /i) and enlarged portions of the surface (scale bars 10

^l), tfie latter all to same scale. 65-66. Lirobarleeia elata Gofas, new species, paratype, Santa Maria (Angola), 67-68. L. suhlaevis

Gofas, new species, paratype, Praia das Oinchas (Angola). 69-70. L. pupoides Gofas, new species, paratype, Corimba (Angola).

the part of the world with the highest species richness

in the family Barieeidae. Ponder (1983) recognized 13

species of Barleeia (including one Psetidodiala) distrib-

uted between California and Panama, and 12 species of

Lirobarleeia , distributed in the same mainland area but

also in the Galapagos and Clarion islands.

The description of species from West Africa brings

attention to other centers where the family is well rep-

resented (Fig. 79). The inner part of the Gulf of Guinea

hosts si,\ species representing three genera. The absence

of records from Annobon or Fernando Poo islands reflects

the lack of data, rather than a real difference between

islands.

Southern Angola, with six species representing two

genera, is another center of richness. The ne)rllu'rn limit

of five species in Angola coincides with the shift from a

rocky coast with clear, cool waters, to a prevalently sandy

coastline with warmer, turbid waters north of the city

of Benguela. Rocky shores with a well developed cover

of small photophiious algae (including C'orallina spp. )

are well represented in the south, unlike the north where

hard substrates are scarce and where high turbidity of

waters would smother a dense algal cover. The sampling

effort has been well balanced between l)()th areas, and

the lack of records ot Barleeia in northern Angola is not

a sampling artifact. This supports the view that the gaps

between Angola and the Gulf of Guinea islands, and

between the Gulf of Guinea and Senegal, are real. The

distribution of Barleeia in West Africa may thus reflect

the breakdown of a formerly more continuous range.

To the south, the fauna of Namibia remains virtually

unknown, lint some of the Angolan species, or related

ones, may be present there; both Barleeia and Lirobar-

leeia are present in the southernmost Angolan sample,

less than 100 km from the Namibian border. Despite

being a temperate area with rocky shores. South Africa

has only two poorly know n species, Barleeia caffra (Sow-

erby, 1897) and B. stnithi Bartsch, 1915, w hich are def-

initely outside the West African radiation.

The island of St. Helena may host a number of species

of Barieeidae. However, the systematic position of most

of the species described by Smith (1890) remains uncer-

tain. Barleeia eongenita Smith, 1890 lacks spiral sculp-

ture and superficially resembles true Barleeia, but its

protoconch sculpture (Fig. 63-64) is similar to that of

Tropidorissoia taphrodes. The species described by Sniilli

as Rissoa and tentatively referred by Tomlin and Shac-

kleford (1915) to Tropidorissoia may be either rissoids

or barleeids (see under Tropidorissoia).

Considering the Gulf of Guinea and Angola together,

there are fewer species (12 species) than in West America

but these are more diverse at the generic level: Tropi-

S. Gofas, 1995

Page 35

^^

Figures 71-78. Kadulae, general views ot several complete rows (left) and enlarged views of central and lateral teeth (right; all

scale bars 10 )i). 71-72. Barlccia pervulgata Gofas, new species, Chapeu Armado (Angola), 73-74. Pseudodtala niso Gofas, new

species. Cap de Naze (Senegal). 75-76. Tropidorissoia taphrodes Tomlin & Shackleford, 1915. Praia das Conchas (Sao Tome).

77-78. Lirnbarlccia snblacvi.s Gofas, new species, Praia das Conchas (Angola).

Page 36

THE NAUTILUS, Vol. 109, No. 1

Barleeia

3 spp. X

Pseutioduila

Isp

Figure 79. Distrihiitidii ol the Barleeidae in the Eastern At-

lantic: dashed contour, Barleeia, hatched, Tropidorissoia, and

Liriiharlceia: stars, Pseiuhnhala

dorissoia is present in addition to the three genera found

in West America. Tluis, West Africa as a whole is the

area of tfie world with the highest number of genera in

the family. The representation in .Senegal (three species,

two genera) and in the Cape Verde Islands (three species,

all congeneric) is less spectacular

Larval development without a planktonic stage has

been observed only in B. unifasciata and in B. perviil-

gata n. sp., but can be extrapolated to all the other species

with similar protoconch characters. Despite this con-

straint, the insular species are able to disperse between

moderately distant islands. The three Barleeia species

from the Cape Verde were collected on si.x islanils 10 to

100 km apart, and would i)rohal)ly have been foimd

everywhere in the archipelago had all ten islands been

sampled. The two species of Tropidorissoia and Barleeia

tomenais described from Sao Tome were also foimd in

Principe, at a distance of 150 km (Kernaiides and Kolan,

199.3) Conversely, there seems to be an allopatric dis-

tribution for I'seudodiala acqttinuctialis u. sp., foimd in

the inner part of the (Julf of Guinea, and P. corollaria

found in Sao Tome. This implies that an efficient alter-

native to dispersal during pelagic larval development is

used by some barleeids. The most likely scheme is that

the egg capsules of Barleeia, which are very light and

attached to algae in shallow subtidal environments, are

easily torn off the substrate and float away.

ACKNOWLEDGMENTS

The author thanks Jose Templado (MNCN) and Emilio

Rolan (of Vigo, Spain) for providing the Cape Verde

material, Anders Waren (Swedish Museum of Natural

History, Stockholm) for preparing and scanning the rad-

ulae, Philippe Bouchet (MNHN) and anonymous ref-

erees for helpful conunents. The SEM micrographs of

protoconchs were taken in liniversity of Malaga (Spain)

by J. -J. Cuenca and G. Caballero, or in Centre Interu-

niversitaire de Microscopic Electronique (Paris) by Mrs.

D. Guillaumin.

LITERATURE CITED

Bellon-Hunibert, C. 1974. Les Molhisqnes testaces niarins du

Maroc. Premier supplement, Travaux de Ihistitut Scien-

tifique Cherifien, 37:l-l-t4.

Borja, A 1986a. La alimentacion y distribucion del espacio

en tres moluscos gasteropodos: Rissoa parva (da Costa),

Barleeia unijaxiata (Montagu) > Bitliuni reticulatum (da

Costa). C^aliiers de Biologie Marine 27:69-75.

Borja, A. 1986b. Biologia > ecologia de tres moluscos gaster-

opodos intermareales: Risnua parva. Barleeia unijaseiata

y Bittium reliculatum. I Estructura y dinamica de las

poblaciones. (^ahiers de Biologie Marine 27:491-.507.

Borja, A 1987. Biologia y ecologia de tres moluscos gaster-

opodos intermareales; Rissoa parva, Barleeia unifasciata

y Bittium reticulatum. 2. (^recimiento ('ahiers de Biol-

ogie Marine 28:351-360.

Cosel, R. von, 1982. Ergebnissedeulscli-portuguesisclierSam-

melreisen auf den Kap\erdischen In.seln (Repuhlica de

Cabo Verde). Vorlaufige Liste der niarinen Mollusken.

Courier Eorschung-lnstitut Senckenberg 52:15-25

Fernandes, F. and E. Rolan. 1993. Moluscos niarinos de Sao

Tome y Principe: .■\ctualizaci6n bibliografica \ nuevas

aportaciones. Iberus ll(l):31-47.

Hertz, J. 1994. Review of the tvpe species ol Liroharlecia

Ponder, 1983. The Veliger 37(1): 1 10-1 16

Lebour, M,\'. 1934. Ri.ssoid larvae as lood ol the \<iung her-

ring Tile eggs and lar\ae of the Pl\ mouth Rissoidae Jour-

nal ol the Marine Biological ,\ssocialion ol the I nited

Kingdom 19:52.3-.540,

Monterosalo, T, .A ili, 1889, Coquilles marines niarocaines.

Journal de Conchy liologie 37:20-40 and 112-121

I'oiKJei, \\ F 1983 Review ol the tJenera ol the Barleeidae

(Mollusca Castropoda: Ris.soacea). Records of the .\ustra-

lian Museum 35:231-281,

Ponder, W F 1985 .\ re\ iew of the Genera of the Rissoidae

(Mollusca: Mesogaslropoda: Rissoacea) Records of the

.\ustralian Museum suppl,4: 221 p

Pouiler, V\', F, 1988, The Truncali'lloidcan ( =Rissoacean)

radiation — a preliminar\ ph\logcM\ Malacological Re-

view suppi 4:129-164

Rol.'in, F and J TetTipladu 1994 t na nue\a especie del

S. Gofas, 1995

Page 37

genero Tropidorissoia (Mollusca; Gastropoda: Barleeidae)

para la costa oesteafricana Bollettino Malacologico 29:

237-242.

Smith, E. A. 1890, Report on the marine niollnstan fauna of

the island of St Helena. Proceedings of tlie Zoological

Society of London 1890:247-317, pls^ 21-24,

Southgate, T 1982, The biology of Bar/eeia i(?n/ascia/a (Gas-

tropoda: Prosobranchia) in red algal tufts in S,W, Ireland,

Journal of the Marine Biological .Association of the United

Kmgdoni 62:461-468,

Tomlin. J R le B and I, J Shackletord 1914 The marnie

Mollusca ot Sao Thome, I Journal ot tA)ncholog\ 14:239-

276,

Tomlin, J, R, le B, and L J. Shackleford, 1915, The marine

Mollusca of Sao Thome. II Journal of C'onchologv 14:307-

309, pi. 5

THE NAUTILUS 109(l):38-40, 1995

Page 38

Shell Growth and Decollation in Terrestrial Gastropods

Edmund Cillenberger

Natiniiaal Natiuirhistorisfli Museum

P.O Box 9517

NL 2300 RA Leiden, Nctherlaucis

[2ik1 address, see Hovel]

G. David E. Povel

Institute of Evolutionary and

Ecological Sciences

Rijksuniversiteit Leiden

P.O Box 9516

NL 2300 RA Leiden, Netherlands

Vermeij (1993:34) described decollation of snail shells as:

"... resorption of the inner shell wall in the early part of

high-spired snail shells makes the shell so thin in this

region that the apical whorls drop off. The remainder of

the shell is then plugged by an apical wall or pad. ' A

more refined mechanism has been described for the ter-

restrial pulmonale snail Rtimuia decullata (Linne, 1758)

(Pulmonata: Subulinidae), in which siiell wall resorption

occurs only locally, resulting in small areas of the shell

being predisposed to breaking; prior to the actual de-

collation, an internal septum is formed just below the

weakened part of the shell (Hochpikhler & Kothbaucr,

1975). While studying shells belonging to the terrestrial,

mainly Greek, genus Albinaria Vest, 1867 (Gastropoda

Pulmonata: Clausiliidae), using X-rays, a different mode

of decollation was discovered in some species.

X-ray radiographs were made with an "Enraf Nonius

Diffractis 581", using an Agfa-Gevaert Osray 50DW

X-ray film, with an exposure time of 10 seconds at JGniA

and 30 KV. To facilitate comparisons of individual shells

for shell wall thickness, up to c. 50 specimens were pho-

tographed simultaneously on a single negative.

A comparison of conspecific specimens of various sizes

revealed that all juvenile shells, irrespective of the actual

number of whorls, as well as some specimens that ap-

peared to be adults, were equally thin-walled (Figures

1, 2). Shells with relatively thick walls were never ju-

veniles, they always appeared to be fully grown. When

an apical septum was present in such shells, either sep-

arating the apical whorls from the lower ones or sealing

the upper part of the shell in cases where the apical

whorls were missing, such shells had thick walls only

EiRures 1-5. Specimens ot Albinaria spralti (1-4) and .A. nhcli (5), sliovving various stages in shell groutli anil development. 1.

Juvenile shell. 2. Fully-grown thin-walled shell. 3. Undamaged shell 4. .\pieally damaged, secondarily thickened shell. 5. Shell

with the very rare eoniliiiialion of intact, thin, apical ulioris and a seiondariK lliickened wall lielow a septum Scale bar (below

figure 1 ) = 1 mm

E. Gittenberger and G. D. E. Povel, 1995

Page 39

Figures 6-10. 6,7. Pcniena clau>,iliacjoiinis. 6. Fiilly-grown, partly decollated shell 7. Dt-tail of another specimen, with the

protDciinch present, shovvuig an extremely thin area (arrows) above the septum. 8-10. Tahcilia licinrichi. 8. Juvenile 9. Decollated

and 10. non-decollated adult shells (arrow indicating extremely thin area). Scale bars = 2 mm.

below the septum. Above the septum, the shell whorls,

or what remained of them in partially decollated spec-

imens, were always as thin as in conspecific juvenile

specimens.

These observations suggest that, in a first phase, these

snails produce all the shell whorls, one after the other,

of about equal thickness. Once the shell wall has been

formed, there is no additional thickening in this initial

phase of shell growth, which ends with the formation ot

the characteristic body whorl, thought to be indicative

of adult size. In a second phase, either the entire shell is

thickened from within (Figure 3), or an apical septum

is formed and the secondary thickening is restricted to

the whorls below this septum (Figure 4). The latter cat-

egory contains shells with relatively thin-walled apical

whorls, prone to irregular breaking, usually resulting in

conspicuously decollated shells (F"igure 4: slightly decol-

lated).

In Albinaria, septum formation is usually followed by

decollation, as it is in A. rebeli Wagner, 1924. This species

occurs abundantly in a small valley in Crete, where the

snails can easily be observed, hanging on the rock-faces.

Among lumdreds of full-grown, decollateil specimens,

six adult shells with the apical whorls still present were

collected. These were X-rayed. All of them contained an

apical septum, above which the whorls are thinner than

below it (Figure 5). No traces of shell wall resorption

were seen.

Septum formation may also be related to incidental

damage of the apical whorls, as is most probably the case

in A. spratti (Pfeiffer, 1846). Upon X-ray examination,

an apical septum was observed in 6S of 208 shells of this

species. Either decollation or fairly serious damage to

the uppermost whorls was observed in these specimens.

The remaining 140 specimens were undamaged and

lacked a septum. Fully grown specimens of A. spratti

may have a relatively thick shell wall from the body

whorl up to the protoconch (Figure 3).

In both A. rcbcli and A. spratti, septum formation

might be hypothesized to be a mechanism to protect

against desiccation by evaporation. In A. rebeli septum

formation might be either an autonomous development

or triggered by a specific, relatively high permeability

of the uppermost whorls. In A. spratti it might be in-

dicative of repair, resulting from an accidentally dam-

aged apical portion of the shell.

Relatively thin apical whorls, due to partial secondary

shell thickening below a septum, and often resulting in

decollation, also occur in Perricria dausiliaeformis Tap-

parone Canefri, 1878 (Figures 6,7) and Taheitia hein-

richi Gittenberger, 1989 (Figures 8-10), two terrestrial

snail species from New Guinea. They belong to the fam-

ilies Megaspiridae [after Zilch, 1959:376] or Coelociidae

[after Nordsieck, 1986:111] (Pulmonata) and Truncatel-

lidae (Prosobranchia), respectively. In Perrieria dausi-

liaeformis, multiple septa are formed during the lifespan

Page 40

THE NAUTILUS, Vol. 109, No. 1

of an iiulivickial, resulting in repetitive decollation, as in

Ruttiina decoUata. We studied adult, but iicit fully de-

collated specimens of both species, in which shell whorls

were present above the septum. Extremely thin shell

walls were observed just above the septum, suggesting

that local resorption had occurred there prior to septum

formation (Figures 7,10).

In view of the low number of gastropod shells studied

with X-rays, we may expect that the decollation mech-

anism reported here might be more widely spread among

gastropods.

St\ lciiiiiiiatii|)ti(ira), Archi\ liir Mdilu.sl^t'nfciinde 106:119-

121.

Nordsieck, H 1986. The system of tlie St\ lomniatophora (Gas-

tropoda), with special regard to the s\stematic position of

the C:laiisiliidae, II Importance of sliell and distriljution.

Archiv fiir Mollusl<eni<uiide 117:93-116.

V'ermeij, G. J. 1993. A Natural History of Shells. Princeton

University Press, Princeton. 207 pp.

Zilch, \ 1959. Euthyneura. In: Wenz, W. Gastropoda. Hand-

buch der Paliiozoologie 6 (2), pp. 201-400

LITERATURE CITED

Hochpochler, F. and H Kothhauer 197,5. Der Mechanismus

der Dekollation bei Rumina dccoHata (L.) (Gastropoda:

THE NAUTILUS 109(l):41-42, 1995

Page 41

Cannibalism in the Terrestrial Slug Deroceras laeve

Jianming Shen

Department of Zoology

l'iiiversit\ of Tennessee

Kiinxville. TN 37996 USA

Cannibalism occurs in many taxa of animals under di-

verse contexts (Elgar & Crespi, 1992). Whatever the caus-

es of its evolution, cannibalism often has significant in-

fluences on population d\ namics and ma\ incur changes

in developmental, morphological, behavioral, and life

history characteristics of a species (reviewed in Fox, 1975;

Polis, 1981; Elgar & Crespi, 1992). Cannibalism in gas-

tropods was recently reviewed b\ Baur (1992), who clas-

sified it into three categories: "(1) adult and ju\enile

gastropods preying on conspecific adults and juveniles;

(2) adult gastropods pre>ing on conspecific eggs; (3) lar-

vae and newK hatched snails pre\ ing on conspecific eggs'

(Baur, 1992: 103). In that re\iew, Baur described can-

nibalism in the terrestrial slug Deroceras laeve (Miiller,

1774) as adults preying on conspecifics, citing Rollo and

Wellington (1979). However, although describing it as a

very aggressive species, Rollo and Wellington (1979) made

no remark on cannibalism in Deroceras laeve. Anecdotal

notes similarly seem to suggest that cannibalism in Der-

oceras laeve occurs as adults or juveniles cannibalizing

other adults and juveniles. For example, Karlin and Nae-

gele (1960:14) noted that "it was impossible to hold more

than a few specimens of laeve in a rearing cylinder since

they exhibited strong cannibalistic tendencies" and that

"on several occasions, an indi\idual of this species was

actually seen to attack and feed upon a smaller slug of

the same species". Pillard (1985:14) also obser\ed "mem-

bers of Deroceras laeve feeding on conspecifics".

During the past few years, while raising a large num-

ber of Deroceras laeve in the laboratory for other pur-

poses, I observed numerous cases of cannibalism in this

species. In this note, I describe these observations with

respect to Baur's three categories noted above.

I. Adults or juveniles cannibalizing other adults or

juveniles. Although this is the form of cannibalism that

has been described by previous authors as occurring in

Deroceras laeve, I observed it only once when one larger

juvenile (extended body length ^ 17 mm) killed a much

smaller juvenile (extended body length ~ 8 mm) and

subsecjuently consumed most of the victim. There was

no food available in the container. As suggested by Karlin

and Naegele (1960), this form of cannibalism may be

due to overcrowding or an inadequate diet. On the other

hand, I also observed numerous attacks, sometimes re-

petitive attacks, among adults and juveniles held in the

same container, supporting previous observations by Rol-

lo and Wellington (1979) that Deroceras laeve is very

aggressive. However, such aggression rarely led to can-

nibalism since most victims were able to escape the at-

tacks very quickly. While high proportions (as high as

56%) of slugs collected from the field were observed with

severe wounds, I do not interpret these as indications of

cannibalism in natural populations of this species.

2. Adulls (and juveniles) cannibalizing eggs. In captiv-

ity, this typically occurs when the diet is nutritionally

unbalanced or when food is not available. Most slugs

described here were raised in petri dishes with 3-5 mm

of potting soil on the bottom. L nder such rearing con-

ditions, adults or juveniles rarely consumed eggs present,

provided that food (fresh lettuce) was available. Only on

a few ( <5) occasions, was a slug found to have consumed

one of the eggs it just laid w hen plenty of fresh food was

present. On the other hand, when slugs were raised in

petri dishes lined with only wet Kimwipes, 14 of the 16

slugs observed for longer than 3 weeks consumed pro-

vided eggs as well as lettuce. Juveniles that ate both eggs

and lettuce grew much faster than those that ate only

lettuce (unpublished data).

3. Halchlings cannibalizing eggs. This is the most com-

mon form of cannibalism that I observed in Deroceras

laeve. I disco\ered that newK hatched slugs (1-3 weeks

old) of field collected parents often ate their sibling eggs.

I also observed this in the hatchlings of laboratory raised,

isolated slugs. Egg cannibalism by Deroceras laeve hatch-

lings occurred regardless of rearing conditions. The pro-

pensity for cannibalism seems to vary both among pop-

ulations and among individuals from the same popula-

tion. Some hatchlings appeared to prefer eggs to fresh

lettuce. Of 25 hatchlings supplied with both eggs and

lettuce, four consumed eggs first. Others readily con-

sumed both eggs and lettuce. In an experiment where

hatchlings were fed only with eggs, however, 5 of a total

of 23 hatchlings died after 5-14 days without eating an

egg. While it is unclear whether trophic or nurse eggs

exist in Deroceras laeve, cannibalistic hatchlings often

consumed eggs that did not have a developing embryo.

A few hatchlings consumed eggs that had early devel-

oping embry OS. Contaminated or infected eggs were nev-

er consumed.

Cannibalistic hatchlings of Deroceras laeve display a

typical behavioral pattern when consuming eggs. They

first break the outer coat of the egg, then eat their way

into the inside of the egg and gradually eat part or all

of the egg from the inside out. Sometimes a hatchling

moves its whole body inside the egg and eats from the

inside. It typically takes more than one hour for a new

hatchling to consume most of an egg that is about the

same mass or slightly larger than the hatchling. A can-

nibalistic hatchling may abandon an egg at any time.

Page 42

THE NAUTILUS, Vol. 109, No. 1

Table 1. Weight comparison between cannibalistic and non-

cannibalistic hatchlings of Deroceras laeve. Means and standard

deviations (in parentheses) of the weight at hatching and the

weight one week after hatching are presented along with the

difference between the two measurements (i.e.. weekly growth

rate).

Weight at hatching (mg) 1,240(0.269) 1.388(0.247) 0.2130

Weight after a

week (mg) 5.507(2.514) 3.325(1.536) 0.0369

Difference (mg) 4.267(2,344) 1,9.38(1,400) 0,0182

especially if it is disturbed during the beginning or end-

ing stages of egg consumption.

DISCUSSION

Egg cannibalism by hatchlings is the main form of can-

nibalism in Deroceras laeve. Adults and juveniles can-

nibalizing other adults and juveniles or eggs may occur,

but probably only under severe conditions. As described

above, egg cannibalism by hatchlings does not seem to

be induced by abnormal conditions. Therefore, it could

be common in natural populations. In species such as

Deroceras laeve, however, it may be difficult to observe

egg cannibalism by hatchlings in the field. Eggs of Der-

oceras laeve are not only small (mean ~ 3 mg) but also

typically buried in the soil or laid beneath boards and

other objects, and hatchlings usually do not emerge to

the surface immediately after hatching.

Egg cannibalism by hatchlings is also known in several

other terrestrial gastropods (Baur 1992). For example,

newly hatched snails of Arianta arbustoruni (Linne, 1758)

and Helix pomatia (Linne, 1758) feed exclusively on

eggs (Baur, 1987a, 1988). Baur (1992) suggested that the

age specificity of egg cannibalism in gastropods may be

associated with an age-specific occurrence of digestive

enzymes. Caruubalistic hatchlings often gain consider-

able nutritional and energetic benefits. In Arianta ar-

bustorum and Helix pomatia. such benefits are appar-

ently so significant that cannibalistic hatchlings do not

distinguish between sibling and non-sibling eggs (Baur,

1987b, 1990a, 1990b). My observations indicate that can-

nibals in Deroceras laeve enjoy a similar nutritional ad-

vantage: cannibalistic hatchlings were more than 1.6 times

larger than non-cannibalistic ones a week after hatching,

and the growth rate of cannibals was on average 2.2

times the growth rate of non-cannibals. As shown in

Table 1, both of these values are statistically significant

(p<0.05). It also seems that cannibalistic hatchlings of

Deroceras laeve do not distinguish between kin and non-

kin eggs (unpublished data).

Many questions may be asked about cannibalism in

Deroceras laeve. How common is cannibalism in natural

populations of this species? How variable is the propen-

sity of cannibalism both within and among populations?

Does this variation of cannibalistic propensity have a

genetic basis? What are the costs and benefits of can-

nibalism, and what might be its evolutionary conse-

quences? Experimental investigations are needed to an-

swer these questions.

ACKNOWLEDGMENTS

I thank Drs. G. F. McCracken and M. G. Harasewych

as well as two anonymous referees for helpful comments

on the manuscript. I am grateful to Dr. M. L. Pan for

providing laboratory space. Financial support was pro-

vided by the Theodore Roosevelt Memorial Fund of the

American Museum of Natural History and Sigma Xi, the

Scientific Research Society.

LITERATURE CITED

Haur, B. 1987a. Effects of early feeding e.vperience and age

on the cannibalistic propensity ot the land snail .\riunta

arhu.sluruin. Canadian Journal of Zoolog) 65:.3068-.3070.

Baur, B 1987b Can cannil>alistic hatchlings of the land snail

Arianta arlmsturum distinguish between sib ami non-sib

eggs? Behavior 103:259-265.

Baur, B. 1988. Age-specific food preferences in hatchlings of

Helix pnmatia (L), Snail F"arming Research 2:14-19,

Baur, B 199()a Kgg cannibalism in hatchlings of the land

snail Helix puniatia: nutritional adsantage ma> outweigh

lack of kin recognition Malacological He\ iew 23:103-105.

Baur, B 199()b Possible benefits ot egg canniiialism in the

land snail :\rianta arbusturum. Functional Ecologx 4679-

684,

Baur, B, 1992, C^annibalisni in gastropods. In: Elgar, M. .\.

and B. J. Oespi. (eds.). Cannibalism: Ecology and Evo-

lution among Diverse Taxa Oxford I'niversity Press. Ox-

ford. England, p. 102-127.

Elgar, M. A. and B J Crespi (eds ) 1992. Cannibalism: Ecol-

ogy and Evolution among Diverse Taxa Oxford I'niversity

Press, Oxford, England. 361 pp

Fox, L. R, 1975, (Jannibalism m natural [xipulalions ,\nnual

Review of Ecology and Systematics 6:87-106,

Karlin, E. J. and J. A. Naegele. 1960. Biology of the Mollusca

of greenhouses in New York state. Cornell LIniversity Ag-

ricultural Experiment .Station Memoir 372:1-35.

Pillard, D A 1985 Xhicus trail following by the slug Der-

oceras lacvc (Miiller) Journal of the Teimessee .Academy

of Science 60; 13- 15.

Polis, G 1981 The evolution and dynamics of intraspecific

predatit)n. ,\muial Review ol licology and S\'Steniatics 12:

22.5-251,

Rollo, C, D, and W (;, Wellinglon, 1979, Intra- and inter-

specific agonistic behavior among terrestrial slugs (Pul-

monata: St\ lininalophora) Canadian Journal ol Zoology

57:846-855,

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THE NAUTILUS

Volume 109, Ntunhers 2 and 3

October 12, 1995

ISSN 0028-1344

A qiiaiierhj devoted

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THEt^NAUTILUS

CONTENTS

Volume 109, Numbers 2 and 3

October 12, 1995

ISSN 0028-1344

Gary A. Coovert

Holly K. Coovert

Revision of the Subraspecific ClassiBcation of

Marginelliform Gastropods

43

Marine niological Lsiioratory/

WoofJs Hol« Oceannnraphic Instilutian

OCT;-

ij. MA 02543

THE NAUTILUS 109(2 & 3):43-110, 1995

Page 43

Revision of the Supraspecific Classification of

Marginelliform Gastropods

Gary A. Coovert

Holly K. Coovert

Dayton Museum of Natural History

2600 DeWeese Parkway

Davton, Ohio 45414 USA

ABSTRACT

The supraspecific classification of marginelliform gastropods,

formerly treated as the single family Marginellidae, is revised

This revision is based on all available information for all Recent

species plus a representative sampling of the fossil fauna, and

includes all Recent and fossil genus-group names. All published

information is summarized, and additional dissections, concen-

trating on the foregut and radular morphology, are reported,

nearly doubling the number of species known anatomically

Fundamental differences between two groups of marginelli-

form gastropods are discussed. Cystiscids have internal shell

whorls partially resorbed and columellar plications reduced

internally. Their radular morphology differs fundamentally in

having narrower, more numerous rachidian plates, a different

subradular membrane structure, separate odontophoral carti-

lages, and in the plate development on the posterior end They

differ anatomically in tfie possession of a typical neogastropod

radular sac. True marginellids have unmodified internal whorls

and columellar plications continuous internally. Their radulae

are shorter and broader, odontophoral cartilages are fused an-

teriorly, and a marginellid buccal pouch is present. The Oli-

vidae have the same specializations as the cystiscids and these

two groups are considered to have a common ancestry. On this

basis, the Cystiscidae is recognized as a valid family and not

considered to be closely related to the Marginellidae. The an-

cestry of the Marginellidae is discussed and the family is con-

sidered to be related to the Volutidae. The Cystiscidae is divided

into four subfamilies, three of which are described as new Nine

Recent genera, plus one fossil genus, are recognized. Plesio-

cystiscus. new genus, is described on the basis of a triserial

radula and other primitive characters The Marginellidae is

divided into two subfamilies and three tribes, two of which are

new. A total of 22 Recent genera are recognized in this family,

plus nine fossil genera. Serrataginella. new genus, is described

on the basis of its unique radular and shell morphology A

systematic section lists synonyms and type species of each genus,

along with a diagnosis, description, and summaries of the dis-

tribution and fossil record. The phylogeny of both families is

discussed, and keys to Recent genera in both groups are pre-

sented.

Key ivords: Cystiscidae, Marginellidae, marginelliform gastro-

pods, systematics, supraspecific classification, new genera, new

subfamilies, anatomv

INTRODUCTION

The best classifications are thus based on the widest pos-

sible variety of comparative data.

Richard E. Blackwelder (1964;25)

The higher classification of the family MarginelUdae

has long been in a state of confusion. Especially useful

for an overview of the taxonomic history are the papers

of Coan (1965) and Coomans (1976). The early mono-

graphs of G. B. Sowerby II (1846), Reeve (1864-1865),

Weinkauff (1878-1879), and Tryon (1882-1883), as well

as Tomlin's (1917) catalog, treated all members of this

family under the single genus Marginella. Many popular

works still treat all of them under this single genus, while

others use generic names inconsistently or erroneously,

basing them primarily on superficial similarities of the

shell. Coan's (1965) important reclassification was a ma-

jor improvement, but is now outdated by new infor-

mation. The confusion over the classification stems from

the fact that shell characters alone were used in the early

classifications, or were largely used in later classifications.

Although many good conchological characters exist, they

have generally been misinterpreted, or, in some cases,

unrecognized. Information on radulae and external anat-

omy was widely scattered in the literature, and internal

anatomical data, only recently available, is represented

by a very small sample.

The senior author has spent the last 13 years working

toward a revision of the supraspecific classification of the

family Marginellidae on a world-wide basis, studying all

aspects of these animals, including shell characters, ex-

ternal anatomy, radulae, internal anatomy, and obser-

vations of live animals, including egg capsules. During

this process, many new characters were discovered. We

feel enough information has finally been gathered that

a stable supraspecific classification can be presented,

summarizing this research and presenting a complete

classification, including complete synonymies of all Re-

cent and fossil genus-group names, along with an inter-

pretation of the phylogeny of this group.

Page 44

THE NAUTILUS, Vol. 109, Nos. 2 & 3

MATERIALS AND METHODS

Since most species were originally described on the basis

of the shell, this character complex is of primary im-

portance. Because the eventual aim is to completely

monograph this group at the species level, initial work

involved acquiring every original description for each

Recent species named, as well as all significant subse-

quent descriptions. A comprehensive species list was

compiled from the main catalogs of the family (i.e. Red-

field, 1870; Tomlin, 1917; Wagner & Abbott, 1978) and

all issues of The Zoological Record and other standard

nomenclators were consulted, resulting in the publication

of a comprehensive bibliography (Coovert, 1988e). This

body of information primarily dealt with the shell, and

much effort was expended on studying and analyzing

the conchological characters of all Recent species (Coo-

vert, 1988d). The type species of all genus-group taxa

were closely scrutinized (Coovert, 1986a, 1986b, 1986d,

1986e, 1987a, 1987c, 1987d, 19871, 1987h, and 1988b).

A vast amount of information was accumulated, includ-

ing the original descriptions of every nominate species,

most subsequent descriptions, a large number of pho-

tographs of type specimens and other material, drawings,

tabular sheets recording locality data and shell measure-

ments for thousands of specimens, keys to species groups,

graphs of morphometric shell data, comparative charts,

and personal observations and notes. All taxa were pro-

visionally determined to be valid, synonymous, or un-

described, and subsequently assigned to a genus. Further

research refined these synonymies and generic alloca-

tions. From this large database, a comprehensive species

catalog of every valid Recent species was compiled, bas-

ing all generic allocations and synonymies on the work

outlined herein, not on previous assignments. The generic

descriptions, diagnoses, and species counts contained in

the systematic section of this paper are summarized from

this database. A total of 766 valid Recent species, in-

cluding 77 undescribed species, is currently recognized

for this entire group.

Literature on fossil species was not as comprehensively

studied as that on the Recent fauna, but the original

descriptions of all genus-group names were studied and

a clear determination of each type species was made. A

large sampling of additional literature on fossil species

was obtained and studied, representing all regions world-

wide. Specimens and descriptions of fossil species from

the western Atlantic were particulary well represented.

Generic allocations of fo.ssil species are based on our own

work, and do not necessarily agree with previous assign-

ments. Our approach to the fossil fauna is conservative.

Only specimens or literature personally examined were

included in our database. A good representation of the

fossil fauna has been obtained, and is presumed complete

as far as genus-group names are concerned.

During this 13-year period, shell specimens from a

wide variety of .sources, including a large number of

museums and individuals (.see acknowledgements), were

studied, largely concentrating on the Recent fauna.

Shell measurements, including maximum length par-

allel to the axis and maximum width perpendicular to

the axis, were made with a Spencer AO binocular dis-

secting microscope using a calibrated optical reticle for

minute to small shells under 5 mm. For larger shells, an

electronic digital Max-Cal caliper was used with a res-

olution to 0.01 mm. Internal plications were initially

studied by noting internal features on shells with gastro-

pod drill holes or breakage. More detailed studies were

conducted by grinding a hole on the ventral surface of

the shell with a fine metal file. Some shells were trans-

versely sectioned with a fine jeweler's saw, ground to a

smooth surface on a fine-grit whetstone, then polished

on a glass plate with toothpaste as a fine abrasive.

Methods used for the extraction, illustration, and anal-

ysis of radulae are given in complete detail in Coovert

and Coovert (1987) and only summarized here. Most

radulae were extracted without damaging the shell by

soaking and flushing the shell with a solution of 10%

potassium hydroxide (KOH) using a micropipette. After

rinsing with acidic water, radulae were transferred to a

microscope slide and mounted in glycerin jelly. Drawings

were made using an optical reticle grid in a standard

light microscope. Radular data, reported for each genus

in the systematic section, includes a shell length to rad-

ular width ratio plus the Radular Index. The Radular

Index is derived by dividing the number of rachidian

plates by the number of cusps per plate, and requires a

complete radular ribbon, w hich was not always available.

Routine extraction using KOH destroys many details of

the delicate subradular membrane, especially in true

marginellids. For a detailed study of these features, rad-

ulae were removed during dissection without the use of

KOH and studied in glycerin before mounting (see be-

low). Study of the developing posterior end of the radula

was made from previously mounted material. All per-

sonally prepared material, totalling 224 specimens of 97

species, was mounted using glycerin jelly and studied

with a standard microscope. Through the courtesy of

Dean Hewish, SEMs of radulae from 99 specimens of 67

mostly Australian species, with complete locality, shell,

and morphometric data, were made available. A review

and summary of published information is given in Coo-

vert (1989b), with additional radular data presented in

Coovert and Coovert (1990). A combined total of 190

radulate species, plus an additional 30 species known to

lack a radula, have been studied in detail.

The external anatomy from published sources was

summarized in Cloovert (1987g). Many photographs and

drawings, provided by colleagues, were also studied. Live

material was examined using a Spencer AO binocular

dissecting microscope, then carefully drawn in color. In

more remote localities, a Rupen wide-field 16 X hand

lens was used. Photographs were taken w hen possible.

Data on the external anatomy of 183 species is sum-

marized in this paper.

All published data on the internal anatonn was studied

in great detail. (Charts of potentially useful taxonomic

characters of all 13 anatomically known species were

G. A. Coovert and H. K. Coovert, 1995

Page 45

constructed, patterned after those in Harasewych and

Kantor (1991). Using this information as a guide, and

carefully choosing taxa to augment this published data,

dissections were completed on 11 additional species, plus

one previously reported species that lacked sufficient data.

We concentrated on features of the foregut, as these were

determined to be of most value taxonomically . Specimens

were dissected using a Spencer AO binocular dissecting

microscope. Gross dissections were completed on mul-

tiple specimens of a species when available, with a total

of 26 specimens being dissected (Table 1). Detailed, an-

notated drawings were made during all stages of dissec-

tion. Preservation varied among material due to a wide

range of sources and preservatives. Larger animals were

extracted from shells with a fine jet of water after briefly

soaking them in liquid dish soap. Smaller shells were first

measured, then cracked using a small hand vice. Odon-

tophoral cartilages were carefully extracted from the

buccal mass of radulate species, manually cleaned of

adherent tissue, and mounted on microscope slides using

glycerin jelly as a mounting medium. The subradular

membranes of these radulae were cleaned of extraneous

tissue after extraction and studied in glycerin prior to

permanent mounting in glycerin jelly. Potassium hy-

droxide was not used in the preparation of these carti-

lages or radulae at any stage. A list of the dissected

material, with locality data, shell measurements, and oth-

er details, is presented in Table 1.

The following revision is based on all available infor-

mation of every Recent species plus a representative

survey of the fossil fauna. Most importantly, the type

species of all marginelliform genus-group names, both

Recent and fossil, were studied and fully clarified. Gen-

era are based on groups of related species, not just their

type species. A very conservative attitude was taken to-

wards recognition of higher taxa. Shared, derived char-

acters were searched for to unite groups rather than using

minor differences to split groups. The rank of subgenus

is not utilized in the classification presented as this would

require additional research and necessitate formal rec-

ognition of additional taxa. We prefer the useage of

informal "species groups." The proposal of new taxa has

been avoided unless strongly supported.

ABBREVIATIONS AND CONVENTIONS

The following abbreviations and conventions are used in

the svstematic section and text:

=?

?

t

t

aff.

AMNH

AMS

synonymy questioned (species)

synonymy questioned (genus)

(preceding genus), denotes extinct genus

(following species), denotes fossil species

affinis. having affinity with but not identical

with

American Museum of Natural History

Australian Museum, Sydney

ANSP Academy of Natural Sciences, Philadelphia

BM(NH) British Museum of Natural History

DMNH Delaware Museum of Natural History

GAG collection of Gary A Coovert

ICZN International Code of Zoological Nomencla-

ture

M original designation by monotypy

MHNG Museum d'Histoire Naturelle, Geneve

MNHP Museum national d'Histoire naturelle, Paris

NMV National Museum of Victoria

OD original designation

OD (M) original designation as type species, but also

monotypic

OD (T) original designation as type species, but also

tautonomous

SD subsequent designation as type

SD (M) subsequent designation as t\pe species, but

also monotypic

sic to signify exact transcription

T original designation by tautonomy

TS type species

USNM National Museum of Natural History, Smith-

sonian Institution

In the systematic section, the following sequence is

used for each generic synonym:

Genus in original combination, followed by author, year,

page number(s)

TS as cited by original author of genus (or if not originally

cited, as cited by author of SD)

TS in original combination of author of species, if dif-

ferent from above

Senior synonym if applicable

Designation of type species (i.e. M, OD, OD (M), OD

(T), SD, SD (M), T); SD followed by author, year, page

number(s)

In the systematic section, nomina nulla (misspellings)

are not included. Most nomina nuda, unless they have

a direct bearing, are also not included. Citing these is

undesirable because it imparts a false sense of validity.

The comments in Tomlin (1917:242) are applicable.

Throughout the remainder of this paper, the term

marginellid will be used in the strict sense for the family

Marginellidae as defined and restricted in the systematic

section herein. The term cystiscid will be used for the

family Cystiscidae as defined in the systematic section

herein. The term marginelliform will be used in the

broad sense of marginellid, equivalent to the traditional

concept of the family Marginellidae as conceived by

previous authors, and including both marginellids in the

strict sense and cystiscids.

Terminology pertaining to shell morphology, radular

morphology, and anatomy has been used differently and

without standardization by previous authors. Consistency

and uniformity are important enough to warrant careful

selection and concise definition of terms. In addition,

several new terms are introduced and defined.

Page 46

THE NAUTILUS, Vol. 109, Nos. 2 & 3

Tablo 1. Sex, voiu-licr niiiiiher, localit) data, coniment.s. and shell measurements of disseeted material All dissected specimens,

incliidini; shells and slide preparations, lia^c been retained in the collectidii of (;ar\' A. (-oovert.

Shell

Species

Sex

Vouchers

Localit v/C'omments

Length Width

(mm) (mm I

Bullata Imllata (Born, 177S) male

Dt'titintargo elninwola iC<.>u-

rad, 1834)

female

Dentinmrgn cbiirncola (Con- male

rad, 1834)

Hyalina pallida (Linne, 1758) male

Marginella glabella (Linne, lemaie

1758)

Marginella sebaitiani Marche- male

Marchad & Rosso, 1979

Marginella sebastiani female

Persicula interruplolineata male

(Megerle von Miihlfeld, I81(i)

Persicttia intcrrupiulineala

Persicula interrupt vlineata

Persicula interruptolineata

Persicula interruptolineata

Persicula masirana Roth ii Pet-

it, 1972

Prinmnt aff aletes Rolh, 1978 male

.Note 2

Prunum aff. aletes

Prunum aff. aletes

Prunum guttalum (DilKwn

1817)

Prunum gutlatum

Prunum gutlatum

Prunum prunum ((JmeliM,

1791)

letnalc

GAC Acc. #I(>

90

GAC 1177A

GAC 1448

GAC Acc. #4-

94 C

GAC M2548

GAC: M2549A

GAt; M2549t;

GAC 1581 A

(;AC 1 17(IB

(;ac: 1.582

41.8

38.8

39.8

14.7

23.8

21.4

23.4

9,6

Baliia de Todos Santos, Bahia State, Brazil, 5 52.1 27.8

meters depth; ex. E. C". Rios; ex. isopropyl

alcohol; Note 1

Stock Island, on N. side of U.S. Rt. 1, E. end 6.9 3.1

of island, Monroe Co., Florida, under algae

covered rocks in 0.3 m, Sept. 7, 1984, col-

lectors G, A & H, K. Coovert; ex. isopro-

pvl alcohol

Stock Island, on N, side of L'S Rt, 1, E, end — —

of island, Monroe Co,, Elorida, hand-

dredged in sea grass beds in 0,3 m, Nov,

30, 1988, collectors G, A, & H, K, Coovert;

ex, isopropyl alcohol

(no data); ex. Robert Lipe; ex. ethyl alcohol, — —

animal only

"Banco Canario-Saharia,' (apparently a fish-

ing bank between Spanish Sahara and the

C^anary Islands), March 20. 1991; ex Jose

Aguiar Morales; ex. methyl alcohol, subse-

quently softened with trisodium |)hosphate

(same data as above)

(same data as above)

N. end of Playa La Galera, Isia de Margarita,

Venezuela, NE side of ba\ , hand-dredged

on sandy mud with small patches of Turtle

Grass, Oct, 22, 1992, collectors G A, & H

K, Coovert; ex, isopropyl alcohol

(same data as above)

(same data as above)

(same data as above)

(same data as above)

Masirah Island. Oman, shallow water, low — —

tide, collector Donald T Bosch, ex Robert

Lipe; ex. eth> 1 alcohol; animals only

small island just off Tamarindo Beach, S. of 18.8 12.1

Tamarintlo Diria Hotel, CJuanacaste Prov-

ince, Costa Rica, under rocks, low tide,

Oct. 18, 1989; collector Peter Bright; ex.

isopropyl alcohol

(same data as above) 19.4 12.2

(same data as above) 20.4 13.4

•Airport Reef, I'tila Island, Bay Islands. Hon- — —

duras, shallow water under rocks; ex Rob-

ert Lipe, ex, isoprop\ 1 alcohol, animal oiiK

chamiel behind Lagoon Motel, Marathon, 16,5 9,7

Key Vaca, Monroe Co , Florida, under

rocks, snorkeling in 0-1 m, Sept. 5, 1984,

collectors G. A. it H K Coovert; ex. iso-

propyl alcohol

(same data as above); Sept 1, 19S1, late juve- 14.8 8,0

iiile

Mal;i Redonda, Isla de Margarita, Venezuela, 30,7 15.8

shallow sand and mud flats with scattered

Turtle (irass. low tide, (Kt 24, 1992, ex

John Wolfe; ex, isoprop\ 1 alcohnl

G. A. Coovert and H, K. Coovert, 1995

Page 47

Table I . ( 'oiiliiiiifil

Sex

Species

Pri/num rooscvciti (Bartscli ^ male

Rehder, 1939)

Note 3

Pruninu rooscvciti male

Volvarinu aft (ucnii (Kiener, male

1834)

Note 4

Voloariiui aft dvcna lemale

Volvarinu aft menu lemale

Shell

N'ouchers

.ocality /Comments

Lengtli Wkltli

(mm) (mm)

#26- Walkers Clay, Batiamas, snorkeiing at 1 m, — —

grass and algae, night, June 17, 1988, col-

lector Robert Lipe; ex. isopropyl alcohol;

animal only

#2(> (same data as above) 16,9 1 I 0

15 channel behind Lagoon Motel, Marathon, 11.3 5.0

Key Vaca, Monroe Co., Florida, under

rocks, snorkeiing in 0-1 m, Sept. .5, 1984,

collectors G. A & H. K. Coovert; ex iso-

propyl alcohol

G.^C 1170C (same data as above); Sept. 4. 1984 11,3 .5,0

G.'\(: 1170E (same data as above); Sept, 4, 1984 10 1 4 5

GAC: Ace

88A

GAC; Ace

88B

GAC 1172

Note 1: Radula figured in Coovert and Coovert (1990:2, fig. 1).

Note 2: Shells were significantly larger and much more brilliantly and differently colored than the nominate deep-water form (from

3 to 82 m). More research is necessary to establish the status of this taxon.

Note 3: The radula of this species is closest to P. carneum (Storer, 1837), the animals greatly differing in the external coloration.

The radulae are very distinct from the conchologically very similar Prunum amabile (Redfield, 1852), also from the Bahamas,

whose radula was described in Coovert and Coovert (1990:5-6).

Note 4: See comments in Coovert and Coovert (1990:35) regarding the status of the Florida form of this species.

COMPARATIVE ANALYSIS OF SHELL

MORPHOLOGY

Contrary to the opinion of many previous authors, there

are many conchological characters that are very useful

in the classiBcation of margineliiform gastropods. Those

most useful for higher categories are emphasized.

External varix: The external varix or "margin," for

which these shells are named, is one of the more im-

portant shell characters. Other authors have used the

terms "margin," "marginal varix," "varix, "labial varix,"

"outer varix, ' or "varical callus, " not clearly differen-

tiating between a true external varix and a merely in-

ternally thickened lip. The term external varix is herein

used in preference over the above terms as it more clearly

makes this differentiation. A shell with an external varix

has a weakly to distinctly raised margin of callus depos-

ited externally along the labial edge, easily seen by close

dorsal examination of the shell.

In all marginelliforms, the outer lip is at least some-

what strengthened or thickened internally in the adult,

but many genera completely lack an external varix. This

fact has received very little attention in the literature

and most original descriptions fail to mention the pres-

ence or absence of this important character. Shells lack-

ing a varix are even and smooth along the external labial

edge, without a discernable raised margin of callus. The

outer lip in such shells is usually marked by a change in

angle when viewed from the apex and is often a different

color. Presumably, a strengthened lip, either internally,

externally, or both, imparts adaptive value in protecting

the snail from predation.

Most cystiscid genera, including "Cystiscus" (the group

with a triserial radula), Cystiscus s. str., Crithe. Gib-

berula, and Canalispira, completely lack an external

varix. Only the cystiscid genus Persicula varies in this

character, with some species possessing a well-developed

e.xternal varix, whereas others completely lack it. Gran-

ulina and Piignus are the only cystiscid genera that al-

ways possess a strong external varix. Because the very

strong axial costae in Extra resembles varices, the pres-

ence of this character is indeterminate. Based on other

conchological characters, Extra has been placed with

Crithe and Cystiscus, both non-varicose genera.

Marginellid genera, as opposed to cystiscid genera,

nearly always possess a strong external varix. Hyalina, a

notable exception, usually lacks an external varix. The

genera Prunum and Volvarina vary in this character,

some species possessing a strong varix, some a weak varix,

and other species completely lacking it. The remaining

marginellid genera always possess at least a weak external

varix. This usually very strong, distinct varix often con-

tinues around the anterior edge of the siphonal canal. A

few species have an extremely thickened external varix

that is doubled or "duplicate."

Ontogenetic lip development, from initial formative

stages in the juvenile, through subadult, to the fully adult

stage, has seldom been reported in the literature. We

have studied late juvenile and subadult shells for many

years in an attempt to ascertain stages of labial devel-

opment. The external varix is initially formed by an

outward upturning of the thin lip edge in most species

with a varicose adult shell. Callus deposition subsequent-

ly occurs internally and externally, resulting in the thick-

Page 48

THE NAUTILUS. Vol. 109, Nos. 2 & 3

ened, adult varix. We have observed this sequence in the

cystiscids Persicula cingulata (Dillwyn. 1817) and P.

persicula (Linne. 1758), and in the marginellids Erato-

idea margarita (Kiener, 1834), Marginella rosea La-

marck, 1822, and Glabella adansoni (Kiener, 1834). Be-

cause the situation in Prunum and Volvarina is more

complex, we transversely sectioned, polished, and studied

many shells from these genera. Preliminary results in-

dicate that some varicose species of Prunum, but not

Volvarina, form the varix as described above. This can

often be seen in cross-sections, with the initial varix and

subsequent concentric growth rings showing clearly.

Other varicose species of both Prunum and Volvarina

initially curve the thin, fragile, developing lip edge in-

wardly, subsequently adding further callus deposits in-

ternally to strengthen and thicken the edge. Only later

are significant external callus deposits added in those

species with a varicose adult shell. Preliminary results

with shell cross-sections of Prunum and Volvarina in-

dicate a difference in how this occurs. Varicose Prunum

species with an initially incurved lip produce the varix

by adding shell material to both surfaces while extending

the growing edge. This can be seen in cross-section as

very fine concentric growth rings centered around an

initial growing point. Varicose Volvarina species, on the

other hand, produce the varix by callus deposition in-

ternally and externally, without much elongation of the

labial edge and lack obvious concentric growth rings.

The same difference appears to exist between non-var-

icose Prunum and Volvarina species, but the external

varix simply does not form in these taxa. Additional study

of this character is needed before definite conclusions

can be drawn, but preliminary results indicate the ex-

istence of a potentially useful taxonomic character.

Labial Denticles: Many marginelliform shells have an

outer lip with distinct denticles, referred to herein as

labial denticles or denticulate lip. This is preferred over

such terms as "teeth," "crenulations,"or "crenate." Pres-

ence or absence of labial denticles is sometimes of generic

value but is usually only of specific value. Lirae are a

different type of labial structure and are very useful

taxonomically. They are usually erroneously referred to

as "denticles" or "teeth" in the literature, but the dif-

ference is very important. Denticles are small bumps or

tooth-like projections on the inside edge of the outer lip

and generally do not extend into the aperture. Lirae are

thin spiral ridges on the inside of the outer lip oriented

perpendicularly to the lip edge and extending well inside

the aperture. They resemble denticles only at the outer

edge of the lip. The only marginelliforms possessing lirae

(but not in all species) are the cystiscid genera Persicula,

Canalispira, and Gihbertda. These lirate genera usually

lack an external varix.

Columellar Plications: Another extremely important

character complex is the number and formation of col-

umellar plications. The terms "fold," "plica," and "plait,"

used by other authors, are considered inappropriate for

a spirally wound ridge of shell material that is di'posited

upon the columella and is not an actual fold of shell

material. The term columellar plication is thus preferred.

The anteriormost plication of marginelliform shells does

not differ fundamentally from the adjacent plication, and

is thus not differentiated by a separate term, such as

"siphonal fold or "basal told." This anteriormost pli-

cation is herein considered the first plication, thus count-

ing anterior to posterior. In many animal groups, serial

features are likewise counted in a standard anterior to

posterior direction. A few authors, mostly of older works,

numbered plications from posterior to anterior, resulting

in the anteriormost plication being variously numbered.

A survey of marginelliform gastropods reveals that the

first columellar plication borders the anterior edge of the

columella in all cases (usually continuing and merging

with the external varix) except Afrivoluta, which has a

very volute-like columella. Two fundamentally different

plicational conditions occur in marginelliform species.

These are correlated with the type of internal whorls

present.

Internal Whorls: One group has typical neogastropod

internal whorls that are simply previous body whorls

subsequently enclosed by additional whorls without fur-

ther modification. The columellar plications remain un-

modified for their entire length. This type of plicate

internal whorl has also been observed in the families

Turbinellidae, Cancellariidae, Mitridae, Costellariidae,

and Volutidae. The internal whorls in true marginellids

are maintained at their original thickness with no ap-

parent resorption or further modification and are herein

termed unmodified internal whorls (figure 1). Taxa with

unmodified internal whorls have columellar plications

remaining at their original strength and number through-

out subsequent whorls. These plications can be found

intact internally upon the entire length of the columella.

They are here termed continuous columellar plications

and are normally oriented nearly perpendicular to the

shell axis. The usual number of plications in this group

is four, but varies from two to six among different species,

although not normally varying within a species. In those

species with five or six plications, the fifth and sixth

continue internally for at least a hall whorl, and often

are fully continuous. These are also considered contin-

uous plications in this paper. In some species, a weak

"false fifth plication" (or less commonly a "false fourth"

or "false sixth ) is present. This may be either a parietal

lira that does not continue into the aperture for more

than an eighth of a whorl, or a denticle, but neither are

true plications because they are not continuous. Within

marginelliform gastropods, unmodified internal whorls

and continuous columellar plications are found only in

species herein assigned to the Marginellidae s. str.

Another group has extremely thin, largely resorbed

internal whorls, olten partially missing due to complete

local resorption or breakage. Proceeding inward, the first

three (anterior) columellar plications in the aperture are

quickly reduced to two, the third simply diminishes with-

in a half whorl internally. The remaining two are further

reduced to a single, sharp, very oblique columellar edge

G. A. Coovert and H. K. Coovert, 1995

Page 49

Figures 1-4. Internal shell vvhorls. 1. Prunum priinum (Gmelin, 1791), showing unmodified internal whorls. GAC, no data Length

30.9 mm 2. Persicula persicula (Linne, 1758), showing cystiscid internal whorls. GAC Ace. # 54-87, Joal River, Senegal, on sand

flats at mouth of river. Length 20.0 mm. 3. Plesiocystiscus jewettii (Carpenter, 1857), showing cystiscid internal whorls. GAC Ace.

# 14-90, California. Length 5.7 mm. 4. Cranulina hadria (Dall, 1889), showing modified cystiscid internal whorls. GAC 824,

Tampa Bay, Pinellas Co., Fla.. 0.3 — 0.6 m, shallow grass flats. Length 2.0 mm.

within one full internal revolution. This single, sharp

columellar edge essentially becomes the axis of the shell.

The shape of the internal whorls are very different from

the original juvenile body whorls that were subsequently

resorbed after being enclosed by the next whorl. Original

juvenile shell color, patterns, and apertural columellar

plications, generally more than two, are all completely

resorbed. Among marginelliform gastropods, this type of

internal whorl is found only in species here assigned to

the Cystiscidae. They are here termed cystiscid internal

whorls (figures 2, 3). Columellar plications in species

possessing cystiscid internal whorls differ fundamentally

from the continuous plications found in species with un-

modified internal whorls, primarily because they are

quickly reduced to a single columellar edge, and are not

continuous internally. They are here termed internally

reduced columellar plications. Posterior to the three in-

ternally reduced columellar plications are one to ten

parietal lirae superficially resembling columellar plica-

tions. These were not differentiated from columellar pli-

cations by most previous authors. Parietal lirae are some-

what smaller and weaker than the three anterior plica-

tions, do not continue into the aperture for more than a

quarter turn, and gradually diminish posteriorly. This

results in a total of up to 13 "plications," of which only

the anterior three are considered true, but highly mod-

ified, plications. This columellar arrangement, composed

of internally reduced columellar plications plus parietal

lirae, is herein termed multiplicate. This term has been

used by some previous authors, but was not clearly de-

fined. In a few cases, there may be parietal denticles

posterior to the parietal lirae, but these are even shorter.

A very few species of Cystiscus have only two columellar

plications (one species, possibly not a cystiscid, is reported

to have only a single plication).

A modification of the cystiscid internal whorl has been

studied in species herein assigned to the genus Granulina.

This modified type is characterized by the first two col-

umellar plications (proceeding inwardly) abruptly end-

ing within one full revolution internally. At this point, a

thin, axially oriented edge begins much as in species with

cystiscid internal whorls. However, the two posterior col-

umellar plications continue and are situated at the an-

terior edge of the more bulbous portion of the mostly

resorbed internal whorl. These usually translucent inter-

nal whorls are extremely thin and fragile and break very

easily. The anterior end of these whorls appears to be

open. We believe that this is a modification of the cys-

tiscid internal whorl and is herein referred to as a mod-

ified cystiscid internal whorl (figure 4). The four colu-

mellar plications of Granulina superficially resemble

continuous plications, but because the first two quickly

terminate, they are considered to be internally reduced

columellar plications typical of all other cystiscids. The

posterior two continue, and are herein referred to as

pseudo-continuous plications because they apparently do

not continue for more than a whorl or two internally.

Members of the genus Granulina lack the gradually

weakening parietal lirae posterior to the columellar pli-

cations, and thus they do not have a multiplicate colu-

mella typical of other cystiscids.

Apertural Characters: The extent to which the aperture

is occupied by the columellar plications is useful taxo-

nomically. Genera with a multiplicate columella usually

have more than half the apertural length occupied by

the plications plus parietal lirae. Marginella, Glabella.

Dentimargo, and Eratoidea have four strong, continuous

columellar plications occupying more than half the ap-

ertural length, as do Austroginella and many of its rel-

atives. Most remaining typical marginellids have four

plications occupying less than half the apertural length.

Page 50

THE NAUTILUS, Vol. 109, Nos. 2 & 3

In a few genera, though, the pHcations are crowded an-

teriorly. Columellar plications are also very useful ta.x-

onomically at the species level. In addition to the number

of plications, other useful characters include their height,

thickness, shape in cross-section, extent of protrusion from

the aperture, their angle, whether they are fused or sep-

arate, and whether or not they are excavated inside the

aperture. Very little use has been made of these features

in the literature.

Another feature of the aperture is its relative width,

which can often be compared to the lip thickness as a

readily available reference. Most genera have an aper-

ture that is widest anteriorly, but in some the aperture

is broadest medially.

Siphonal Notch: The presence or absence of a siphonal

notch is an important shell character. Called "anterior

notch," "notched anterior canal," or "siphonal canal" by

various authors, the term siphonal notch is preferred, as

it relates to the function of this feature. A distinct si-

phonal notch is present in Persicula and Gibberula, but

absent in the related Canalispira. This notch is a typical

character of Glabella and most Marginella s. str. The

genera Afrivolnta and Marginellona possess a siphonal

notch, although it is weak in Marginellona. Austrogi-

nella has a distinct siphonal notch, but in the related

Mesoginella the notch is very weak to absent (Coovert,

1988b). In all remaining marginelliform genera the notch

is very weak or absent.

Posterior Notch: A weak to distinct posterior notch is

sometimes developed at the junction or commissure be-

tween the body whorl and the posterior end of the lip.

The posterior end of the lip generally forms an abrupt

angle just prior to joining the body whorl, leaving a broad

to narrow notch between the lip and body whorl. Less

commonly, this notch forms as a weak to distinct groove

in the posterior end of the lip itself. Variously termed

"posterior notch," "posterior sinus," "posterior canal,"

"anal canal," or "anal sulcus," posterior notch is pre-

ferred, because it is consistent with "siphonal notch."

The terms "anal canal" or "anal sulcus" imply a func-

tional aspect not borne out anatomically. The presence

or absence of this character is often obscure, reducing

its taxonomic importance. However, many Persicula,

Gibberula, and Canalispira have a distinct posterior notch

formed at the junction of the body whorl and posterior

end of the lip. The jiosterior notch in Persicula is often

accentuated by an adjacent callus deposit on the body

whorl. The deeply channeled, narrow notch in Canal-

ispira is diagnostic. Protoginella has a posterior notch

formed in the posterior corner of the lip itself.

Spire: Spire height is often important taxonomically.

Several groups have a completely immersed spire and

thus an easily recognizable shell sha[)e. In this paper, a

low spire is defined as having a length less than 'A the

shell length (figures 43, 44, 48, 55). A spire of medium

height ranges from V* up to '/i the shell length (figures

57, 73), whereas a tall spire is more than '/i the shell

length (figures 49, 70). Spire height often varies consid-

erably within a genus and is thus more important at the

species level. Spire height is determined by the insertion

point of the posterior end of the lip, and can be refer-

enced by whether or not the posterior end of the lip joins

the body whorl at, above, or below the previous suture.

This is a very useful character within Volvarina, for

example.

Shape: Shell shape is another useful feature, and al-

though somewhat subjective, should be carefully de-

scribed using standard shape names. Marginelliform shells

vary from a nearly round, globose outline to elliptic,

obovate, obconic, conic, biconic, subtriangular, pyriform,

elongate-ovate, subcylindrical, cylindrical, and other

combinations. The shape of the shoulders is very impor-

tant in determining overall shell shape, and can be gently

curved to strongly rounded or carinate.

Size: Shell size is often very valuable as a species-level

character and, in a general way, for higher categories.

Shell size is here reported as minute for shells up to 2.4

mm in length, small for shells over 2.4 mm up to 6.0

mm, medium for shells over 6.0 mm up to 13.0 mm,

moderately large for shells over 13.0 mm up to 25.0 mm,

large for shells over 25.0 and up to 50.0 mm, and very

large for shells over 50.0 mm in length.

Surface Features: Shell texture is another very useful

conchological feature. Although most marginelliform

shells are perfectly smooth and glossy, some have very

characteristic weak to strong a.xial costae. This term is

preferred over "axial ribs" or "longitudinal ribs. This

character is often useful at the genus or the species level.

Pronounced surface texture is very rare in the family,

but Pugnus and a few Granulina have a minutely sculp-

tured surface. Rivon}arginella species have minute pus-

tules ventrally that are part of the callus deposit. Many

other species have callus deposits ranging from a thin

parietal wash to distinct deposits on the spire, posterior

parietal area, or anteroventrally. Ajrivoluta has a very

distinctive callus pad (Coovert, 1987a). The placement

of callus deposits is species specific and thus diagnostic

in the Persicula cornea group (Coovert, 1987b). Many

Granulina. Crithe. and other species have a character-

istic callus deposit here termed a collabral parietal callus

ridge (figures 55, 56). This is a narrow ridge of callus

deposited parallel to the lip and usually lying just outside

the aperture. It often interesects the columellar plica-

tions, causing them to appear excavated. Heavier callus

deposits at vulnerable points are interpreted as a pro-

tective adaptation against predatory boring. Preliminary

observations suggest a strong correlation betw een typical

bore hole locations in uncallused species and locations of

callus deposits in related species.

(]olor: Shell color and pattern are also useful taxonomic

characters. Most small specie's are translucent to opaque

white. Mo.st larger species are white, but some range

from gray or greenish-gray, to brown, amber, flesh-col-

ored, yellow, orange, pink, rose, or even red. The base

color may he overlain with various pattern elements of

G. A. Coovert and H. K. Coovert, 1995

Page 51

a different color. These can consist of spots, irregular

blotcfies, spiral lines or bands, axial or oblique bands or

streaks, oblique lines, or various combinations of these

features.

COMPARATIVE ANATOMY OF

MARGINELLIFORM GASTROPODS

External Anatomy: Features of the external anatomy

of 82 species of marginelliform gastropods have been

reviewed and summarized (Coovert, 1987g). Since then,

a number of additional species have been reported in

the literature (Coovert, 1988e:37). Our additional re-

search, plus notes and excellent photographs of the living

animals shared by others, provided a clearer picture of

the external anatomy of this group. Data on 183 species

are summarized in this paper. The four basic morpho-

logic types are summarized first, along with a discussion

of general features of the externa! anatomy. Further

details of the external anatomy of each genus can be

found in the systematic section.

The four distinct animal "types" delineated and sum-

marized in Coovert (1987g) are based primarily on fea-

tures of the head. The most common marginelliform

animal is termed a "Type 2 animal," which has a simple,

unmodified head described by some authors as "bifur-

cate" (figures 7, 8). The long, slender tentacles emerge

from the anterior end of the head, resulting in a bifurcate

appearance. Viewed from above, as most photographs

are oriented, this bifurcation is not readily apparent. The

long, slender tentacles have an eye located on a slight

swelling at their bases. The siphon, as in all marginelli-

form gastropods, is a simple tube formed by the inrolled

edges of an elongated expansion of the mantle edge. The

siphon in Type 2 animals is moderately long to very long,

and simple or weakly notched distally. A few cystiscid

and most marginellid genera have a Type 2 animal,

which is considered to be the generalized type.

Type 1 animals, represented by Afrivoluta and Mar-

ginellona, are characterized by a broad head with a

longitudinal medial furrow or channel (figures 5, 6). This

channel was described by Harasewych and Kantor (1991:

10) as a "deep tubular channel along dorsal mid-line of

head," apparently formed by the ventral fusion of the

tentacle bases. These same authors (op. cit., p. 13) noted

that this medial channel had not been reported in A/-

rivoluta, but our careful examination of a color trans-

parency of Afrivoluta pringlei (ex. W. R. Liltved) has

revealed that both genera share this feature. Type 1

animals are further characterized by relatively short ten-

tacles, a simple, moderately long siphon, and a distinct,

subtriangular lateral lappet on each side of the head. A

small red eye is present on each lateral lappet in Afri-

voluta. Eyes are completely lacking in Marginellona.

Type 3 animals are characterized by an elongate head

that is longitudinally split dorsally, with the anterior end

bifurcate (figures 9, 10). This configuration could be de-

scribed as either having the tentacles fused ventrally,

resulting in a longitudinal dorsomedial channel, or al-

ternatively, as having a longitudinally split head com-

pletely lacking tentacles. A detailed anatomical study is

needed to clarify this situation. The siphon is either very

short and not readily apparent, or completely absent.

Eyes are located on the sides of the head, usually in a

conspicuous bulge. Type 3 animals are represented by

the genus Cystiscus and probably also Crithe.

Type 4 animals have a somewhat flattened, longitu-

dinally split head, normally with tentacles (figures 11,

12). The anterior half of the head is completely split,

usually with the anterior tips pointed and divergent, but

capable of closing together. These two lobes are very

flexible and capable of much movement. The posterior

half is only split dorsally, and fused ventrally. Tentacles,

which are rarely absent, are short to long. The eyes are

located on the sides of the head, slightly below the ten-

tacle bases. The siphon is short to long, simple, and pro-

trudes through the siphonal notch at the anterior end of

the shell when the animal is active. Type 4 animals are

represented by the genera Persicula and Gibberula. Can-

alispira animals differ from the typical Type 4 animal

in lacking tentacles and a visible siphon (cf. Systematic

section for more detail).

The siphon of most marginelliform gastropods emerg-

es from the left side of the head, but has been described

as fused to the head in Marginellona by Harasewych

and Kantor (1991;table 3), although these authors stated

that the siphon is "fused to head just left of posterior

margin of mid-line channel" (op. cit., p. 10). It was

described as fused to the mid-dorsal line of the head in

Priintim marginatum by Graham (1966:135). All species

studied by us (Table 3) have a siphon fused to the left

side of the head. There seems to be some difference in

interpretation, but probably all marginelliforms have a

siphon fused to the head at least slightly left of center.

The mantle is another external anatomical feature that

is useful taxonomically. Within higher ta.xa, the degree

of mantle extension over the external shell surface varies

from none to complete coverage. Extended mantles can

be smooth, have low, rounded tubercles or pustules, or

possess elongate papillae that are either simple or

branched. The terms pustulose and papillose are used

here.

Most marginelliform gastropods have a broad, flat foot,

although it may be very narrow in a few species. The

relative size and length of the foot may have some value

taxonomically. Some species have a medially indented

foot, and some (e.g. Hyalina hyalina, H. pallida, and

Prunum martini) have an anterior, transverse pedal

groove. As far as is known, females have a medial, an-

teroventral pedal gland on the sole of the foot. In some

Gibberula species, the anterolateral edges of the foot are

raised and laterally rolled, forming what Gofas and Fer-

nandes (1988:22) described as "parapodia." All these

characters may have taxonomic value and need further

study.

Animal coloration is very useful at the species level,

but has limited value for higher classification. Gofas and

Fernandes (1988) and Gofas (1989a, 1990, 1992) provid-

Page 52

THE NAUTILUS, Vol. 109, Nos. 2 &

Figures 5-8. Marginellid external anatomy. 5, 6. Afrivoluta pringlei Tomlin, 1947, e.xample of Type 1 animal. After Liltved

(1985) plus pers. obs. of photos (ex. W. R. Liltved), Off Danger Point, western Cape Province, at 2-46 m. Ca. X 0.65. 5. Animal

viewed from right side. 6. Dorsal view of head. 7, 8. Prunum aff. aletes Roth, 1978, example of Type 2 animal. GAC 1518, small

island just off Tamarindo Beach, S. of Tamarindo Diria Hotel, Guanacaste Province, Costa Rica, under rocks, low tide Shell length

19.4 mm. 7. .'\nimal viewed from right side. 8. Dorsal view of head. Figures 9-12. Cystiscid external anatomy, 9, 10. Cystiscus

minutissimus (Tenison-VVoods, 1876), example of Type 3 animal. After Murray (1970) and Coleman (1975). 9. .\ninial viewed

from above. Ca. 9.0 X. 10. Dorsal view of head. Ca.l8 X. 11, 12. Persicula interruptohneata (Megerle von Miihlfeld, 1816).

example of Type 4 animal. GAC 1581, N. end of Playa La Galera, Isla de Margarita, Venezuela, NE side of bay, hand-dredged

on sandy mud with small patches of Turtle Grass. Length 14.7 mm. 1 1. Dorsal view of head. 12. .\nimal viewed from right side.

ed color plates of live animals, making excellent use of

coloration or "polychromatism" for species differentia-

tion.

Radula: The radulae of very few species (20 fide Coo-

vert, 1989b) were known at the time of the last generic

revision of this group (Coan, 1965), Much additional

radular information has been subsequently published (see

Coovert, 1989b for sutnmary; Coovert and Coovert, 1990).

This published data, combined with our unpublished

data and observations of SEMs (ex. Dean Hewish), brings

the number of species studied to 190. An additional 30

species are known to be non-radulate. Of the 31 Recent

genera recognized in this revision, the radulae of only

two are unknown.

Three basic radular patterns of marginelliform gas-

tropods have long been recognized by the senior author.

In addition to non-radulate species, the radulate species

fall into two different groups. One group has a relatively

long, narrow radula, the other group has a shorter, wider

radula. The most fundamental differences between these

two groups were not apparent until recently. In fact,

Coovert (1989b:31) erroneously assigned the genera al-

lied to Austroginella to the long, narrow radulate group

based solely on radular width.

The first major difference noted between these two

groups was the discovery of paired, wing-like extensions

of the subradular membrane, found only in species with

a long, narrow radula (and not in genera allied to Aus-

troginella). These structures, which persist in routine

tnaceration with KOH, were first noticed by Dean Hew-

ish (pers. conmi), and later confirmed by us in a number

of species. They were originally called "membranous

wings," and appeared to be attached anteriorly to the

G. A. Coovert and H. K. Coovert, 1995

Page 53

radular ribbon at the bending plane. They are not readily

visible after permanent mounting on microscope slides.

We follow Coan (1965) in useage of the term rachidian

plate rather than "rachidian teeth" because of the wide,

multicusped nature of these structures in marginelliform

radulae.

We discovered that two different conditions of ra-

chidian plate development occur on the posterior end of

the radula where plates are initially secreted. Species

with a long, narrow radula have a series of at least 6,

and up to 12 or more, gradually formed rachidian plates

at the posterior end, which are enclosed in a radular sac.

These plates, initially narrower and rudimentary, are

extremely thin, clear, and very difficult to discern in slide

preparations. Progressing anteriorly, these plates become

wider and more visible, first developing medially, with

lateral portions forming next, finally followed by full

cusp development. We refer to these separate undevel-

oped plates as nascent rachidian plates, and the gradually

developing series as a developmental series. Just anterior

to the nascent plates, a series of up to 40 or more fully

formed plates are distinctly brown-tinged, followed by

fully developed, clear, normal-appearing plates. In con-

trast, species with a short, wide radula have only one or

two nascent plates posteriorly. These plates are fully

formed with completely developed cusps, but are ex-

tremely thin, clear, and often very difficult to discern in

microscope slide preparations. The first plate often ap-

pears as a "ghost image." We term these preformed

nascent rachidian plates. Just anterior to these nascent

plates, a series of 4 to 5 fully formed brown-tinged plates

are followed by fully developed, clear, normal-appearing

plates. The buccal pouch, characteristic of marginellids

in the strict sense (see Internal Anatomy section), contains

the posterior end of the radula in this group. Thus, two

fundamentally different conditions exist in both overall

structure and development of the posterior end of mar-

ginelliform gastropod radulae.

In our search for taxonomic characters of marginel-

liform radulae, we noted that Ponder (1970:70) reported

two completely separate odontophoral cartilages in Gib-

berula (as Diluculum sp.). A pair of anteriorly fused

odontophoral cartilages (described as a single cartilage

in Volvarina taeniolata. see Fretter, 1976:329) were de-

scribed from all other studied species. None have been

illustrated. We dissected Persicula interruptolineata

(Megerle von Miihlfeld, 1816), a close relative of Gib-

berula, to confirm the presence of separate odontophoral

cartilages. Rather than underlying the radular ribbon as

in typical marginellids, the two separate odontophoral

cartilages of Persicula each fit into one of the "mem-

branous wings" that flank the radular ribbon (figures 15,

16). Further w ork determined the shape and attachments

of these wing-like membranous structures. These paired,

hood-like extensions of the subradular membrane (more

accurately termed the cuticle, see Hyman, 1967:236),

which serve to receive the odontophoral cartilages, are

here referred to as odontophoral cartilage hoods (figures

13-16). Nothing resembling these structures has been

Table 2. .Anatomical charactt-r states utilized in Table S

Buccal poucli

0 — absent

I — present

2. Neogastropc id radular sac 0 — present

1 — absent

3 General radular t\ pe 0 — cystiscid

1 — marginellid

2 — absent

4 Specific radular type 0 — absent

3 — Type 3 radula

5 — Type 5 radula

6 — Type 6 radula

6m — modified Type 6 radula

7 — Type 7 radula

.5 Odontophoral cartilages 0 — separate

1 — anteriorly fused

2 — fused both anteriorly and

posteriorly

3 — absent

6. Siphon 0 — attached left of head

1 — fused to head

7 Valve of I.eiblein 0 — present, no bypass tube

1 — present, with bypass tube

2 — absent

8, Esophageal caecum 0 — absent

1 — present

9. Gland ol Leiblein 0 — small, narrow gland, no ter-

minal bulb

1 — large, sacculate gland, no

terminal bulb

2 — with terminal bulb and long,

convoluted duct

10 Duct of gland of Lei- 0 — empties into esophagus pos-

blcui terior to nerve ring

1 — passes through nerve ring,

emptying into anterior end

of proboscis either into buc-

cal cavity or anterior end of

anterior esophagus

11 Paired salivary glands 0 — ascinous

1 — tubular

12. Paired salivary gland at- 0 — attached to esophagus

tachment 1 — free of esophagus

13. Paired salivary glands, 0 — embedded in walls of esoph-

ducts agus

1 — attached to esophagus

2 — free of esophagus

14. Accessory salivar\ glands 0 — absent

1 — single gland present

reported in marginellids with a short, wide radula. A

radula from a specimen of Priinum aff. aletes Roth, 1978,

which was not macerated in KOH, revealed anterolateral

extensions of the radular ribbon cuticle. These, however,

were narrower lateral flanges that differ functionally from

the odontophoral cartilage hoods of Persicula. These ex-

tensions of the cuticle apparently help hold the radular

ribbon in correct alignment with the underlying odon-

tophoral cartilage. We term these structures marginal

cuticular flanges (figures 17-20). Unhke odontophoral

cartilage hoods, they are lost during routine maceration

Page 54

THE NAUTILUS, Vol. 109. Nos. 2 & 3

Table 3. Suiiiniar) ol aiiatniiiical icatures of margiiielliform gastropods ('liaractcr stales given in Table 2 Sources of data given

in Tabic 4 ''' indicates character state unknown or uncertain.

Cvstiscidae

Character states

10

11

12

13

14

Pcrsicula inlerruplolineala

Pcrsicula masirana

Gibberula sp.

0?

Marginellidae

G. A. Coovert and H. K. Coovert, 1995

Page 55

odontophoral cartilage

hood

marginal cuticular

flange

14

odontophoral cartilage

Figures 13-16. Persicula interruptolineata (Megerle von Miihlfeld, 1816), showing details of cystiscid radular morphology. GAC

1581, data in Table 1, Rachidian width enlarged ISC^c for clarity. 13. Left lateral view, odontophoral cartilages removed, only

posterior end of radular sheath shown. 14. Same, dorsal view. 15. Dorsal view, odontophoral cartilages in place, muscles and

connective tissue removed. 16. Same, anterior view. Figures 17-20. Pntnum aff. aletes Roth, 1978, showing details of marginellid

radular morphology. GAC 1518, data in Table 1. 17. Left lateral view, odontophoral cartilage removed. 18. Same, dorsal view.

19. Dorsal view, odontophoral cartilage in place, muscles and connective tissue removed. 20. Same, anterior view.

in KOH, suggesting additional differences in composi-

tion.

Morphological evidence suggests that there are two

fundamentally different types of radulae. The first type

has a very long, triserial or uniserial, C-shaped radula,

composed of 80 to 200 or more very narrow, usually

overlapping, arched rachidian plates, with relatively few

(3 to 15) cusps. This type of radula has a pair of odon-

tophoral cartilage hoods, each receiving a separate odon-

tophoral cartilage, which flank the radular ribbon an-

teriorly. The posterior end of the radula begins as a

developmental series of nascent plates and is enclosed in

a typical neogastropod radular sac. This type of radula

is found in all species here assigned to the Cystiscidae

and is termed a cystiscid radula (figures 13-16). The

second type has a uniserial, J-shaped ribbon composed

of 10 to 80 broad, overlapping, flat rachidian plates,

usually with numerous cusps (9 to 80, rarely 4 to 7). This

type of radula has marginal cuticular flanges, and the

odontophoral cartilages are fused anteriorly into a single

structure underlying the radular ribbon. The posterior

end of this type of radula starts with 1 or 2 preformed

nascent rachidian plates and is enclosed in a marginellid

buccal pouch. This second type is found in all radulate

species here assigned to the Marginellidae s. str., and is

termed a marginellid radula (figures 17-20).

Coovert (1989b:31-33) divided marginelliform radu-

lae into seven distinct types based on characters of the

rachidian plates and the presence or absence of lateral

teeth. In addition to these seven types, which are dis-

cussed below, two new types are recognized. The ori-

entation of rachidian plates refers to the large, dorsal

Page 56

THE NAUTILUS. Vol. 109, Nos. 2 & 3

Marginellid Radulae

Type 5

Modified Type 6

Cystiscid Radulae

Type 1 Type 2

Type 3

Figures 21-24. Cystiscid radular types 21. Pleniocystisciis jeivettii (Carpenter, 1857). example of Type 1 radula G,'\C radula no

093, Pirates Cove, San Luis Obispo, California, intertidal. Ex. M1823, shell length 4.9 mm. 22. Cystiscus angasi (Crosse, 1870),

example of Type 2 radula. Hewish radula U, Lonsdale Bight, Victoria, Australia. Ex. NMV F53748, shell length 1.7 mm. 23.

Pcrsicula persicula (Linne, 1758), example of Type 3 radula. GAC radula no, 183, Niodior, Senegal. Ex. AMNH 173759, shell

length 19.8 mm 24. Granulina hadria (Dall, 1889), example of Type -4 radula. GAC radula no. 038, Tampa Bay, Pinellas Co.,

Florida, sand and mud. Ex GAC Ace # 27-82, shell length 2.2 mm. Figures 25-30. Marginellid radular types 25. Atistroginella

muscaria (Lamarck, 1822), example of Type 5 radula GAC radula no. 181, Lakes Entrance, Victoria, Australia. Ex, DMNH 12435,

shell length 13 3 mm. 26. Serrata Iranslata (Redfield, 1870), example of Modified Type 6 radula GAC radula no 304. Raiatea,

French Polynesia. Ex. GAC Ace, # 2-91, shell length 4.3 mm. 27. Prunum apicinum (Menke, 1828), example of Type 6 radula,

GAC radula no. 147, Sunshine Key, Florida Keys, Monroe Co,, Florida, Ex, Peggy Williams colln,, shell length 95 mm. 28.

Margincllona gigas (Martens, 1904), example of Type 7 radula. After Thiele (1904;pl, 9, fig, 64), Ex, Holotype, shell length 80 —

100 mm. 29. Serrataginclla spryi (Clover, 1974), example of Type 8 radula, G.f^C radula no, 305, off Nacala Bay, N, Mozambique,

dredged in 20-30 m, sand, Ex. GAC; Ace. # 6-93, shell length 8,0 mm 30. Hydroginella tridenlata (Tate, 1878), example of Type

9 radula G.AC radula no 306, Margaret River, Western Australia, in sand, 4 6 m Ex G.^C \cc. # 13-90, shell length 7,8 mm.

section of the ribbon, not the veiitrally folded anterior

end. This orientation is such that the cutting edge of each

rachidian plate is posterior, and the basal edge, where

the plate is attached to the membrane, is anterior.

Type 1 through Type 4 radulae are all c\ sticid radulae,

possessing the characteristics previously defined.

Type 1 ("Triseriai Type") radulae (figure 21 ) are triserial,

with simple, thin, weak, lateral teeth that narrow to a

single, posteriorly-pointed cusp. The long, narrow radula

is composed of 66 to 103 rows of overlapping rachidian

plates. These plates are very narrow, weakly arched, with

3 to 7 sharp cusps along cutting edge. The basal edge of

each rachidian plate is slightly concave, resulting in a

weakly Ll-shaped plate, Odontophoral cartilage hoods

were noted by us as well as by Dean Hewish (pers, comm, ),

but the nature of the odontophoral cartilages is unknown.

The genus previously referred to as "Cystiscus" (Coo-

vert, 1989b), described herein as new, is the only one to

possess a Type 1 radula (radulae of 3 species studied).

Type 2 radulae (figure 22) are uniserial, consisting of a

G. A. Coovert and H. K. Coovert, 1995

Page 57

long, narrow series of over 80 to 220 rachidian plates.

These plates are overlapping (rarely separate), very nar-

row, weakly to strongly arched, with 4 to 15 very strong

cusps along cutting edge. The basal edge of each plate

is slightly concave, nearly straight, or convex medially,

resulting in a crescent-shaped, subrectanguiar, or asym-

metrically V-shaped plate. Odontophoral cartilage hoods

were noted in many species of Cystisctis. but were un-

known and unobserved at the time Crithe radulae were

extracted. The nature of the odontophoral cartilages is

unknown. Cystiscus (radulae of 14 species studied) and

Crithe (radulae of 1 species studied) both have a Type

2 ("Crithe / Cystiscus Type") radula. The radula of

Extra is unknown, but presumed to be similar.

Type 3 radulae (figure 23) are uniserial, consisting of a

long, narrow series of over 80 to 209 rachidian plates.

These moderately to strongly overlapping plates are nar-

row, moderately to strongh arched, with 5 to 14 sharp

cusps along cutting edge. The central cusp is often the

strongest. The basal edge of each plate is strongly con-

cave, resulting in a U- or V-shaped plate. The reports of

wishbone-shaped rachidian plates (Barnard, 1962:14-15;

Ponder, 1970:70; Ponder, 1973:fig. 2) are inaccurate and

due to an artifact of observation (see discussion in Coo-

vert, 1989b: 12-13). Odontophoral cartilage hoods were

noted in all three included genera, with a pair of separate

odontophoral cartilages noted in Persicula and Gibber-

ula. Persicula (radulae of 14 species studied), Gibberula

(radulae of 21 species studied), and Canalispira (radulae

of 2 species studied) all have a Type 3 ("Persicula Type")

radula.

Type 4 radulae (figure 24) are uniserial, consisting of a

long, narrow series of at least 90 to 159 rachidian plates.

These usually completely non-overlapping plates are nar-

row, moderately arched, with 9 to 12 clustered, raised

denticles located in staggered positions on the dorsal sur-

face, with a single (or asymmetrically paired) strong

central cusp protruding on the cutting edge. The basal

edge of each plate is weakly concave, resulting in a

weakly V-shaped plate. The rachidian plates are asym-

metrical and alternate as mirror images (figure 24). Al-

though odontophoral cartilage hoods have not been ob-

served in representatives of this group, their radulae are

otherwise typically cystiscid. A broad, membranous flange

on either side and extending the whole length of the

radula, effectively doubling the width, has been observed

(Dean Hewish, pers. comm.) when using the enzyme

Pronase for maceration. This is a less destructive method

than the use of KOH. This flange is certainly homologous

to odontophoral cartilage hoods, but the nature of the

odontophoral cartilages is unknown. No further anatom-

ical information is available. Granulina (radulae of 12

species studied) and Fugnu.s (radulae of 2 species studied)

both have a Type 4 ("Granulina Type") radula.

Type 5 through Type 9 radulae are all marginellid rad-

ulae, possessing the characteristics as previously defined.

Type 5 radulae (figure 25) are uniserial, consisting of a

relatively short, broad ribbon of 19 to 75 rachidian plates.

These usually overlapping plates are moderately wide,

weakly arched, with 8 to 22 strong cusps along cutting

edge. The central cusp is often the strongest. There are

often pits on adjacent plates to receive the tips of some

of the cusps, especially the central. The basal edge of

each rachidian plate is generally slightly concave, re-

sulting in a rectangular to chevron-shaped plate. The

odontophoral cartilages in Mesoginella and Austrogi-

nella were described as fused anteriorly (Ponder, 1970:

62; Ponder & Taylor, 1992:320), but no detailed data

exists as to marginal cuticular flanges. Protoginella (rad-

ula of 1 species studied), Alaginella (radulae of 9 species

studied), Austroginclla (radulae of 5 species studied),

Mesoginella (radulae of 16 species studied), Ovaginella

(radulae of 2 species studied), and Balanetta (radula of

1 species studied) all have a Type 5 ("Austroginclla /

Mesoginella Type") radula. The radula of Closia is un-

know n, but presumed to be similar.

Type 6 radulae (figure 27) are uniserial, consisting of a

relatively short, broad ribbon of 24 to 75 overlapping

rachidian plates. These plates are broad, nearly flat, with

9 to 45 (exceptionally 5) sharp cusps along cutting edge.

There are often pits on adjacent plates to receive the tips

of some or all of the cusps. The basal edge of each plate

is generally straight, resulting in an elongate, rectangular

"comb-like" plate. The odontophoral cartilages are fused

anteriorly. They are fused posteriorly in most species

(but not in Prunum aff. aletes and P. guttatum), leaving

a narrow, medial, longitudinal posterior slit in most Prun-

um and Volvarina (figures 38-42). (See following section

on internal anatomy.) A single cartilage was described

for Volvarina taeniolata (cf. Fretter, 1976:329). The

marginal cuticular flanges of Prunum aff. aletes, de-

scribed above, are very similar in other species of Prun-

um we examined. Prunum (radulae of 23 species stud-

ied), Volvarina (radulae of 42 species studied), Rivo-

marginella (radulae of 2 species studied), Bullata (radula

of 1 species studied), and Cryptospira (radulae of 4 spe-

cies studied) all have a Type 6 ("Prunum / Volvarina

Type") radula. Most of these are discussed and figured

in Coovert and Coovert (1990).

Modified Type 6 radulae (figure 26) are similar to typical

Type 6, but differ in having a shorter ribbon composed

of only 13 to 35 plates, averaging more cusps (22 to 59)

that project from a somewhat sinuous posterior edge, and

in often having thinner and more fragile rachidian plates.

Pits on adjacent plates to receive cusp tips were not

observed as in many typical Type 6 radulae. The odon-

tophoral cartilages of Serrata (as Haloginella) were de-

scribed by Ponder (1970:66) as fused except for a very

short posterior portion. Marginal cuticular flanges are

unknown. Serrata (radulae of 8 species studied) is the

only genus to have a Modified Type 6 ("Serrata Type )

radula.

Type 7 radulae (figure 28) are uniserial, consisting of a

relatively short, very broad ribbon of 56 to 80 overlap-

ping rachidian plates. These plates are very broad, nearly

Page 58

THE NAUTILUS, Vol. 109, Nos. 2 & 3

flat, with numerous (58 to 85) sharp cusps along cutting

edge. The basal edge of each plate is generally straight,

resulting in an elongate, rectangular "comb-like" plate.

Odontophoral cartilages and marginal cuticular flanges

are unknown. The monotypic genera Marginellona and

Afrivuluta both have a Type 7 {" Afrivoluta Type") rad-

ula.

Type 8 radulae (figure 29), here described, are uniserial,

consisting of a relatively short, broad ribbon of 38 over-

lapping rachidian plates. These plates are broad, nearly

flat, with 9 to 10 sharp cusps on the cutting edge. The

entire posterior edge, including all edges of the main

cusps, with numerous (total of ca. 70) small, secondary

cusps, giving the main cusps a serrated appearance. Pits

for reception of adjacent cusp tips were not observed.

The basal edge of each plate is slightly indented medially,

resulting in a very shallowly V-shaped plate. Odonto-

phoral cartilages and marginal cuticular flanges are un-

known. The Type 8 ("Serrataginella Type") radula is

unlike any others known, and is represented by "Mar-

ginella" spryi Clover, 1974, type species of a monotypic

genus herein described as new.

Type 9 radulae (figure 30), first described by Bouchet

(1989:79, fig. 3), are uniserial, consisting of a very short,

narrow, greatly reduced ribbon of 10 to 30 rachidian

plates. These weak plates overlap and are nearly flat,

with 4 to 7 cusps on the cutting edge. Pits for reception

of adjacent cusp tips are absent. The basal edge of each

plate is straight, resulting in a subquadrate plate. Odon-

tophoral cartilages and marginal cuticular flanges are

unknown. Hydroginella (radulae of 4 species studied) is

the only genus to have a Type 9 ("Hydroginella Type")

radula. One species is known to be parasitically associated

with fishes (Bouchet, 1989). Judging from the very sim-

ilar radulae of other species, it seems likely that all may

have such an association.

A third major group, comprised of non-radulate spe-

cies, completely lacks a buccal mass, odontophore, and

radula. This was confirmed by dissections of species in

the following genera: Dentimargo (Ponder, 1970:67, plus

species reported herein), Marginella s. str. (Graham, 1966:

139, Coan & Roth, 1976:220, plus species reported here-

in). Glabella (Coan & Roth, 1976:220), and Hyalina (Coan

& Roth, 1976:220, and herein). Other species have been

subsequently reported to lack a radula, some based on

unsuccessful radular extraction with KOH (see Coovert,

1989b tor review). Thus, a total of 31 species have been

inferred or demonstrated to lack a radula. We have at-

tempted routine radular extraction on separate occasions

from two specimens of the type species of Eratoidea, E.

margarita (Kiener, 1834), and have not found a radula.

Although not definitive, it is suggested tiiat this genus is

also non-radulate. This assemblage of non-radulate gen-

era is an artificial group, with Hyalina l)ei()Mging to a

different lineage (see discussion in systematic section).

Internal Anatomy: Prior to our studies, only 13 species

of marginelliform gastropods had been studied anatom-

ically. Harasewych and Kantor (1991:tables 2, 3) sum-

marized anatomical data of the 11 species known at that

time. Ponder and Taylor (1992) subsequently presented

data for two species of Austroginella. Few taxonomic

conclusions could be drawn from such a relatively small

sample because the anatomy varied so greatly. Our dis-

sections added anatomical data for 11 additional species,

and further data for one previously reported species (Ta-

bles 1-4). Combined with conchological and radular data,

this greatly expanded body of anatomical information

enabled taxonomic groupings to become evident, making

a reliable and stable classification possible. Details of the

foregut were emphasized because of their greater taxo-

nomic value. Our primary objective is a presentation of

the higher classification, therefore, more detailed reports

on the anatomy of individual species will be published

elsewhere. A discussion of foregut anatomy follows, along

with a summary of known anatomical information for

these 24 species.

The proboscis of marginelliform gastropods is rela-

tively short and pleurembolic. The retractor muscles are

inserted on the sides of the proboscis, resulting in the

basal portion being invaginated, forming a proboscis sac

(Fretter & Graham, 1962:150). The proboscis varies

somewhat in shape among genera. The small mouth is

always situated terminally at the anterior end. Most gen-

era have a proboscis that is rounded or blunt distally,

and, when retracted, is generally about two-and-one-half

to four times as long as broad. When feeding, the pro-

boscis may be extended to a length approximating the

length of the shell in at least some species (Fretter, 1976:

329). Both species of Dentimargo studied have a distally

pointed proboscis about four times as long as broad when

retracted (figure 33). Serrata aff. mustelina has a more

bulbous, distally rounded proboscis about one-and-a-half

times as long as broad. Gibberula and Persicula (figures

35, 79) both have a short, broad proboscis about two

times as long as broad.

The mouth opens directly into the buccal cavity. No

jaws have been observed in any marginelliform species.

In radulate species, the buccal cavity leads posterodor-

sally to the anterior esophagus and posteroventrally to

the buccal mass. In non-radulate species, the buccal cav-

ity leads with little or no differentiation to the anterior

esophagus.

The most significant hallmark of the family Margi-

nellidae, in the strict sense, is the possession of a "buccal

pouch" or "buccal caecum" in radulate species. Both

Fretter (1976:333) and Harasewych and Kantor (1991:

16) recognized the significance of this structure and its

presence only in marginellids and toxoglossans. The buc-

cal pouch (figure 31), the term we prefer for this struc-

ture, was first discovered and named by Graham (1966:

137), who noted that "in marginellids the most interest-

ing feature of the gut is the situation of the buccal mass,

wholly withdrawn into a caecum from its typical place

on the floor of the buccal cavity (op. cit., p. 147)." The

l)uccal mass, described as "rather reduced" by Graham

(1966:137) and consisting of the odontofihoral cartilages.

G. A. Coovert and H. K. Coovert, 1995

Page 59

Table t. Sources of anatomical data utilizcil in Tabic 3

Species (current nonicuclaturcl

Source

Cystiscidae

Pcnicula intcrruptolincata (Megcrle von Miihlleld, 1816)

Pcrsiciila masirana Roth 6i Petit, 1972

Cibherula sp

Marginellidae

Marginvllona gigaa (Martens, 1904)

Serrata aff mustclina (Angas, 1871)

Amtroginclla johnstoni (Petterd, 1884)

Austroginclla muscaria (Lamarck, 1822)

Mesoginrlla pygmaca (G. B. SouerhN II, 184(i)

Volvarind all ciicnci (Kiener, 1834)

Volvarina tacmulata Morcli, ISHO

Priinnm aff. alctcs Rotli, 1978

Pruntim guttalum (Dill\v\ii. 1817)

Pruntim marginatum (Born, 1778)

Pniniim martini (Petit, 18.53)

Pninum prunum (Gmelin, 1791)

Pruniuii rooscvciti (Bartscli & Relider, 1939)

Bullala huUata (Born, 1778)

Htialina lujalina (Tfiiele, 1913)

Hi/alina pallida (Linne, 17.58)

Dcntimargu cairoma (Brookes, 1924)

Dcntimargo cburneola (Conrad, 1834)

Marginclla dcsjardini Marche-Marehad, 1957

Marginella glabella (Linne, 1758)

Marginclla scliantiani Marclie-Marcliad & Rosso, 1979

herein; see Table 1 for data

herein; see Table 1 for data

Ponder (1970), as Diluculuni sp,

Harasewych and Kantor (1991)

Ponder (1970), as Volvarina (llalogindia) mustelina, the New

Zealand species differs from the Australian species; see

Coovert (1989b: 16) for comments

Ponder and Tavlor (1992)

Ponder and Taylor (1992)

Ponder (1970), as Mesoginella iSinuginclla) pygmaea

herein; see Table 1 for data

Fretter (1976)

herein; see Table 1 for data

herein; see Table 1 for data

Graham (1966), as Marginella mariginata

Marcus and Marcus (1968), as Mariginella fratercultis

herein; see Table 1 for data

herein; see Table 1 lor data

herein; see Table 1 for tlata

Eales (1923), as Marignella hyalina

Coan and Roth (1976) jihis herein; see Table 1 for data

Ponder (1970), as \'olvarinella cairoma

herein; see Table 1 for data

Graham (1966)

herein; see Table 1 for data

herein; see Table 1 for data

radula, and associated connective tissue and muscles, is

contained in the buccal pouch. What characterizes the

buccal pouch is the sphincter at its anterior opening into

the buccal cavity and the fact that it is a separate caecum

or sac that is blind posteriorly, where the radula is ini-

tially secreted. This is further discussed by Graham (1966;

138), who noted that radular formation occurs "at the

innermost end of the pouch, which therefore corresponds

to the radular sac of more normal gastropods," Ponder

(1970:76) also noted that the buccal pouch is derived

from the radular sac. Fretter (1976:329) observed that

the "teeth arise at the innermost end of the pouch where

there is no clear demarcation of the radular sac typical

of prosobranchs." The blind end of a buccal pouch thus

corresponds to the posterior end of the radular sac. Our

observations of Prunum, Volvarina, and Bullata species

confirmed the presence of a buccal pouch. The genera

Serrata. Mesoginella, and Austroginclla, dissected by

Ponder (1970) and Ponder and Taylor (1992), likewise

possess a true marginellid buccal pouch. In the true mar-

ginellids, the odontophore extends beyond the posterior

end of the radula.

The absence of a buccal pouch in non-radulate species

has been confirmed by dissections of species of Hyalina,

Dentimargo, and Marginella (figures 32-34). Graham

(1966:139) described what he believed to be a vestigial

buccal pouch in Marginella desjardini. Ponder (1970:

76) considered this to be the accessory salivary gland.

Our dissections of Marginella glabella (figure 34) and

M. sebastiani, both close relatives of M. desjardini, con-

firmed this contention, because an accessory salivary gland

was found in both species, but neither had a vestigial

buccal pouch. Ponder's (1970: fig. 3B) dissection of Den-

timargo cairoma, and our dissection of D. eburneola

(figure 33), show the complete absence of a buccal pouch.

There is insufficient information about Hyalina hyalina

in Eales (1923) to draw any conclusions. Coan and Roth

(1976:220) likewise did not provide sufficient detail for

H. pallida, although they did mention the absence of a

radula and odontophore. Our dissection of H. pallida

(figure 32), confirms the absence of a buccal pouch.

A typical neogastropod radular sac enclosing the pos-

terior end of the radula was found in both species of

Persicula we dissected (figure .35). In Persicula inter-

ruptolineata and in the Gibberula studied by Ponder

(1970:fig. E, r.s), the radular sac is readily distinguished

because it protrudes from the posterior end of the pro-

boscis in a characteristic downward curve (figures 35,

79). It is further characterized by a slightly bulbous pos-

terior end which encases the nascent end of the radular

ribbon (cjv. previous section). The radular sac extends

well behind the odontophore and is an extension of the

radular sheath, formed from the subradular membrane.

The buccal mass in Persicida masirana is morphologi-

Page 60

THE NAUTILUS, Vol. 109, Nos. 2 & 3

gland of Leiblein

esophageal caecum

esophagus

duct of gland

of Leiblein

mouth

buccal pou

paired salivary glands

buccal mass

31

gland of Leiblein

esophagus

mouth

duct of gland

of Leiblein

paired salivary glands

accessory salivary gland

Figures 31-32. Anatomical features dI llie foregiit of Pruiiini Data in Tahle I 31. Pninum aff. aleies Rotli, 1978 12-4 X. 32.

Hyalinu pallida (Linne, 1758). 15.3 .\,

cally the same, but the proboscis is simply longer, com-

pletely containing the radular sac. The characteristic

shape and protrusion of the radular sac seems to be fairly

typical for most Neogastropods (including Olividae), with

the prominent exception being the Marginellidae, s. str.

The radular sac thus differs in several respects from the

buccal pouch of iiiargiiiellids, especially in the lack of

an anterior sphincter. All cystiscid genera studied have

the same general type of radula, previously termed a

cystiscid radula ((jv.). Based on the similar na.sceiit ends

of their radulae, all cystiscid genera are a.ssumed to pos-

sess a true radular sac similar to that found in Persictila

and Gibbcrula. A radular sac has actually been observed

in several incompletely cleared cystiscid radulae, in-

cluding the triserial group "Cystiscus," named as a new

genus herein

Odontophorai cartilages, which we prefer over "bol-

sters, " "odontophore, or "cartilages," is a more specific

term and clearly differentiates these structures from the

odontophore, which is often used as a general term for

the entire radular mass. These cartilages support and

stiffen the radular mass and are composed of large Ley-

G. A. Coovert and H. K. Coovert, 1995

Page 61

esophagus

nerve ring gland of Leiblein

mouth

accessory salivary gland

duct of gland

of Leiblein

pairea salivary gland

mouth

esophagus

accessory salivary gland

duct of gland

of Leiblein

gland of Leiblein

paired salivary glands

34

mouth

valve of Leiblein

esophagus

buccal mass

accessory salivary gland

gland of Leiblein

35

Figures 33-35. Anatomical features of the foregut of Marginellini and Persiculinae^ Data in Table 1 33. Dentimargo ebiirneola

(Conrad, 1834). 22.4 X. 34. Marginella glabella (Linne, 1758). 5.2 X. 35. Persicula interruptolineata (Megerle von Miihlfeld,

1816). 16.8 X.

Page 62

THE NAUTILUS, Vol. 109, Nos. 2 & 3

39

42

Figures 36-42. Odontophoral cartilages of cystiscids and marginellids Data in Table 1 36. Persicula interruptolineata. GAC

1581A. Length 0 97 mm 37. Persicula masirana. GAC Ace. # 4-94B Length LOl mm 38. Volvarina aff. avena. GAC 1172B

Length 0 88 mm. 39. Prtuwm aff. aides. GAC 1518B. Length L50 mm. 40. Prunum guttatum. GAC 1172A. Length 0.75 mm.

4L Prunum prunum. GAC 1582. Length 2.12 mm. 42. Prunum roosevelti. GAC Ace # 26-88A. Length 1.67 mm.

dig cells (chondroid tissue) by which they are easily rec-

ognized. Two fundamentally different conditions also

exist in this character complex.

In all true marginellids examined, the two odonto-

phoral cartilages are fused at least anteriorly, effectively

functioning as a single structure underlying the radula.

Ponder (1970:62) described the odontophoral cartilages

in Mesoginella pijgmaea as "fused into a single structure

in the front half of the odontophore but posteriorly are

connected by a transverse muscle in the usual way."

Ponder (1970:66) described similarly fused cartilages in

Serrata aff. mustelina, which are separated for only a

short distance posteriorly. The two species of Austrogi-

nella discussed by Ponder and Taylor (1992:320) are

likewise described as having anteriorly fused cartilages.

The odontophoral cartilages in Marginellona were not

described by Harasewych and Kantor (1991). Fretter

(1976:329) simply described a single cartilage for Vol-

varina taeniolata. Our dissection of V. aff. avena re-

vealed a pair of cartilages fused both anteriorly and

posteriorly (figure 38). They are slightly bowed medially,

leaving a medial slit about half the length of the entire

structure, with the anterior margins raised and thick-

ened, and truncated anteriorly. We suspect that V. tae-

niolata may be similarly structured. The closely related

genus Prunum shows a similar condition. These struc-

tures were insufficiently described in Graham (1966) and

Marcus and Marcus (1968) to draw any conclusions. Our

dissections of four species of Prunum revealed that they

all have strongly raised, thickened anterolateral margins,

effectively forming a trough or channel for the radular

ribbon (figure 20). The odontophoral cartilages in P.

roosevelti (figure 42) are relatively narrow, rounded an-

teriorly, completely fused on the very long and narrow

anterior half, and slightK bowed and narrowly fused

posteriorly, confining a medial slit to the posterior half.

The odontophoral cartilages in Prunum prunum (figure

41) are relatively short and broad, fused anteriorK and

posteriorly, with a narrow medial slit. The anterior end

is shallowly emarginate. Prunum guttatum (figure 40)

and P. aff. aletes (figure 39) differ because their odon-

tophoral cartilages are \'-shaped, and narrowly fused

anteriorly, but not posteriorly. The fused cartilages in P.

guttatum (figure 40) ha\e the posterior tips slightly con-

vergent but not touching, with the anterior end truncate.

In P. aff. aletes (figure 39), the anterior end is truncate

to shallowly emarginate, and the posterior tips are di-

vergent, forming a distinct \'-shape, but in situ (figure

19) connective and muscle tissue hold these posterior tips

closer together. In all cases, this fused pair of odonto-

phoral cartilages underlies the main part of the radula

(figures 19, 20). The radula wraps around the anterior

G. A. Coovert and H. K. Coovert, 1995

Page 63

end of the odontophore, tliis point being effectively

termed the bending plane. The shorter anterior portion

of the radular ribbon, often showing broken cusps, thus

hes under the anterior end of the odontophore, giving it

the characteristic J-shape. Our data indicates that the

odontophoral cartilage shape is consistent within each

species, but differs between them, and may prove to be

a useful taxonomic character.

All cystiscid genera examined have two completely

separate odontophoral cartilages. Each cartilage fits into

one of the lateral odontophoral cartilage hoods of the

subradular membrane, flanking and not underlying the

radula (figures 15, 16). Ponder (1970:70) described the

odontophoral cartilages and muscle attachments for a

species of Gibberula. Our dissections of Persicula inter-

ruptolineata (figure 36) revealed two separate subtrian-

gular cartilages that are broad and rounded anteriorly,

and narrowed posteriorly. In P. masirana (figure 37), the

separate cartilages are elongate, rather uniform in width

throughout, and pointed anteriorly. The radulae of near-

ly all other cystiscid genera have been described earlier

in this paper, and most have odontophoral cartilage hoods.

On this basis, we infer that they possess two separate

odontophoral cartilages.

The anterior esophagus in radulate genera is situated

dorsally within the proboscis, the buccal mass lying below

it. In non-radulate species, the anterior esophagus is more

centrally placed. Upon exiting the retracted proboscis

posteriorly, the esophagus generally makes a sharp S-bend

prior to passing through the nerve ring. A valve of Lei-

blein, when present, is located anterior to the circum-

esophageal nerve ring. Ponder and Taylor (1992:321)

described the valve of Leiblein in Austroginella as being

partially encased in the nerve ring anteriorly (figure 80).

Serrata and Mesoginella have a well-developed valve

located just anterior to the nerve ring. These three genera

have a narrow, non-glandular bypass tube that is adher-

ent ventrally to the valve of Leiblein. Marginellona has

a large valve of Leiblein situated anterior to the nerve

ring (figure 80). The two Persicula species we dissected

also have a distinct valve of Leiblein, placed well anterior

to the nerve ring (figure 35). No bypass tube was found.

Gibberula has a similarly placed valve of Leiblein, but

has a separate glandular tube that bypasses the valve

(figure 79). Volvarina, Hyalina, Pninuvi, Btillata. Den-

tiniargo, and Marginella all lack a valve of Leiblein.

Marcus and Marcus (1968:65) discussed an "individually

inconstant valve of Leiblein" in Prunum martini. In

some Prunum species we dissected, what at first ap-

peared to be a possible valve of Leiblein simply turned

out to be food lumps in the esophagus that were incon-

sistent in their location or occurrence. We believe that

the so-called "individually inconstant valve of Leiblein '

in Prunum martini were simply food lumps in the esoph-

agus of some individuals.

The esophagus narrows upon entering the nerve ring

anteriorly, resuming its original size after exiting pos-

teriorly. A much enlarged sac-like esophageal caecum

was found just posterior to the nerve ring in all species

of Volvarina. Prunum (figure 31), and Bullata studied.

This structure was first described in Prunum margina-

tum as an "oesophageal caecum" (Graham, 1966:135,

fig. 1). Eales (1923) did not mention this structure in

Hyalina hyalina, nor was it mentioned by Coan and Roth

(1976) for H. pallida. Our dissection of H. pallida (figure

32) revealed a thin-walled, but very distinct, esophageal

caecum. An esophageal caecum is absent in all other

marginelliform genera that have been studied anatom-

ically.

The most conspicuous foregut gland is the gland of

Leiblein, also called the "unpaired gland, "unpaired

foregut gland, or "poison gland." This gland occurs in

three different forms. In Persicula (figure 35) and Gib-

berula (figure 79), this gland is a small, short, often slight-

ly folded gland emptying directly into the posterior end

of the mid-esophagus, well posterior to the nerve ring.

In Marginellona (figure 80), the gland of Leiblein is

extremely large, sacculate, broad anteriorly and tapering

posteriorly (Harasewych and Kantor, 1991:11). It also

empties directly into the mid-esophagus posterior to the

nerve ring. All the remaining marginelliform genera

studied anatomically have a gland of Leiblein with a

long, convoluted duct. The gland in these genera ends

distally in a terminal bulb that is either rounded or weak-

ly to distinctly pointed distally. The terminal bulb is

narrow, elongate, and rounded distally in Der^timargo

(figure 33). In Marginella, it is shorter, more swollen,

and ends in an acuminate, pointed tip (figure 34). In

Serrata. Mesoginella, and Austroginella (figure 80), the

relatively large duct from the gland of Leiblein, which

is about the same diameter as the esophagus, empties

into the posterior end of the mid-esophagus just posterior

to the nerve ring but does not pass through the nerve

ring. These three genera also possess a well-developed

valve of Leiblein with a ventral by-pass tube. In Prunum

(figure 31), Bullata, Volvarina, Hyalina (figure 32), Den-

timargo (figure 33), and Marginella (figure 34), the long,

convoluted duct from the gland of Leiblein narrows as

it passes through the nerve ring, then continues as a very

narrow tube to the anterior end of the proboscis. These

six genera all lack a valve of Leiblein. The duct from

the gland of Leiblein empties either into the buccal cav-

ity or into the extreme anterior end of the anterior esoph-

agus, which is usually in very close proximity to the point

where the ducts of the paired salivary glands empty into

the buccal cavity. Ponder (1970:77, 79) described this

duct in Marginella desjardini as opening "ventrally into

the oesophagus a little in front of the nerve ring." This

is in error because Graham (1966:139) clearly described

this duct as opening "to the gut at almost the same level

as that at which the salivary ducts reach the same po-

sition," which is at the "extreme anterior end of the

proboscis, where they discharge, ... into the buccal cavity. '

This is mentioned because apparently a figure in Ponder

(1973:fig. 3N) is based on this misconception. This does

not weaken Ponder's hypothesis for the formation of the

poison gland (Ponder, 1970:77-80), but it indicates that

this intermediate stage is presently unknown.

Page 64

THE NAUTILUS, Vol. 109. Nos. 2 & 3

All inarginelliform genera have either tubular or as-

cinous paired salivary glands associated with the esoph-

agus. In many cases, tubular and ascinous salivary glands

appear to be found in different species of the same genus,

and thus may be of limited taxonomic value in these

genera. In all cases, the paired glands send fine, narrow

ducts to the anterior end of the proboscis, where they

empty into the buccal cavity. The placement of the glands

and the attachment of ducts differ between species. In

Marginellona. the paired salivary glands are found dor-

sal to the gland of Leiblein (figure 80), their ducts run-

ning anteriorly along the floor of the proboscis, free of

the esophagus. Serrata, Mesoginella, and Austroginella

all have glands that are attached to tfie esophagus just

anterior to the valve of Leiblein, the ducts embedded in

the esophageal walls for their entire length. Volvarina

taeniolata has the glands free in the body cavity and the

ducts free of the esophagus. In V. aff. avena. the paired

salivary glands are relatively large and fully contained

within the retracted proboscis, the ducts attached ante-

riorly to the esophagus. In Prunum. the paired salivary

glands are free of the esophagus. In P. prunum and P.

guttatum, the ducts are attached to the walls of the

esophagus. In the other Prunum species studied, the ducts

lay ventrally along the floor of the proboscis, free of the

esophagus. Details on the salivary glands of Bullata bul-

lata are unknown. The ducts in H. pallida are free of

the esophagus, the glands apparently tubular but poorly

preserved in the specimen studied. Both Dentimargo

species have the glands largely contained within the pro-

boscis. They are presumed to be tubular in D. cairoma.

but are ascinous in D. eburneola. Oddly, these glands

are not placed laterally in D. eburneola as is usual, but

instead are placed dorsally and ventrally, one gland sit-

uated above the esophagus, the other below the accessory

salivary gland (figure 33). In both species, the glands and

their ducts lay free. The paired salivary glands of the

three species of Marginella are located outside the pro-

boscis near the nerve ring, free from the esophagus. In

M. glabella, the ducts are attached to the duct of the

gland of Leiblein (figure 34), unlike those of M. sebas-

tiani. The paired salivary glands in Persicula and Gib-

berula are attached to the esophagus just anterior to the

valve of Leiblein (figure 35). The ducts of Persicula are

attached to the outside walls of the esophagus, but in

Gibberula this is unknown. Ponders illustration (1970:

fig. 3K) seems to indicate that these ducts are embedded.

Information on these glands anil their ducts is summa-

rized in Table 3.

The acces.sory salivary gland, when present, is always

single and empties into the buccal cavity. It is present

in all cystiscids studied, being ascinous in both species

of Persicula, but tubular in Gibberula. Within margi-

nellids, a tubular accessory salivary gland is present in

Serrata, but absent in Austroginella and Mesoginella. A

large, ascinous gland is present in Volvarina aff. avena,

but appears to l)e tubular in V. taeniolata as described

by Fretter (1976:329). Both species of Hijalina possess

an accessory salivary gland, which is tuliular in H. hy-

alina and apparently so in H. pallida (unreported in Coan

& Roth, 1976 and poorly preserved in our specimen). An

accessory salivary gland is present in only one species of

Prunum, being tubular in P. prunum. The gland is pres-

ent and tubular in Dentimargo and Marginella, but ab-

sent in Marginellona. Further work, including histology,

is needed. See Table 3 for a summary of the occurrence

of this gland.

Our primary objective was to discern useful taxonomic

characters, and we feel that the most useful features are

those of the foregut. Thus, the remainder of the digestive

system was not studied in any greater detail by us, nor

was the reproductive system. A brief summary of pub-

lished information on these other systems follows.

The presence or absence of an anal gland should be

further investigated. This structure was reported as pres-

ent and extremely large in Marginellona gigas by Har-

asewych and Kantor (1991:11). Serrata aff. mustclina,

Mesoginella pygmaea, Dentimargo cairoma (cf. Ponder,

1970), Austroginella johnstoni, A. muscaria (cf. Ponder

& Taylor, 1992:322), and Volvarina taeniolata (cf. Fret-

ter, 1976:329) were all reported as having an anal gland.

The only marginelliform reported to lack an anal gland

is the species of Gibberula reported by Ponder (1970:

71). Published accounts of the remaining species did not

report the presence or absence of this structure.

Main features of the reproductive system were sum-

marized in Harasewych and Kantor (1991:tables 2, 3).

Published information on the reproductive system is in-

complete for a number of species described anatomically.

Probably of most interest in the female reproductive

system is the presence of three glandular structures or

seminal receptacles joining the pallial oviduct between

the albumen gland and capsule gland in Dentimargo

cairoma (cf. Ponder, 1970:74), whereas only one is pres-

ent in Gibberula (op. cit., p. 75). Two such structures are

the usual number, as recorded in Mesoginella pygmaea

and Serrata aff. mustelina by Ponder (1970:74), Mar-

ginellona gigas by Harasewych and Kantor (1991:11),

Marginella desjardini by Graham (1966:140), and Vol-

varina taeniolata by Fretter (1976:fig. 2A). Information

is lacking for the other species. In the male reproductive

system, the penis is normally simple, lacking appendages

or lobes. But Ponder (1970:72, fig. 4C) described a bi-

lobed penis for Dentimargo cairoma that has a sharply

pointed posteroapical appendage, which he termed a

sheath. Our dissection of D. elmrneola likewise revealed

a sharply pointed apical appendage, but the penis was

not bilobed. The penis in Bullata Imllata is an extremely

large, simple, flattened, spatulate structure, about two-

thirds the body length. All other species studied have a

simple, unlobed penis. Refer to Harasewych and Kantor

(1991:tables 2, 3) for a sunnnary of additional characters.

Published information on marginelliform egg capsules

was reviewed in Cxwvert (1986c and 1988e:42). Most

marginelliform gastropods produce plano-convex or

hemispherical egg capsules, but in Persicula cornea (La-

marck, 1822) and Marginella goodalli (G. B. Sowerby

I, 1825) they are stalked. In most cases, each egg capsule

G. A. Coovert and H. K. Coovert, 1995

Page 65

contains a single embryo, but Pruniim apicimwi (Menke,

1828) and Dentimargo cairoma sometimes have a second

embryo, while egg capsules of Volvarina avena contain

about five embryos. Development is direct in all known

cases. All marginelliform species have a paucispiral pro-

toconch indicating direct development.

Discussion: The division of marginelliform gastropods

into two fundamentally different groups became obvious

during the course of our research. These two groups were

initially differentiated on gross radular features, but fur-

ther research revealed differences in radular structure,

anatomy, and internal columellar plications. In fact, these

two groups can be differentiated on the basis of their

internal shell whorls alone. These distinctly different

groups are the cystiscids and the true marginellids. A

brief summary of the differences follows.

Cystiscids have a shell that possesses what we have

termed cystiscid internal whorls (figures 2, 3), or modified

cystiscid internal whorls in Graniilina (figure 4). These

internal whorls are mostly resorbed by the animal, re-

sulting in extremely thin, modified whorls. The anterior

two or three columellar plications are reduced internally

to a single, axially oriented, sharp edge. Within the ap-

erture, the columella bears parietal lirae posterior to these

plications, thus the columella is multiplicate In Gran-

ulina, the posterior two columellar plications continue

internally for a whorl or two and does not have a mul-

tiplicate columella. Cystiscid radulae (figures 13-16) are

long, narrow, and C-shaped, with over 80 and up to 200

or more rachidian plates (with lateral teeth in one genus).

The narrow rachidian plates have few cusps and are

weakly to distinctly arched. The two odontophoral car-

tilages each fit into an anterolateral odontophoral carti-

lage hood (modified in Granulina). which flank the rad-

ula (figures 15, 16). The posterior end of the radula begins

as a series of nascent rachidian plates. It is encased and

secreted in a typical neogastropod radular sac, which

extends beyond the end of the odontophore (figures 15,

35).

True marginellids, on the other hand, have unmodified

internal whorls (figure 1). The plications are continuous

internally and the columella is not multiplicate. Margi-

nellid radulae (figures 17-20) are a much shorter, usually

broad, uniserial, J-shaped ribbon with 10 to 80 rachidian

plates. These moderately to very broad, flat to weakly

arched plates usually have many cusps. The subradular

membrane is weakly expanded anterolaterally into mar-

ginal cuticular flanges, which help hold the radula in

correct alignment with the underlying, anteriorly fused

odontophoral cartilages (figures 19, 20). The posterior

end of the radula begins with only one or two preformed

nascent rachidian plates which are secreted in the blind

end of a buccal pouch. The odontophore extends past

the posterior end of the radula (figure 19).

After discerning the two fundamentally different

groups of marginelliform gastropods, we searched for

possible relationships among other neogastropod fami-

lies. This revealed that all of the above cystiscid char-

acters are shared with the Olividae. Thus, we feel that

the cystiscids have a shared ancestry with that family,

and not with the Marginellidae (see final Discussion sec-

tion for more details). We have come to the conclusion

that the cystiscids and the true marginellids are not close-

ly related and must be considered as separate and distinct

families. A systematic section, outlining the classification

of these two groups, follows.

SYSTEMATICS

Key to the Families of Marginelliform Gastropods

la. Columellar plications not continuous internally,

at least anterior two columellar plications re-

duced to one nearK' axiall)- oriented columellar

edge within one full revolution internally; col-

umella usually multiplicate with 3 (rarely 2) pli-

cations, plus often up to 14 parietal lirae poste-

riorly, but may have only 1 or 2 pseudocontin-

uous plications posterior to the anterior 2 inter-

nally reduced plications; internal whorls cystiscid

or modified cystiscid type (figures 2, 3, 4) very

thin and often partially missing due to partial

resorption family CYSTISCIDAE

lb. Columellar plications continuous internally, not

reduced; columella with 2 to 6 plications, at least

the first 4 continuing internally, the 5th and 6th

often not continuing internally (and are thus

parietal lirae); internal whorls unmodified type

(figure 1) complete and of original thickness,

without evidence of resorption

family MARGINELLIDAE

Family CYSTISCIDAE Stimpson, 1865:55

Diagnosis: Shell minute to large, white, uniformly col-

ored, or patterned; surface smooth, sculptured, or axially

costate; spire flat to immersed, or low to tall; protoconch

paucispiral; lip thickened, smooth or denticulate; exter-

nal varix present or absent; siphonal notch present or

absent; posterior notch present or absent; columella mul-

tiplicate, with combined total of 2 to 13 plications -I-

parietal lirae or with 2 internally reduced columellar

plications -I- 1 or 2 pseudo-continuous plications; internal

whorls cystiscid or modified cystiscid type. Type 2, 3, or

4 animal; operculum absent. Cystiscid radula. Type 1,

2, 3, or 4. Mantle cavity with monopectinate ctenidium

and bipectinate osphradium. Proboscis pleurembolic; jaws

absent; typical radular sac present.

Remarks: The family was erected by Stimpson (1865:

55) for a single included species based on unusual features

of the head and radula. Subsequent authors included this

group in the Marginellidae. Coan (1965:186) recognized

this group as a subfamily, again based on unusual features

of the head and radula, but did not include such genera

as Gibberula (in part), Persicula. and Canaiispira. Rules

of priority require this name to be used, which is un-

fortunate because the type specimen of the type species

Page 66

THE NAUTILI'S, Vol 109, Nos. 2 & 3

is lost (Coovert, 1986d) and the species is otherwise poorly

known. We currently recognize 283 species in the Cys-

tiscidae, of which 26 are undescribed.

Key to the Recent Genera of the Family Cystiscidae

la. Distinct siphonal notch present; spire low to im-

mersed; shell minute to small 2

lb. Siphonal notch absent, anterior end rounded; spire

immersed, low, or distinctly produced; shell mi-

nute to large 3

2a. Adult shell completely lacking an external vari.x;

spire low to nearly immersed, but never com-

pletely immersed; shell usually white, uniformly

colored in some, rarely with a few narrow spiral

bands Gibberula

2b. Adult shell completely lacking an external varix

or with an obscure to distinct, raised external

varix; spire usually completely immersed, rarely

very low; shell usually uniformly colored or with

distinctive pattern of spots, tear-drop markings,

longitudinal or oblique streaks or zig-zag lines;

narrow spiral bands, if present, very numerous;

rarely white, if so, with pattern Persicula

3a. External varix completely absent; columella mul-

tiplicate, with combined total of 2 to 17 plications

-I- parietal lirae; with cystiscid internal whorls. . .4

3b. External varix present, distinct, raised, usually

strong; columella with 2 internally reduced an-

terior plications + 1 or 2 pseudo-continuous pli-

cations; with modified cystiscid internal whorls ... 8

4a. Posterior corner of aperture sharply, narrowly,

and deeply channeled; aperture usually lirate,

smooth in some; sutures often channeled or im-

pressed, if filled with callus, then still distinct;

spire distinctly produced Canalispira

4b. Posterior corner of aperture rounded, not chan-

neled; aperture smooth or denticulate, not lirate;

sutures callused over, indistinct; spire immersed

or low 5

5a. .Shell with very strong, sharply crested axial cos-

tae; strongly developed parietal callus "shield"

present; spire sunken, but not immersed, with

nuclear whorls visible; 5 columellar plications

present; lip heavily thickened, flared posteriorly,

not denticulate Extra

5b. Shell lacking strong axial costae, although may

have very weak axial growth lines or very faint

wrinkles; parietal callus wash often present, but

lacking strongly developed parietal "shield"; spire

produced or completely immersed, but not as

above; combined total of 2 to 17 columellar pli-

cations -t- parietal lirae; lip narrowly to strongly

thickened, smooth or weakly denticulate 6

6a. Suture of last whorl expanding rapidly for final

half whorl, then abruptly swept posteriorly just

before lip, giving characteristic shape to shell;

spire nearK' flat to low, not immersed; shell shape

usually cylindrical, elliptic, or oblong

Plesiocystiscus

6b. Suture of last whorl not unusually expanding or

narrowing; spire low to completely immersed;

shell shape rarely cylindrical, usually elliptic, ob-

ovate to broadly obovate, or subtriangular 7

7a. Spire usually immersed; plications usually square

in cross-section, often appearing excavated due

to collabral parietal callus ridge, which intersects

them just outside of aperture Crithe

7b. Spire usually at least slightly produced; plications

usually rounded in cross-section, rarely appear-

ing excavated, if so, not due to collabral parietal

callus ridge Cystiscus

8a. Shell usually smooth, if sculptured, very obscure;

spire immersed; shell elongate to broadly elliptic,

obovate, subtriangular, or pyriform, not cylin-

drical Granulina

8b. Shell with distinct, punctate sculpture; spire flat

to immersed; shell broadly cylindrical and

abruptly narrowed anteriorly to broadly subtrian-

gular Pugnus

Subfamily PLESIOCYSTISCINAE Coovert and Coo-

vert, new subfamily

Diagnosis: Shell minute to small, white, hyaline; last

whorl rapidly expanded then lip abruptly swept poste-

riorly giving characteristic shape; spire flat to low; lip

thickened posteriorly, smooth, lacking lirae or denticu-

lation; external varix absent; siphonal notch absent; pos-

terior notch absent; columella multiplicate with com-

bined total of 3 to 8 plications + parietal lirae; internal

whorls cystiscid type. Type 2 animal; tentacles and si-

phon moderately long; mantle translucent, in some spe-

cies not readily extending over external shell surface;

foot relatively narrow. Type 1 radula. Internal anatomy

unknown.

Genus Plesiocystiscus Coovert and Coovert, new genus

(figures 3, 21, 43)

Type species: Marginella jewettii Carpenter, 1857a;

OD, herein (figure 43).

Diagnosis: Same as for subfamily.

Description: Shell (figures 3, 43) minute to small, (adult

length 1.5-6.0 mm). Color white, hyaline; surface smooth,

glossy. Shape cylindrical, elliptic, or obovate; weakly to

distinctly shouldered. Suture rapidly descending on last

half (if last whorl, then abruptly sweeping upward just

before lip, giving characteristic shape to adult shell. Spire

nearly flat to low. Aperture moderately narrow, broader

anteriorly. Lip thin anteriorly, gradually thickening to-

ward shoulder, smooth, lacking denticulation or lirae,

external varix absent. Siphonal notch and posterior notch

ab.sent. Shell often with a weak parietal callus wash or

weak parietal callus deposits. (]olumella multiplicate, with

combined total of 3-8 plications -l- parietal lirae, usually

G. A. Coovert and H. K. Coovert, 1995

Page 67

occupying less than half to slightly more than half the

aperture length. Shell with cystiscid internal whorls.

External anatomy: 4 species studied. Type 2 animal,

with moderately long tentacles; eyes at base of tentacles;

siphon simple, moderately long; mantle translucent and

barely visible, possibly not extending over external shell

surface in some species; foot relatively narrow, about as

wide as shell and l'/2 times shell length; animal variously

marked with tiny dots of various colors, pattern of in-

ternal mantle showing through translucent shell.

Internal anatomy: Unknown.

Radula (figure 21): 3 species studied. Type 1, triserial,

with simple, thin, weak laterals that narrow to a single

cusp. Ribbon long, narrow, composed of 66-103 rows of

plates. Rachidian plates overlapping, very narrow (0.017-

0.028 mm wide), weakly arched, with 3-7 sharp cusps

on posterior edge. Anterior edge of rachidian plate slight-

ly concave, resulting in weakly U-shaped plates. Shell

length; radular width ratio (based on rachidian plate

width) = 159-200. Radular Index = 10.9-34.3.

Included species: P. abbotti (Jong & Coomans, 1988),

P. atomus (E. A. Smith, 1890), P. bubistae (Fernandes,

1987), P. cinereus (Jousseaume, 1875), P. consanguineus

(E. A. Smith, 1890), P. gutta (Gofas & Fernandes, 1988),

P. jansscni (Jong & Coomans, 1988), P. jewettii (Car-

penter, 1857), P. josephinae (Fernandes & Rolan, 1992),

P. larva (Bavay, 1922), P. palantirulus (Roth & Coan,

1968), P. polituhis (Dall, 1919), plus 4 undescribed spe-

cies.

Distribution and Habitat: Indo-Pacific (1 species), E

Pacific (4 species), western Atlantic (5 species), W. Af-

rican (6 species). Intertidal to 80 m.

Fossil Record: Eocene of France, Miocene and Pliocene

of Florida, and Pleistocene of California, to Recent.

Nomenclature, Synonymy: Photographs of the syntypes

of the type species, Marginella jewettii. were published

by Palmer (1958:pl. 24, figs. 19-21). A lectotype was

designated and illustrated in Coan and Roth (1966:pl.

51, fig. 66). The genus name is derived from the Greek

plesios. near, in the sense of plesiomorphic, of characters

near the ancestor, combined with Cystiscns. This indi-

cates the relationship to (and previous inclusion in) the

genus Cystiscus, type genus of the family. The triserial

radula, found in many other neogastropod families, but

unknown in any other cystiscids, is the most primitive

in this family. Gender masculine.

Remarks: This is the only genus of marginelliform gas-

tropods (previously referred to as "Cystiscns") with a

triserial radula. This group is considered to be the most

primitive in the family based on the radula and possession

of a generalized Type 2 animal. Radular and shell fea-

tures, including presence of odontophoral cartilage hoods,

possession of cystiscid internal whorls, and lack of an

external varix, place this group in the Cystiscidae.

Gofas and Fernandes (1988:19) described the mantle

of P. gutta (Gofas & Fernandes, 1988) as "translucent

and hardly visible when it extends over the shell." Roth

and Coan (1968:66) observed that "the mantle was never

extruded over the top of the shell of C. jewettii" and

Behrens (1984:241) likewise described the mantle as not

covering the shell in the same species. Additional obser-

vations on this point would be useful.

Subfamily CYSTISCINAE Stimpson, 1865:55

Diagnosis: Shell minute to small, white; spire immersed

to low; surface smooth or axially costate; lip thickened,

smooth or denticulate; external varix absent; siphonal

notch absent; posterior notch absent; columella multi-

plicate, with combined total of usually 2 to 8 plications

+ parietal lirae; internal whorls cystiscid type. Type 3

animal; mantle smooth, at least partially extending over

external shell surface. Type 2 radula. Internal anatomy

unknown.

t Genus Topaginella Laseron, 1957

Topaginclla Laseron, 1957:288

Type species: Marginella octoplicata Tenison-Woods,

1877 t; OD (M)

Diagnosis: Shell minute, pyriform, strongly narrowed

anteriorly; spire low; lip thickened, strongly denticulate;

external varix absent (?); distinct axial costae present;

siphonal notch absent; columella multiplicate, with com-

bined total of usually 8 plications + parietal lirae.

Fossil Record: Miocene of Australia.

Nomenclature, Synonymy: The type species, T. octo-

plicata, was figured in Cotton (1949;pl. 18).

Remarks: The type species seems to be unique. The

multiplicate columella and lack of a siphonal notch place

it near Cystiscus, but the pyriform shape, exserted spire,

denticulate lip, and fine axial costae ("striated growth

lines" of Cotton, 1949:218) render it distinct. The absence

of an external varix needs to be confirmed.

Genus Cystiscus Stimpson, 1865

(figures 9, 10, 22, 44)

Cystiscus Stimpson, 1865:55

Euliginella Laseron, 1957:282 [TS: Marginella angasi Crosse,

1870; OD]

Type species: C. capensis Stimpson, 1865 (non Mar-

ginella capensis Krauss, 1848), = Marginella cystiscus

Redfield, 1870 (nom. nov.); M (figure 44).

Diagnosis: Shell minute to small, white, hyaline; spire

immersed to low; lip thickened, smooth or weakly den-

ticulate; external varix absent; siphonal notch absent;

posterior notch absent; lacking collabral parietal callus

ridge; columella multiplicate, with combined total of

Page 68

THE NAUTILUS, Vol. 109, Nos. 2 & 3

49

Figures 43-5 1 . Shells of type species of cystiscid genera, ventral views. 43. Plesiocystiscus jeicettii (Carpenter, 1857) GAC. M1823,

Pirates C;ove San Luis Obispo, California, intertidal. Length 5 3 mm. 44. Cystiscus cystiscus (Redfield, 1870). Holotype (destroyed).

False Bay. Cape of Good Hope, South Africa, 37 m, on gorgonians. After Stimpson (1865:pl 8, fig. 2). Length 3.6 mm. 45. Cnthe

atomaria (louid. 1860. ANSP 307869, 1.6 km W.S.W, of Pulau Blankang Malti. Singapore, Malaya. Length 2.0 mm. 46. Extra

extra Jous.seaume, 1894. P. W. Clover colln., Perim, South Yemen. Length 0.9 nun. 47. Pcrsicula persicula (Linne, 1758). GAC

G. A. Coovert and H. K. Coovert, 1995

Page 69

usually 2 to 8 plications + parietal lirae, 1st plication

usually strong and raised. Type 3 animal; mantle smooth,

at least partially extending over external shell surface

Description: Shell (figure 44) minute to small (adult

length 1.0-6.0 mm). Color white, hyaline; surface smooth,

glossy. Shape rarely cylindrical, usually elliptic, obovate,

or subtriangular; weakK shouldered. Spire completely

immersed to low. Aperture narrow to broad, usually wid-

er anteriorly. Lip slightly to distincth thickened, flared

posteriorly in some species, smooth on inside edge to

weakly denticulate, lacking lirae, external varix absent.

Shell lacking a siphonal notch and posterior notch. Shell

with weak parietal callus wash or weak parietal callus

deposits in some species, but lacking collabral parietal

callus ridge. Columella multiplicate, with combined total

of 2-8 plications + parietal lirae, rarely up to 17 in which

the posteriormost are denticles; one species with onl\ 1

plication. Plications usually occupying less than half the

length of the aperture, but most of the aperture in some.

Plications excavated just inside aperture in a few species,

usually evenK rounded, first plication usualK' raised and

very strong. Shell with CNStiscid internal whorls.

External anatomy (figures, 9, 10): 13 species studied.

Type 3 animal with eyes at side of head, usually on lateral

swelling; mantle smooth, at least partially extending onto

external shell surface, in some species nearly covering

shell; foot relatively narrow, about as wide as shell and

1-1.5 X shell length; head and mantle usually uniformly

colored, often bright red, orange, or yellow, or brown or

black, internal mantle color pattern often showing through

translucent shell.

Internal anatomy: Unknown.

Radula (figure 22): 14 species studied. Type 2, uniserial,

ribbon long, narrow, composed of 80-200 plates. Ra-

chidian plates overlapping (rarely separate), very narrow

(0.009-0.036 mm wide), weakly to strongly arched, with

5-15 very strong cusps on posterior edge. Anterior edge

of rachidian plate slightly concave, resulting in crescent-

shaped plates. Shell length: radular width ratio = 94-

217. Radular Index = 10.7-25.7.

Distribution and Habitat: Neozelanic (1 species). So

Australian (21 species), Indo- Pacific (11 species), W. At-

lantic (2 species). South African (23 species). Intertidal

to 370 m.

Fossil Record: Eocene of France, Miocene and Pliocene

of Australia, to Recent.

Nomenclature. Synonymy: The type species C. cijstis-

cits is discussed and figured in Coovert (1986d) and here-

in (figure 44). The radula of this species, along with three

others, was reviewed in Coovert (1989b:5-6). SEMs of

an additional eight Australian species (Dean Hewish,

pers. comm), including C. angasi, the type species of

Eitliginella. have been examined by us. All these species

are considered congeneric based on similar radulae and

shell characters. Euliginclla was considered synonymous

by Coan (1965:190).

Remarks: Originally described in its own family by

Stimpson, apparently based on the radula and unusual

features of the head, this genus again becomes the nom-

inate genus of a now more clearly defined family.

Genus Crithe Gould, 1860

(figure 45)

[TS: M. nipponica Habe. 1951;

Crithe Gould, 1860:384

Microviilina Habe, 1951:105

OD]

Type species: C. atomaria Gould, 1860; M (figure 45)

Diagnosis: Shell minute to small, white, hyaline; spire

usually immersed, rarely low; lip thickened, smooth,

lacking denticulation; external varix absent; siphonal

notch absent; posterior notch absent; columella multi-

plicate, with combined total of 6-8 plications -I- parietal

lirae, plications usually excavated inside aperture due to

collabral parietal callus ridge. Presumably a Type 3 an-

imal.

Description: Shell (figure 45) minute to small (adult

length 1.6-3.2 mm). Color white, hyaline; surface smooth,

glossy. Shape narrowly to broadly obovate or subtrian-

gular; weakly to moderately strongly shouldered. Spire

usually completely immersed, rarely low. Aperture nar-

row, wider anteriorly. Lip moderately thickened, es-

pecially posteriorly, thinner anteriorly, smooth on inside

edge, lacking denticulation and lirae, external varix ab-

sent. Shell lacking a siphonal notch and posterior notch.

Shell usually with a distinct collabral parietal callus ridge

that intersects plications. Columella multiplicate, with

combined total of 6-8 plications -t- parietal lirae, usually

occupying about two-thirds the aperture length. Plica-

tions usually appearing excavated just inside aperture

due to callus deposits from anterior end of collabral pa-

rietal callus ridge. Shell with cystiscid internal whorls.

External anatomy: Presumably a Type 3 animal; man-

tle extension undetermined; generally uniformly colored,

bright yellow, rose, or orange-red, or black with 2 zones

of yellow on internal mantle.

M1558, Dakar, Senegal, West Africa, in sand, low tide. Length 20.0 mm. 48. Gibherula onjza (Lamarck, 1822). Neotype, MHNG,

Senegal. After Gofas (1990:f)g. 2). Length 6,5 mm. 49. Canalispira olivellaeformis Jousseaume, 1875. Lectotype, MHNP, locality

unknown. After photos in Kilburn (1990:fig. 5). Length 4,2 mm, 50. Granulina isseli (G. & H, Neviii, 1875), GAG Ace. # 53-87,

Red Sea coast of Egypt between El Ghurdaga and 80 km south of El Quseir. Length 1.6 mm 51. Ptignus parvus Hedley, 1896.

GAG M2135, east of North Ft., Flinders Island, Bass Strait, Australia, 39°43'S, 148°36'E. Length 2.0 mm.

Page 70

THE NAUTILUS, Vol. 109, Nos. 2 & 3

Internal anatomy: Unknown.

Radula: 1 species studied (3 specimens). Type 2, uni-

serial, ribbon long, narrow, composed of 189-220 plates.

Rachidian plates overlapping, very narrow (0.007-0.008

mm wide), weakly arched, with 4 strong cusps on pos-

terior edge. Anterior edge of rachidian plate slightly

concave, resulting in an asv mmetrically V-shaped plate.

Shell length: raduiar width ratio = 234-270. Radular

Index = 47.2-55.0.

Distribution and Habitat: Indo-Pacific (6 species). In-

tertidal to 58 m.

Fossil Record: Pliocene of Australia to Recent.

Nomenclature, Synonymy: The type species of Crithe

and Micwviilina, and their generic synonymy, were dis-

cussed in Coovert (1986e and 1987c). The holotype of

C. aiomaria was figured in Johnson (1964:pl. 8, fig. 12)

and Coovert (1986e;fig. 1).

Remarks: The columellar plications, which are square

in cross-section and excavated due to the collabral pa-

rietal callus ridge, and the usually immersed spire, serve

to differentiate this group. The radulae are diagnostic.

Genus Extra Jousseaume, 1894

(figure 46)

Extra Jousseaume, 1894:98, 101

Type species: £. extra Jousseaume, 1894; M (figure 46)

Diagnosis: Shell minute, white, semitranslucent; prom-

inent axial costae present; spire sunken but not immersed;

lip strongly thickened, smooth, lacking denticulation,

flared posteriorly; siphonal notch absent; posterior notch

absent; distinct parietal callus "shield" present; columella

multiplicate, with combined total of 5 plications + pa-

rietal lirae, plications slightly excavated inside aperture

due to parietal callus deposits.

Description: Shell (figure 46) minute (adult length 1.3-

1.4 mm). Color white, body whorl semitranslucent, sur-

face with 17-20 prominent, sharply crested axial costae

Shape obovate, truncate posteriorly; strongly shouldered

Spire sunken but not immersed. Aperture narrow, slight-

ly wider anteriorly. Lip very strongly thickened, es-

pecially posteriorly, arched above apex and flared pos-

teriorly, smooth on inside edge, lacking denticulation and

lirae, apparently lacking an external varix. Shell lacking

a siphonal notch and posterior notch. Shell with a distinct

parietal callus "shield" that is free anteriorly with slight

chink behind Columella multiplicate, with combined

total of 5 ])licati(jns 4 parietal lirae occupying about %

the aperture length. Plications slightly excavated just in-

side aperture due to callus deposits from the parietal

callus shield. Internal whorls unknown but presumed to

be cystiscid type.

External anatomy: Unknown.

Internal anatomy: Unknown.

Radula: Unknown.

Distribution and Habitat: Indo- Pacific (1 species), re-

corded only from the western Indian Ocean. Apparently

shallow water.

Fossil Record: K.nown only from the Recent.

Nomenclature, Synonymy: See Coovert (1987d) for dis-

cussion.

Remarks: This monotypic genus is provisionally placed

in the Cystiscinae based on the 5 excavated plications

and lack of labial denticulation. The last strong axial

costa has the appearance of an external varix, which, if

present, might eliminate this genus from the Cystiscinae.

The slightly sunken but non-immersed spire and the

parietal "shield of this group appear to be unique.

Subfamily PERSICULINAE Coovert and Coovert, new

subfamily

Diagnosis: Shell minute to large, white, uniformly col-

ored, or patterned; spire immersed or low to tall; lip

thickened, smooth or Urate; external varix present or

absent; siphonal notch present or absent; weak to distinct

posterior notch present; columella multiplicate with

combined total of 3-13 plications -I- parietal lirae; in-

ternal whorls cystiscid type. Type 4 animal; tentacles

short to long, rarely absent; siphon short to long; eyes

situated laterall> on head slightly below the base of the

tentacles; mantle usually not extending over external shell

surface. Type 3 radula. Odontophoral cartilages separate;

valve of Leiblein present, with or without bypass tube;

esophageal caecum absent; gland of Leiblein short, small,

emptying directly into posterior end of esophagus; paired

salivary glands ascinous or tubular, attached to esophagus

just anterior to the valve of Leiblein, ducts attached to

walls of esophagus; single accessory salivary gland pres-

ent, ascinous or tubular; anal gland absent.

Genus Persicula Schumacher, 1817

(figures 2, 11-16, 23, 35-37, 47)

I'crsicula Scliunuiclier, 1817:235

Piwhijhathrun Claskoiii, 185;3:356[TS: P. rnarginclloidcuni Gus-

koiii, 185.3, = Erato cijpracuidcs il. B .\(lanis, 1845; SD

(MlTryon, 1885:270]

Pcrsicida, suhg Ralncca Gray, 1857:37 [TS: /'. inlerrupta (La-

marck, 1822), = Marginella intcrrupta Lamarck, 1822,

= Volulci iulcrruplolineata Megerle von Miihifeld, 1816;

Ml

Type species: P. variabilis Schumacher, 1817, = Voluta

persicula Linne, 1758: M (figure 47)

Diagnosis: Shell small to large, uniformly colored, or

white to colored and patterned, nuxlerately thick to thick-

shelled; spire usualK inunersed; lip thickened, vveakK to

strongly lirate; external varix present or absent; distinct

siphonal notch present; posterior notch present; colu-

mella nniltiplicatc with combined total of 4-13 plications

G. A. Coovert and H. K. Coovert, 1995

Page 71

+ parietal lirae. Type 4 anitnai; teiitatles long; siplion

usually long; mantle apparently not extending over ex-

terna! shell surface.

Description: Shell (figures 2, 47) small to large (adult

length 4.5-34 0 mm). CJolor white, tan, pinkish- or yel-

lowish-tan, to gray, with pattern of spiral lines, rows of

dots or tear-drop shaped markings, or longitudinal wavy

or zig-zag markings which often form triangular peaks,

less often uniformly Desh-colored or nearly white due to

pattern reduction; surface smooth, glossy, few species

with weak axial costae. Shape narrowly to broadly ob-

ovate, or narrowly to broadly elliptic; weakly to mod-

erately strongly shouldered. Spire usually completely im-

mersed, rarely slightly produced. Aperture narrow,

slightly wider anteriorly Lip moderately thickened, es-

pecially posteriorly, thinner anteriorly, very weakly to

strongly lirate, completely lacking an external varix or

with strong to weak external varix Shell with a distinct

siphonal notch and a very weak to very distinct posterior

notch. Moderately-thick to thick-shelled, some species

with distinct parietal or apical callus deposits, a few with

a very strong parietal callus "shield." Columella multi-

plicate, with combined total of 4-13 plications + parietal

lirae usually occupying more than half the aperture

length. Shell with c\stiscid internal whorls.

External anatomy (figures 11, 12): 15 species studied.

Type 4 animal with moderately long to long tentacles;

siphon simple, moderately short to usually very long;

mantle apparently not extending over external shell sur-

face; foot narrow to moderately broad, about as wide as

shell to considerably broader and about IVz times shell

length; animal variously marked with tiny dots, spots,

blotches, flecks, or streaks of opaque white, black, or

various colors.

Internal anatomy (figures 35-37): 2 species studied. As

in family diagnosis. Valve of Leiblein without bypass

tube, paired salivary glands ascinous, single accessory

salivary gland ascinous.

Radula (figures 13-16, 23): 14 species studied. Type 3,

uniserial, ribbon long, narrow, composed of 80-209 plates.

Rachidian plates overlapping, narrow (0.011-0.093 mm

wide), moderately to strongly arched, with 6-14 sharp

cusps on posterior edge, the central cusp usually the

strongest. Anterior edge of rachidian plate strongly con-

cave, resulting in \J- or V-shaped plates. Shell length:

radular width ratio = 146-572. Radular Index = 6.7-

21.8.

Distribution and Habitat: S. Australian (3 species), Indo-

Pacific (11 species), E. Pacific (7 species), W. Atlantic

(20 species), W. African (9 species), South African (1

species).

Fossil Record: Eocene of France and Alabama, upper

Oligocene and Miocene of W. Atlantic, to Recent. In-

tertidal to 370 m.

Nomenclature, Synonymy: The type species of Rabicea,

V. interrtiptolineata, is congeneric with P. persicula,

based on similar shell, radular (Coovert, 1989b:22), and

external anatomical features. {]oan and Roth (1966282)

discussed this synonymy. Pachybathron has been treated

as a valid genus by many recent authors, including Coo-

mans (1972), who discussed the group, its early confused

familial placement, the type species, and its synonymy.

The cited example in Adams and Adams (1853:194) does

not constitute a type fixation. Tryon (1885:270) thus pro-

vided the earliest type fixation by subsequent designa-

tion. The features of the external anatomy, radula, and

the brown triangular markings of the shell indicate a

close relationship to some of the other small western

Atlantic Persicuta. The parietal callus shield is considered

to be no more than a species-group character. Thus,

Pachybathron is here treated as a synonym. Subgeneric

recognition may eventually be justified, but this would

necessitate formal recognition of other species groups, a

step considered unwise at this time.

Remarks: Erroneously reported to have a Type 2 animal

(Coovert, 1987g, 19871) based on strictly dorsal views,

closer examination of several species revealed the split

head typical of a Type 4 animal (figure 11). The two

halves of the head are capable of closing together at will,

thus appearing unsplit. The long siphon usually distin-

guishes this group anatomically from the closely related

Gibberiila. The shell of Persicula is usually patterned,

often has a distinct external varix, and the spire is usually

immersed. These conchological features serve to separate

the two groups and are presented in the key to Recent

genera. See Coovert (1987b) for a discussion of the Per-

sicula cornea group.

Genus Gibberula Swainson, 1840

(figures 48, 79)

Gibberula Swainson, 1840:323

Grantila Jousseaume, 1875:167, 246 [TS: C. bei^soni (Reeve,

1865), = Marginclla bcnsoni Reeve, 1865; SD (M) Coan,

1965: 190]

Kogoinea Habe, 1 951:) 03 [TS: Marginclla nuvenipruvincialis

(Yokoyama, 1928), = Erato ii(>vcniprovincialis\oko\dma.,

1928; OD]

t Dcntiginella Laseron, 1957:288 [TS: Marginclla rnctula Cot-

ton, 1949 t; OD (M)]

Epiginella Laseron, 1957:279 [TS: £. ablita Laseron, 1957; OD]

t Lataginella Laseron, 1957:288 [TS: Marginclla hitsoni Chap-

man, 1921 t; OD (M)]

Phyloginella Laseron, 1957280 [TS: P. coinprcssa Laseron,

19.57; OD]

t Vetaginclla Laseron, 1957:288 [TS: Marginclla donia Cotton,

1949 t;OD{M)]

Dilnculum Barnard, 1962:14 [TS: D. inopinaliun Barnard, 1962,

= Marginclla bcnsoni Reeve, 1865, OD]

Type species: G. zonata Swainson, 1840, = Vulvaria

oryza Lamarck, 1822; M (figure 48)

Diagnosis: Shell minute to medium size, white or col-

ored, some species with spiral bands; spire low to medium

height, not immersed; lip thickened, weakly to strongly

Page 72

THE NAUTILUS, Vol, 109, Nos, 2 & 3

lirate, smooth in some species; external varix absent; dis-

tinct siphonal notch present; posterior notch present but

often weak; columella multiplicate with combined total

of 3-8 plications + parietal lirae. Type 4 animal; ten-

tacles short; siphon short; mantle not usually extending

over external shell surface.

Description: Shell (figure 48) minute to medium in size

(adult length 1.8-8.5 mm). Color usually white, hyaline,

less commonly tan, yellow, or orange, some species with

spiral lines or bands or sutural band, rarely with oblique

markings; surface smooth, glossy, rarely with fine, weak

axial costae. Shape narrowly subcylindrical to usually

elliptic, obovate, or subpyriform; weakly to strongly

shouldered, some species with weakly raised sutures. Spire

low to medium height, not immersed. Aperture narrow

to moderately broad, wider anteriorly. Lip moderately

thickened, especially posteriorly, thinner anteriorly, usu-

ally weakly to strongly lirate but absent in some species,

completely lacking an external varix. Shell with distinct

siphonal notch and very weak to distinct posterior notch.

Shell with apical callus deposits in some species, rarely

with elongate collabral parietal callus deposits. Colu-

mella multiplicate, with combined total of 3-8 plications

+ parietal lirae occupying Va to nearly the full aperture

length. Shell with cystiscid internal whorls.

External anatomy: 26+ species studied. Type 4 animal,

with short tentacles, which are rarely absent; siphon sim-

ple, short, barely extending above the dorsal edge of the

siphonal notch; mantle apparently not extending over

external shell surface or with a small lobe on the left side

in some species; foot narrow to moderately broad, about

as wide to slightly wider than shell and usually about IVi

times shell length; some species with anterolateral edges

of foot weakly to strongly raised to form "parapodia ;

animal variously marked with tiny dots, spots, or blotch-

es, but not striped or lineated, markings of opaque white,

brown, yellow, orange, or green.

Internal anatomy (figure 79): 1 species studied. As in

family diagnosis. Valve of Leiblein with bypass tube;

paired salivary glands tubular; single accessory salivary

gland tubular.

Radula: 21 species studied. Type 3, uniserial, ribbon

long, narrow, of 80-153 plates. Rachidian plates over-

lapping, narrow (0.008-0.042 mm wide), moderately to

strongly arched, with 5-11 sharp cusps on posterior edge.

The central cusp is usually the strongest. Anterior edge

of rachidian plate strongly concave, resulting in U- or

V-shaped plates. Shell length: radular width ratio = 118-

309. Radular Index = 9.1-21.9.

Distribution and Habitat: Neozelanic (2 species), S. Aus-

tralian (4 species), Indo-Pacific (26 species), E. Pacific (6

species), W. Atlantic (9 species), Mediterranean (10 spe-

cies), W. African (25 species). South African (7 species).

Intertidal to 3,300 m.

Fossil Record: Paleocene and Eocene of France and

W. Atlantic, Miocene and Pliocene of Italy, Australia,

W. Atlantic, and Japan, and Pleistocene of W. Atlantic

and California, to Recent.

INomenclature, Synonymy: Gibberula onjza. type spe-

cies of this genus, is valid and distinct from G. miliaria

(Linne, 1758). Coan (1965:189) (questionably) and Dodge

(1955:79-82) considered these two species synonymous.

But Gofas (1990:117) clearly differentiated the two spe-

cies based on shell and external anatomical characters,

and designated a neotype (figure 48) and lectotype re-

spectively. The geims Granula and its type species M.

bensoni differ from Gibberula (type species G. oryza)

only in size and very minor, species-level shell characters.

Granula was synonymized under Gibberula by Coss-

mann (1899:96) and Wenz (1943:141). Kogomea, like-

wise differing only in minor characters, was synonymized

by Roth and Coan (1971:579) and Rehder (1980:85).

Diluctilum was discussed and synonymized by Kilburn

and Rippey (1982:113, 214). Barnard (1962) described

Dibicubin} and its type species simply because the radula

was known, whereas other species "are merely names

given to the shells of molluscs whose anatomy is unknown

(op. cit., p. 14)." This is clearly untenable. The wishbone-

shaped radula was an error in observation, likewise made

by Ponder (1970:70, fig. 3F). This error was discussed in

Coovert (1989b:12). Gofas (1989a) correctly figured a

number of radulae from Gibberula, drawn from SEMs.

The five Laseron (1957) genera were all based on species-

level characters, and are here synonimized with Gibber-

ula. The t\pe species of all these genera show a distinct

siphonal notch. The type species of Phyloginella was

figured by Kaicher (1992: #6211). Powell (1979:222)

synonymized Lataginella under Kagomea (sic).

Remarks: Gibberula cucullata Gofas and Fernandes,

1988, with strongly developed "parapodia," has a highly

modified head. The bifurcated tips of the head taper into

divergent anterior lobes. These tips can be interpreted

as being either the anterior lobes of the head and thus

lacking tentacles, or the tentacles themselves and lacking

the anterior lobes. Based on the sharply pointed tips and

relative position of the eyes, we interpret these tips as

being the anterior lobes of the head, the animal lacking

tentacles. Gofas and Fernandes (1988:25) suggested that

this species "may deserve separate generic status." Gib-

berula suhbulbosa (Tate, 1878) (based on unpubl. data

ex. R. Burn and D. Hewish) has a similarly constructed

head but apparently lacks the strongly developed "par-

apodia." Until the external anatomy of the t>pe species

of all related genus-group names are known, it is con-

sidered unwise to place these species in a different genus-

level group.

This large, widely distributed group is well charac-

terized by radular, external anatomical, and concholog-

ical characters. See Persicula for additional discussion.

Genus Canalispira Jousseaume, 1875

(figure 49)

(.'(luaiisiiiru JoiisseaiiiiR', 187.5:168, 270

G. A. Coovert and H. K. Coovert, 1995

Page 73

Baroginella Laseron, 1957:286-287 [TS: B. infirma Laseron,

1957; OD]

Type species: C. olivellaeformis Jousseaume, 1875; M

(figure 49)

Diagnosis: Shell minute to medium size, white, hyaline,

rarely with faint pattern, cylindrical-biconic to obovate

or obconic; sutures usually impressed to channeled; spire

medium to tali; lip thickened, smooth or iirate; external

varix absent; siphonal notch absent; deeply channeled

posterior notch present; columella multiplicate with

combined total of 3-6 plications + parietal lirae. Mod-

ified Type 4 animal; tentacles absent; siphon not visible;

foot split anteromedially, covering external shell surface

anterolaterally.

Description: Shell (figure 49) minute to medium in size

(adult length 2.2-10.3 mm). Color usually white, hyaline,

rarely with faint brown maculations, fine oblique lines,

or weak axial streaks; surface smooth, glossy. Shape cy-

lindrical-biconic to narrowly obovate or obconic; usually

very weakly to weakly shouldered. Sutures distinctly to

weakly channeled, deeply impressed, or completely cal-

lused over. Spire medium to tall. Aperture narrow to

moderately broad, wider anteriorly. Lip moderately

thickened, especially posteriorly, thinner anteriorly, out-

er edge incurved on posterior half, usually weakly to

strongly Urate but smooth and lacking lirae in some spe-

cies, completely lacking an external varix. Shell lacking

a siphonal notch but with a sharply, narrowly, and deeply

channeled posterior notch. Parietal callus wash usually

confined to aperture. Columella multiplicate, with com-

bined total of 3-6 plications -f- parietal lirae occupying

half or more of the aperture length. Shell with cystiscid

internal whorls.

External anatomy: 2 species studied. Modified Type 4

animal; tentacles absent; triangular lobes of head pointed

anteriorly; the small black eyes medially placed on each

triangular lobe; siphon not visible; mantle extension un-

determined; foot narrow, about as wide and long as the

shell, split anteromedially, covering the anterior end of

the shell laterally like "parapodia ; animal white or col-

orless. (Largely based on unpubl. notes and drawings of

an undescribed W. Australian species, ex. P. W. Clover).

Internal anatomy: Unknown.

Radula: 2 species studied. Type 3, uniserial, ribbon long,

narrow, composed of 121-136 plates. Rachidian plates

overlapping, narrow (0.016-0.033 mm wide), moderate-

ly to strongly arched, with 7 sharp cusps on posterior

edge. The central cusp is usually the strongest. Anterior

edge of rachidian plate strongly concave, resulting in U-

or V-shaped plates. Shell length: radular width ratio =

176-270. Radular Index = 17.3-19.4.

Distribution and Habitat: So. Australian (1 species),

Indo-Pacific (7 species), W. Atlantic (3 species). South

African (2 species). Intertidal to 540 m.

Fossil Record: Pliocene of Australia to Recent.

Nomenclature, Synonymy: The lectotype of C. olivel-

laeformis was figured by Kilburn (1990:fig. 5) and is

illustrated here (figure 49). We studied the holotype of

B. infirma and it is clearly congeneric, differing only in

the degree of channeling of the sutures. We regard this

character to be of species-level and synonymize Baro-

gijiella with Canalispira.

Remarks: The modified Type 4 animal and lack of a

siphonal notch render this group distinct, but the Type

3 radula clearly places this genus in the Persiculini. More

study is needed of living animals.

Subfamily GRANULININAE Coovert and Coovert, new

subfamily

Diagnosis: Shell minute to small, white, hyaline; surface

smooth or sculptured; spire flat to immersed; lip thick-

ened, smooth or denticulate; external varix present; si-

phonal notch absent; columella with 2 internally reduced

columellar plications, plus 1 or 2 pseudo-continuous pli-

cations; internal whorls modified cystiscid type. Type 2

animal; tentacles long, slender; siphon moderately long;

mantle pustulose, extending over external shell surface;

foot long, narrow. Type 4 radula. Internal anatomy un-

known.

Genus Granulina Jousseaume, 1888

(figures 4, 24, 50)

Granulina Jousseaume, 1888:191

? f Cryptospira (Cyprafo/ina) Cerulli-Irelli, 1911:231 [TS: Val-

uta clandestina Brocchi, 1814 f; M]

Mcrovia Dall, 192186 [TS: Merovia pyriformis (Carpenter,

1865), = Volutclla pyriformis Carpenter, 1865, = ? Mar-

ginella margaritula Carpenter, 1857b; M)

Microginella Laseron, 1957:280 [TS: Marginella anxia Hedley,

1909; OD]

Gibberulina audi., non Monterosato, 1884

Type species: Marginella pygmaea Issel, 1869 {non

Marginella pygmaea G. B. Sowerby II, 1846), = Mar-

ginella isseli G. & H. Nevill, 1875 (nom. nov.); M (figure

50)

Diagnosis: Shell minute to small, white, hyaline; spire

immersed; lip strongly thickened, usually denticulate;

external varix present; siphonal notch absent; collabral

parietal callus ridge often present; columella with 4 strong

plications that are often excavated inside aperture. Type

2 animal; tentacles long, slender; siphon moderately long;

foot long, narrow; mantle pustulose, extending over ex-

ternal shell surface.

Description: Shell (figures 4, 50) minute to small (adult

length 0.8-3.2 mm). Color white, hyaline; surface smooth,

glossy, rarely with fine, weak axial costae or minutely

pitted. Shape narrowly to broadly elliptic, obovate, sub-

triangular, or pyriform; very weakly to weakly shoul-

dered. Spire usually immersed. Aperture narrow to mod-

erately laroad, noticeably wider anteriorly and slightly

wider posteriorly. Lip moderately to greatly thickened,

Page 74

THE NAUTILUS, Vol. 109, Nos. 2 & 3

especially posteriorly, usually distinctly denticulate, rare-

ly smooth, with distinct external varix. Shell lacking a

siphonal notch; posterior notch usually absent, rarely

weakly developed. Shell usually with a weak to distinct

collabral parietal callus ridge that often intersects colu-

mellar plications. (Columella with 4 strong plications, third

and fourth rarely weak, rarely with weak parietal den-

ticles posterior to plications. Plications often appearing

excavated inside aperture due to callus deposits from

anterior end of collabral parietal callus ridge. Plications

occujjying halt or less the aperture length. Shell with

modified cysti.scid internal whorls.

External anatomy: 13 species studied. Type 2 animal,

with long, slender tentacles; eyes at base of tentacles;

siphon simple, moderately long; mantle pustulose, ex-

tending symmetrically over most of external shell sur-

face; foot narrower to slightly wider than shell and about

twice the shell length; animal variously marked with tiny

dots, spots, or areas of concentrated pigmentation, mark-

ings usually black, white, or yellow, but may be orange

or turquoise.

Internal anatomy: Unknown.

Radula (figure 24): 12 species studied. Type 4, uniserial,

ribbon long, narrow, composed of 90-159 plates. Ra-

chidian plates usually completely non-overlapping, very

narrow (0.006-0.012 mm wide), moderately arched, with

9 to 12 raised, clustered denticles located in staggered

positions on dorsal surface, with a single (or asymmet-

rically paired) strong central cusp protruding posteriorly.

Anterior edge of rachidian plate weakly concave, re-

sulting in vaguely V-shaped plates. The rachidian plates

are asymmetrical and alternate as mirror images. Shell

length: radular width ratio = 164-351. Radular Index =

8.2-13.7.

Distribution and Habitat: So. Australian (3 species),

Indo- Pacific (13 species), E. Pacific (3 species), W. At-

lantic (12 species), Mediterranean (8 species), W. African

(3 species), South African (3 species). Intertidal to 1,285

m (single record at 1,700 m).

Fossil Record: Miocene to Pliocene of W. Atlantic,

Pliocene ol Italy, Pleistocene of Japan and California, to

Recent.

Nomenclature, Synonymy: The type species and ge-

neric synonymy oi Granulina was discussed in Coovert

(1987h). Although the "problem of Granulina clandes-

tina" was treated by Gofas (1992:5-6), with a neotype

designated and figured (op. cit., fig. 3), several discre-

pencies remain. Brocchi (1814:642-643, pi. 15, fig. 11)

described and figured a shell with 3 cohmiellar plications,

a slightly exserted spire, and a pyriform shape. This was

considered to be a juvenile 1)\ ( iofas ( 1 992:5), but juvenile

Granulina have four plications, an immersed spire, and

an ovate or globose shape. More study is needed to resolve

these discrepencies. The lectotype of V, pyrijormis. type

species of Merovia, was figured by Cloan and Roth (1966:

pi. 51, fig. 77). Further research is needed to clearly

establish the synonymy of this species with M. margar-

itula. We have studied syntypes ol M . anxia, type species

of Microginella, and regard them to be referable to

Granulina.

Remarks: Absence of a siphonal notch and presence of

a distinct external varix distinguish this genus from all

other cystiscid genera except Piignus. Shell shape and

usual lack of sculpturing will further characterize Gran-

ulina. The very distinct Type 4 radula, modified cystiscid

internal whorls, and Type 2 animal clearly distinguish

these two genera as a separate subfamily. For a discussion

of G. hadria (Dall, 1889), see Coovert (1988a).

Genus Pugnus Hedley, 1896

(figure 51)

Pugnus Hedley, 1896:105-106

Mar^incUopsis Ravav, 1911:241 [TS: M serrei Bavay, 1911;

M]

Type species: P. parvus Hedley, 1896; OD (M) (figure

51)

Diagnosis: Shell minute, white, hyaline; surface mi-

nutely sculptured; spire flat to immersed; lip thickened,

smooth or denticulate; external varix present; siphonal

notch absent; columella with 3 or 4 plications. Type 2

animal; tentacles long, slender; siphon moderately long;

foot long, narrow; mantle pustulose, extending over ex-

ternal shell surface.

Description: Shell (figure 51) minute (adult length 0.75-

2.0 mm). C>olor white, hyaline; surface sculpture of ex-

tremely fine, crossed spiral and axial threads leaving rows

of quadrate or hexagonal pits in between, or with spiral

or axial rows of beads or short, dashed ridges which are

often serpentine or interconnected. Shape broadly cylin-

drical to broadly subtriangular; weakly to strongly nar-

rowed anteriorly, strongly shouldered. Spire flat to im-

mersed. Aperture moderately narrow to broad, wider

anteriorly and slightly wider posteriorly. Lip strongly

thickened, especially posteriorly, distinctly denticulate

or smooth, with distinct external varix. Shell lacking a

siphonal notch and posterior notch. Often with a distinct

parietal callus wash extending out of aperture. Columella

with 3-4 plications, third and fourth often weak. Pli-

cations occupying less than hall the aperture length. Shell

with modified cysti.scid internal whorls.

External anatomy: I species studied. Type 2 animal,

with long, slemler tentacles; eyes at base of tentacles;

siphon simple, moderately long; mantle pustulose, ex-

tending symmetrically over most ot external shell sur-

face; fool narrower than shell and about twice the shell

length; animal variously marked with tiny reddish dots

and si)ots on a translucent background. [Based on pers.

obs. of pliotographs ot /'. serrei (Bavay, 1911), ex. Colin

Redfern].

Internal anatomy: Unknown.

G. A. Coovert and H. K. Coovert, 1995

Page 75

Radula: 2 species studied. Type 4, uniserial, ribbon long,

narrow, of >124 plates. Racliidian plates usually com-

pletely non-overlapping, very narrow (0.006 mm wide),

moderately arched, with 11-12 raised, clustered denti-

cles located in staggered positions on dor.sal surface, with

a single (or as> mmetricaliy paired) strong central cusp

protruding posteriorly. Anterior edge of rachidian plate

weakly concave, resulting in vaguely V-shaped plates.

The rachidian plates are asymmetrical and alternate as

mirror images. Shell length: radular width ratio = 161-

242. Radular Index = 10.3.

Distribution and Habitat: S. Australian (1 species), Indo-

Pacific (1 species), W. Atlantic (2 species). Intertidal to

60 m.

Fossil Record: Known only from the Recent.

Nomenclature, Synonymy: Marginellopsis is here syn-

onymized with Piignus on the basis of very similar rad-

ulae (pers. obs. of SEMs, ex. Dean Hewish) and similar

shell sculpturing between the type species of the two

groups. An undescribed species from Brazil has char-

acters of both groups, and P. maesae Roth, 1972 is in-

termediate in shell shape. Special attention should be

paid to the distinctive surface sculpturing uniting them

all. These differences are provisionally considered to be

of specific value only. See Roth (1972:107) for remarks

on differing familial placement.

Remarks: For general comments, see remarks under

Granulina.

Family MARGINELLIDAE Fleming, 1828:328

Diagnosis: Shell minute to very large, white, uniformly

colored, or patterned; protoconch paucispiral; lip thick-

ened, smooth or denticulate; external varix present or

absent; siphonal notch present or absent; columella with

2-6 plications; internal whorls unmodified type. Type 1

or Type 2 animal; operculum absent. Marginellid radula.

Type 5, 6, modified Type 6, Type 7, 8, or 9, or non-

radulate. Mantle cavity with monopectinate ctenidium

and bipectinate osphradium. Proboscis pleurembolic; jaws

absent; marginellid buccal pouch present, absent in non-

radulate species; odontophoral cartilages fused anteriorly

or both anteriorly and posteriorly; valve of Leiblein pres-

ent or absent, with or without bypass tube; esophageal

caecum present or absent; gland of Leiblein large, sac-

culate, and emptying directly into posterior end oi esoph-

agus, or with long, convoluted duct and a terminal bulb,

emptying either into posterior end of esophagus or pass-

ing through nerve ring and emptying into anterior end

of proboscis; paired salivary glands ascinous or tubular,

either attached to esophagus just anterior to the valve of

Leiblein, contained within proboscis, or free, ducts either

embedded in or attached to walls of esophagus, or free;

single accessory salivary gland present or absent, ascinous

or tubular; anal gland present.

Remarks: We currently recognize a total of 483 species

in the Marginellidae, of which 51 remain to be described.

The two subfamilies were recognized in Coan (1965) but

were not clearlv based on anatomical characters.

Key to the Recent Genera of the Family Marginellidae

la. Adult shell very large, over 50 mm in length;

siphonal notch present, broad, weak to distinct;

spire medium height, with large, often some-

what bulbous protoconch; lip narrowly thick-

ened, smooth, without denticulation 2

lb. Adult shell minute to large, rarely very large,

if over 50 mm, then spire immersed or with

strongly thickened, denticulate lip; spire im-

mersed to tall, protoconch not unusually large;

lip narrowly to strongly thickened, with or with-

out denticulation 3

2a. Columella with 4 very strong, nearly transverse

plications; 1st plication well posterior of anterior

border of columella; with very distinct parietal

callus deposit posteriorly; shell often somewhat

dull and crazed or blistered Afrivoluta

2b. Columella with 2 strong, very oblique plica-

tions; 1st plication bordering anterior end of

columella; without distinct parietal callus de-

posits; shell very smooth, glossy . . . Marginellona

3a. Columella with 4 strong plications occupying

more than half the aperture length; if with weak

"false 5th" plication, then with distinct poste-

rior notch; spire low, medium, or tall; external

varix distinct; often with weak to distinct si-

phonal notch present 4

3b. Columella with 2-6 plications, those with 4 or

fewer plications occupying half or less the ap-

erture length; spire immersed, or low, medium,

or tall; external varix distinct, weak, or absent;

siphonal notch usually absent, if present, with

more than 4 plications 12

4a. Shell with distinct to weak siphonal notch 5

4b. Shell completely lacking a siphonal notch, the

anterior end rounded, or notch very weak and

obscure 8

5a. Shell usually white or uniformly colored yel-

lowish-white to \ellowish-orange, rarely with

several obscure pale yellow spiral bands; Aus-

tralian or Neozelanic distribution 6

5b. Shell usually with distinct and colorful pattern

of spots or dots, numerous spiral lines, mottled,

or with axial streaks or lines, or various com-

binations; eastern Atlantic or western Indian

Ocean distribution 7

6a. Posterior notch present, sharply defined; lip

rapidly thickening posteriorly, narrowed slight-

ly then widening again at posterior Vr. lip smooth,

not denticulate; body whorl generally pyriform,

strongly narrowed at anterior Vi; usually with

heavy ventral callusing; spire whorls convex or

shouldered, giving uneven profile; posterior end

Page 76

THE NAUTILUS, Vol. 109, Nos. 2 & 3

of lip joins body whorl well below suture above;

4th plication not distinctly remote . . Amtroginella

6b. Posterior notch weak to absent; lip thickest me-

dially, uniformly thickened throughout; lip

smooth or weakly denticulate; body whorl ob-

ovate, usually straight to convex at anterior Vi;

spire whorls evenly contoured, essentially

straight in profile; posterior end of lip joins body

whorl at or slightly below previous suture; 4th

plication often remote (i.e. farther separated

than other 3) Mesoginella

7a. Shell with distinct to obscure axial costae; ap-

erture usually broadest medially; lip strongly

denticulate Glabella

7b. Shell smooth, completeK lacking axial costae;

aperture usually broadest anteriorly; lip smooth

or denticulate Marginella (part)

8a. Lip flared at shoulder, weakly to strongly alate;

shell usually strongly narrowed anteriorly 9

8b. Lip not notably flared or alate; shell not usually

strongly narrowed anteriorly 10

9a. With a strongly marked posterior notch in cor-

ner of aperture; columellar plications appear

excavated inside aperture due to callus deposits;

weak "false 5th" plication present; lip strongly

thickened, with distinct denticulation

Protoginella

9b. With at most a weak, broadly rounded posterior

notch in corner of aperture; columellar plica-

tions rounded, not appearing excavated inside

aperture; lacking "false 5th ' plication; lip mod-

erately thickened, usually lacking denticula-

tion, if present, usually weak Alaginella

10a. Shell either with intricate pattern of spiral and

axial elements, or shell length > 15 mm

Marginella (part)

10b. Shell usually white or uniformly colored; pat-

tern, if present, consisting of only 1 to 4 narrow

spiral bands; shell minute to medium in size,

1.9-12.0 mm in length 11

11a. Labial denticulation always present, subequal

throughout; spire low to medium height; shell

broadly obconic in shape; some species with

weak to distinct axial costae Eratoidea

lib. Labial denticulation sometimes absent, if pres-

ent, with either a single strong posterior labial

denticle, or posterior denticle distinctly stronger

than others; spire medium to tall; shell usually

narrowly biconic, rarely broadly biconic in

shape; rarely with axial costae Dentimargo

12a. Spire immersed or nearly so 13

12b. Spire low to medium 20

13a. External varix extremely weak to absent; col-

umella with 3 plications; shape narrowly cylin-

drical Volvarina (part)

13b. External varix strong, distinct, if weak to absent,

then with 4 columellar plications; columella with

3-6 plications; shape various 14

14a. Columella with 3 plications that are crowded

anteriorly; shape cylindrical to narrowK' obo-

vate 15

14b. Columella with 4-6 plications, that often oc-

cupy half or more the aperture length; shape

broadly elliptic to obovate, rarely cylindrical to

narrowly obovate 16

15a. Without labial denticulation; bod\' whorl usually

somewhat abruptK narrowed anteriorly

Balanetta

15b. With fine labial denticulation in adult; body

whorl gradual!} narrowed anteriorly

Hydroginella (part)

16a. Columella with 4, 5, or 6 plications; if with only

4 plications, then lip completely lacking den-

ticulation, shell uniformly colored or patterned,

and shell moderately large to large

Cryptospira (part)

16b. Columella with 4 plications; lip usualK' distinct-

ly denticulate; if not denticulate, then shell

white, and small to medium in size 17

17a. Columellar plications usually thin, sharp, an-

teriorly crowded; if not, then lacking labial den-

ticulation; Indo-Pacific distribution 18

17b. Columellar plications thick, moderately heavy,

not crowded anteriorly; western Atlantic dis-

tribution 19

18a. Labial denticulation strong, distinct; shell usu-

ally with distinct color, lip and apical spot darker,

rarely grayish-white; columella with 4 thin,

sharp, anteriorly crowded plications; western

Indian Ocean distribution Closia

ISb. Labial denticulation usually absent, rarely

weakly developed; shell \\ hite, lacking colora-

tion; columella with 4 thin to moderately heavy

plications occupying less than half the aperture

length, crowded anteriorly in some; Australian

or Neozelanic distribution Ovaginella

19a. Shell white; unpatterned or with 2 obscure, bro-

ken spiral bands Prunum (part)

19b. Shell distinctly colored; distinctly patterned with

spots or numerous spiral bands Bullata

20a. Columella with 3 plications that are crowded

anteriorly; weak to strong collabral parietal cal-

lus ridge present posterior to plications, the pos-

terior end often appearing as "false 4th' pli-

cation; lip with fine tlenticulation. . Hydroginella

20b. Columella with 2-6 plications that are not

crowded anteriorly; usually lacking collabral

parietal callus ridge; if present, then lip not

denticulate, or with 2nd collabral callus ridge

inside aperture and 4 plications; lip with or

without denticulation, if present, usually strong

' 21

21a. Columella with 5-6 continuous plications; shell

usually .solid gray, or gray with spiral lines or

a distinct pattern of spiral lines interrupted by

longitudinal streaks, but not white and un-

marked; spire low, depressed; shell broadly el-

liptic, obovate, pyriform, or subtriangular; me-

G. A. Coovert and H. K. Coovert, 1995

Page 77

diiim-sized to large, length 6.2-46.0 mm, Indo-

Pacific distribution Crtjptospira (part)

21b. Columella with 2-4 plications; if with more than

4 plications, then either (1) with a "false 5th"

plication and shell narrow and cylindrical, (2)

color pure white, opaque, with a weak siphonal

notch, or (3) western Atlantic distribution; oth-

erwise not as above 22

22a. Siphonal notch weak but present; Indo-Pacific,

Australian, or Neozelanic distribution 23

22b. Siphonal notch completely absent; if weak but

present, then West .\frican distribution; widely

distributed 24

23a. Columella with 4 plications, 4th plication often

remote; shell translucent white to white, with-

out heavy ventral callus deposits; with distinct

external varix; Australian or Neozelanic distribu-

tion Mesoginella (part)

23b. Columella with 5-6 plications; shell opaque

milky white w ith hea\\ ventral callus deposits;

w ith or without external varix; Indo-Pacific dis-

tribution Volvarina (part- nionilis Group)

24a. With "false 5th" plication present as a trans-

verse denticle or weak parietal lira, neither ex-

tending into aperture appreciably. rareK' ab-

sent; lip denticulate when fully adult; shell cy-

lindrical or subcylindical; distinct external varix

present; shell colored and multi-banded or white;

Indo-Pacific, Australian, or Neozelanic distribu-

tion Serrate

24b. With 4 columellar plications; if 5th present, then

continuous, not a "false 5th" or weak parietal

lira; lip smooth or denticulate; shell \ariously

shaped; if cylindrical or subcylindrical, then lip

smooth, not denticulate; distinct external varix

present or absent; widespread distribution .... 25

25a. Collabral parietal callus ridge present, although

often weak, plus a collabral callus ridge present

inside aperture, both just posterior to 4 strong

columellar plications; lip with strong denticu-

lation; external varix present, lip moderately to

strongly thickened, especially at anterior third;

shape obovate to subp\ riform; color translucent

white or light orange; East African distribution

Serrataginella

25b. Lacking collabral callus ridge inside aperture;

if collabral parietal callus ridge present, then

with onK 3 columellar plications and lip smooth;

otherwise lacking above combination of char-

acters 26

26a. Columellar plications 2, 3, or 4, thin and sharp;

shell very thin, translucent or hyaline, color

white to amber, usually unpatterned, rarely with

faint yellowish-white spiral bands; external var-

ix absent, if present, very weak; lip smooth,

completely lacking denticulation Hyalina

26b. Columellar plications 3, 4, or 5, not unusually

thin and sharp; if only 3, then joined by short

collabral callus ridge posteriorly; shell color var-

ious, shell often thicker, not unusually thin and

translucent; external varix present or absent, if

absent, columellar plications thicker; labial den-

ticulation present or absent 27

27a. Shell with ventral and apical callus wash mi-

nutely granulated; with distinct external varix;

lip smooth, lacking denticulation; aperture

broad; freshwater, SE Asian distribution

Rivomarginella

27b. Shell lacking minute granulations ventrally and

apically; without above combination of char-

acters; not freshwater, widely distributed 28

28a. Labial denticulation present in man\ species,

but may be absent; color uniformly white,

cream, or gra\ , or spotted to streaked in some,

but generally not with narrow darker bands

alone (bands, if present, of pale background

color interrupting darker shell color); columella

with 4-5 plications; heavy ventral callusing

common, especially callus pad near posterior

commissure of lip; if shell white and lacking

labial denticulation, then unpatterned, with 4

columellar plications, and western Atlantic dis-

tribution Priinum

28b. Labial denticulation absent; color translucent

white to orange or brown, usually with yellow,

orange, brown, or reddish-brown narrow bands,

which are darker than shell color, rarely axially

streaked, not patterned with spots; columella

with 3-4 plications; ventral callus deposits weak

or absent Volvarina (part)

Subfamily MARGINELLONINAE Coan, 1965:186, 191

Diagnosis: Shell very large; spire medium height; pro-

toconch large; lip narrowly thickened, smooth, lacking

denticulation; external varix present; siphonal notch pres-

ent; columella with 2 or 4 strong plications. Type 1 an-

imal; tentacles relatively short; eyes reduced or absent;

siphon moderately long. Type 7 radula. Marginellid buc-

cal pouch present; valve of Leiblein present, without

bypass tube; esophageal caecum absent; gland of Leiblein

large, sacculate, emptying directly into posterior end of

esophagus; paired salivary glands tubular, free, ducts free

along floor of proboscis; single accessory salivary gland

absent.

Genus Afrivoluta Tomlin, 1947

(figures 5, 6, 52)

Afrivoluta Tomim, 1947:244

Type species: A. pringlei Tomlin, 1947; OD (M) (figure

52).

Diagnosis: Shell very large, volute-shaped; spire me-

dium height; protoconch large; lip narrowly thickened,

smooth, lacking denticulation; external varix present;

broad, distinct siphonal notch present; distinct parietal

callus deposit present; columella with 4 strong, nearly

Page 78

THE NAUTILUS, Vol. 109. Nos. 2 & 3

56

Figures 52-60. Sliclls of t\pc spocies ol marf^iiicllid genera, ventral \ lews, 52. Afrivohita prin^lci Tonilin, 19-17 i',\C. Ml 81 9,

.'\j»ulhas Hank, Sonlli Alriea, trawled at 110 m Length 115.4 mm 53. Mar^ineUona gigas (Marten.s, 1904). Holotype of Sigalula

pratasemis lU-hder, 19()7, USNM 237018, VV. of Pralas Reef, South China Sea, 20°37'N, 11,5°43'E. in 380 m, gray niud and sand

bottom. After photos in Harasewych and Kantor (1991:fig. 2) and Weaver and DuPont (1970:pl. 40, fig. H, I). Length 54.1 mm.

G. A. Coovert and H. K. Coovert, 1995

Page 79

transverse plications, 1st plication beginning well pos-

terior of anterior end of columella. Type 1 animal; ten-

tacles relatively short; eyes reduced; siphon moderately

long; mantle smooth, extending at least partialK over

external shell surface.

Description: Shell (figure 52) very large (adult length

84.7-130.9 mm). Color tawny to cinnamon or pinkish-

tan. Relatively thin-shelled, surface smooth and semi-

glossy with evident growth lines, often crazed or with

weak to distinct bubbles or blisters in shell surface. Shape

oblong, volute-shaped, broadest at about posterior third,

narrowed anteri<irly, very weakly shouldered. Spire of

medium height, apex swollen, blunt. Aperture broad,

wider anteriorly. Lip narrowly thickened inside and rolled

externally, producing a distinct external varix with a

sharp dorsal margin unconnected to 1st columellar pli-

cation. Lip smooth, not denticulate. Siphonal notch broad,

distinct; posterior notch poorly defined. Parietal callus

deposit near posterior commissure of lip large to mod-

erately large. Columella with 4 very strong continuous

plications that are nearly transverse to axis, 1st plication

beginning well posterior of anterior end. Plications oc-

cupying slightly more than halt the aperture length. In-

ternal whorls unmodified.

External anatomy (figures 5, 6): 1 species studied. Type

1 animal; tentacles relatively short; eyes very reduced,

red, on lateral lappets close to edge; siphon moderately

long; mantle smooth, able to extend over at least anterior

left half of external shell surface; foot broad, about 2 X

shell width, slightly longer than shell, with posteromedial

mound corresponding to parietal callus deposit of shell;

animal uniformly colored yellowish-tan. (Based on pub-

lished data, summarized in Coovert, 1987a and Coovert,

1987g:9; plus personal observations of photographs ex.

W. R. Liltved).

Internal anatomy: L'nknown.

Radula: 1 species studied. Type 7, uniserial, ribbon

short, very broad, composed of 70-80 plates. Rachidian

plates overlapping, very broad (1.75-1.90 mm wide),

nearly flat, with numerous (70-80) sharp cusps along

posterior edge. Anterior edge of rachidian plate generally

straight, resulting in elongate, rectangular "comb-like"

plates. Shell length: radular width ratio = 61. Radular

Index = 0.8.

Distribution and Habitat: South African (1 species).

Recorded from depths of 70 to 500 m. Specimens ob-

tained by commercial bottom trawlers.

Fossil Record: Known only from the Recent.

Nomenclature, Synonymy: The type species and genus

were discussed by C'oovert (1987a). The holotype was

figured by Tomlin (1947:244) and Bruggen (1963:fig. 1).

Remarks: This monotypic genus was discussed by Coo-

vert (1987a, 1987e). The genus was correctly assigned to

the Marginellidae by Barnard (1963). The type species

has a very large, striking shell. The 4 massive columellar

plications are distinctive, with the first situated well pos-

terior of the anterior border of the columella. The very

prominent posterior parietal callus deposit is character-

istic.

Genus Marginellona Martens, 1904

(figures 28, 53, 80)

Marginclla (Marginellona) Martens, 1904:108

Sigaluta Rehder, 1967:182-183 (TS: S. pratasensis Rehder, 1967,

= Marginellona gigas Martens, 1904; OD]

Type species: Marginella (M.) gigas Martens, 1904; M

(figure .53)

Diagnosis: Shell very large, smooth and very glossy,

narrowly obovate; spire medium height with shouldered

whorls; protoconch large; lip narrowly thickened, smooth,

lacking denticulation; external varix apparently present;

a broad, weak siphonal notch present; lacking parietal

callus deposit; columella with 2 strong, very oblique pli-

cations, 1st plication bordering anterior end of columella.

Type 1 animal; tentacles relatively short; eyes absent;

siphon moderately long.

Description: Shell (figure 53) very large (adult length

53.9-L57.0 mm). Color translucent tan, yellowish-brown,

or greenish-brown, aperture darker brown. Thin-shelled,

surface smooth and very glossy. Shape narrowly obovate,

weakly shouldered. Spire medium height, with large bul-

bous protoconch and shouldered whorls, sutures callused

over. Aperture very broad, wider and truncate anteriorly.

Lip narrowly thickened, smooth, not denticulate, ap-

parently with a narrow external varix. Siphonal notch

very broad, weak; posterior notch poorly defined. Shell

with a thin parietal callus wash but no large callus de-

posits. Columella with 2 strong, nearly axially oriented

continuous plications occupying slightly less than half

the aperture length. Internal whorls unmodified.

External anatomy: 1 species studied. Type 1 animal;

tentacles relatively short; eyes absent; siphon moderately

54. Serrata serraia (Gaskoin, 1849). Holotype, BM(NH), Mauritius. After photograph in Kaicher (1981:#2690). Length 8.9 mm.

55. Serrataginella spryi (Clover, 1974). Holotype, BM(NH) 197.3.83, 240 km S. of Porto Amelia, Mozambique, low tide under

stones. After photos in Clover (1974:fig. 6) and Kaicher (1981:#2629). Length 9.5 mm. 56. Hydroginclla dispersa Laseron, 19.57.

Holotype, AMS 10:3.353, Murray Island, Torres Strait, north Queensland, 4-15 m. Length 5.1 mm, 57. Protoginella lavigata (Brazier,

1877). Synt>pe, Katow. New Guinea, mud bottom 13 m .After Hediey (1901:pi 16, fig 5). Length 6 4 mm. 58. Alaginella ochracea

(Angas, 1871). GAG Ace. # 28-89, Little Bay, New South Wales. Length 3,5 mm. 59. Austroginella nmscaria (Lamarck, 1822).

GAG M12.32, Victoria, Australia, on beach Length 15.3 mm. 60. Mesoginella lurbinata (G. B. Sowerby II, 1846). GAG MI655,

Port Stephens, New South Wales. Length 7.2 mm.

Page 80

THE NAUTILUS, Vol. 109, Nos. 2 & 3

long; mantle extension undetermined; foot broad, about

2 X shell width, slightly longer than shell (Harasewych

and Kantor, 1991:10).

Internal anatomy (figure 80): 1 species studied. Same

as for subfamily.

Radula (figure 28): 1 species studied. Type 7, uniserial,

ribbon short, very broad, composed of 56-80 plates. Ra-

chidian plates overlapping, very broad (1.6-2.5 mm wide),

nearly flat, with numerous (58-85) sharp cusps along

posterior edge. Anterior edge of rachidian plate generally

straight, resulting in elongate, rectangular "comb-like"

plates. Shell length: radular width ratio = 43-63. Radular

Index = 0.9-1.0.

Distribution and Habitat: Indo- Pacific (1 species). Oc-

curs from the eastern Indian Ocean off the Nicobar Is-

lands and the South China Sea. Recorded from depths

of 380 to 1,280 m.

Fossil Record: Known only from the Recent.

Nomenclature, Synonymy: The type species of Sigaluta

is a synonym of M. gigas, the type species of Margi-

nellona, thus synonymizing these two groups. See dis-

cussion in Harasewych and Kantor (1991). The holotype

of Sigaluta pratascnsis is figured herein (figure 53) and

in color in Weaver and DuPont (1970:pl. 40, figs. H, I)

and Abbott and Dance (1982:220).

Remarks: Harasewych and Kantor (1991) reviewed this

monotypic genus. This is the largest marginellid known.

The smooth and very glossy shell, 2 very oblique colu-

mellar plications, and lack of a parietal callus deposit

serve to separate the type species from Afrivohita prin-

glei, the only marginellid species with which it is likely

to be confused.

Subfamily MARGINELLINAE Fleming, 1828:328

Diagnosis: Shell minute to very large, white, uniformly

colored, or patterned; lip thickened, smooth or dentic-

ulate; external varix present or absent; siphonal notch

present or absent; columella with 2-6 plications. Type 2

animal; tentacles long to very long, slender; eyes at base

of tentacles on slight swelling; siphon moderately to very

long; mantle smooth, pustulose, or papillose, usually at

least partially extending over external shell surface. Type

5, 6, modified Type 6, Type 8, or 9 radula, or non-

radulate. Marginellid buccal pouch present, absent in

non-radulatc species; odontophoral cartilages fused an-

teriorly or both anteriorly and posteriorly; valve of Lei-

blein present or absent, if present, with bypass tube;

esophageal caecum present or ab.sent; gland of Leiblein

with long, convoluted duct and a terminal bulb, emp-

tying either into posterior end of esophagus or passing

through nerve ring and emptying into anterior end of

proboscis; paired salivary glands a.scinous or tubular, ei-

ther attached to esophagus jusl anterior to the valve of

Leiblein, contained within proboscis, or free, ducts either

embedded or attached to walls of esophagus, or free;

single accessory salivary gland present or absent, ascinous

or tubular.

t Genus Myobanim Sohl, 1963

Mijoharum Solil, 1963:750-751

Type species: M . laevigatum Sohl, 1963 f; OD (M)

Diagnosis: Shell moderately large, surface smooth, glossy

spire medium to tall; sutures callused over; aperture broad

lip thickened, smooth; external varix weakly developed

weak siphonal notch present; strong, narrow posterior

notch present; columella with 2 widely-spaced plications,

the anterior one bordering the anterior end of the col-

umella.

Fossil Record: Late Cretaceous (Upper Maestrichtian)

of Mississippi and Georgia.

Nomenclature, Synonymy: Photographs of the holotype

were figured in Sohl (1963:pl. 90, figs. 19, 20) and Sohl

(1964:pl. 44, figs. 15, 16).

Remarks: Apparently monotypic. The two widely-

spaced columellar plications are unusual. Although orig-

inally placed in the Volutidae, Ponder (1973:331) sug-

gested that this genus might belong in the Marginellidae.

Petuch and Sargent (1986:10) mentioned Myobaruin as

a possible ancestor to the Olividae. The general shell

shape, glossy, callused surface, anterior plication bor-

dering the anterior end of the columella, and the blunt,

paucispiral protoconch all indicate a marginellid. Based

on these conchological features, this genus is here in-

cluded in the Marginellidae, subfamily Marginellinae.

Placement to tribe must await further, extensive study

of the fossil fauna.

Tribe AUSTROGINELLINI Coovert and Coovert, new

tribe

Diagnosis: Shell minute to moderately large, white,

rarely lightly colored or obscurely banded; spire im-

mersed to tall; lip thickened, smooth or denticulate; ex-

ternal varix present; siphonal notch weak or absent; col-

umella with 2-4 plications occupying more than half to

less than half the aperture; some species with a "false

4th" or "false 5th plication. Type 2 animal; siphon mod-

erately long to very long; mantle smooth, pustulose, or

papillose, extending over external shell surface. Type 5,

modified Type 6, or Type 8 or 9 radula. Marginellid

buccal pouch present; odontophoral cartilages fused an-

teriorly; valve of Leiblein present, with bypass tube;

esophageal caecum absent; gland of Leiblein with a ter-

minal bulb and a long, convoluted duct emptying into

posterior end of esophagus; paired salivary glands tu-

bular, attached to esophagus just anterior to the valve of

i>eiblein, ducts embedded in walls of esophagus; single

tubular accessory salivary gland present or absent.

G. A. Coovert and H. K. Coovert, 1995

Page 81

"Serrata Group"

Diagnosis: Shell small to medium, white to translucent,

uniformly colored or banded in some species; spire im-

mersed or low to medium; lip thickened, finely to coarse-

ly denticulate, rarely smooth; external varix present; si-

phonal notch absent; some species with collabral parietal

callus ridge; columella with 3 or 4 plications occupying

less than half the aperture length, often with "false 4th"

or "false 5th plications. Type 2 animal; siphon mod-

erately long; mantle smooth or pustulose, extending over

external shell surface. Modified Type 6, or Type 8 or 9

radula. Anatomy as in tribe, single tubular accessory

salivary gland present.

t Genus Conuginella Laseron, 1957

Connginclla Laserun, 1957;2S8

Type species: Marginella inermis Tate, 1878 f; OD

(M)

Diagnosis: Shell medium size, conical, strongly but

evenly narrowed anteriorly; spire low; lip narrowly thick-

ened, denticulate, produced or angulate posteriorly; ex-

ternal varix present; siphonal notch absent; columella

with 4 plications occupying less than half the aperture

length.

Fossil Record: Miocene of Australia.

Nomenclature, Synonymy: This group was placed by

Coan (1965:190) as a subgenus of Serrata.

Remarks: Related to Exiginella, a fossil group synon-

ymized below with Serrata. Conuginella differs in hav-

ing only 4 columellar plications, a distinctive conical

shape, and a produced posterior corner of the lip. Shared

features are the presence of an external varix, denticulate

lip, and slender shell shape.

Genus Serrata Jousseaume, 1875

(figures 26, 54)

Serrata JoiLSseauine, 1875:167, 230

Haloginclla Laseron, 1957:284 [TS: Marginella nwstelina (Aii-

gas, 1871), = Ilt/alina (\'i)lvarin(i) mtistelina Angas, 1871;

OD]

t Exiginella Laseron, 1957:289 [TS: Marginella winteri Tate,

1878 t: OD]

Type species: Serrata .serrata (Gaskoin, 1849), = Mar-

ginella serrata Gaskoin, 1849; T (figure 54)

Diagnosis: Shell small to medium, white to brown, often

banded, usually cylindrical; spire low to medium; lip

thickened, finely to coarsely denticulate, rarely smooth;

external varix present; lacking a siphonal notch; lacking

parietal callus deposits and ridge; columella with 4 strong

plications plus a weak "false 5th" occupying less than

half the aperture length. Type 2 animal; siphon mod-

erately long; mantle smooth or pustulose, extending over

external shell surface. Modified Type 6 radula.

Description: Shell (figure 54) small to medium in size

(adult length 3.6-13.0 mm). Color white to grayish-white

or brown, often with prominent brown spiral bands, or

rarely pale orangish-yellow bands; surface smooth, glossy.

Shape cylindrical to narrowly elliptic, narrowly subpy-

riform in one species; weakly to strongly shouldered.

Spire low to medium height, rarely very low and obscure.

Aperture narrow, usually wider anteriorly. Lip weakly

to strongly thickened, finely to coarsely denticulate, rare-

ly smooth, with a weak to distinct external varix. Shell

without a siphonal notch or a posterior notch. Shell with

a thin parietal callus wash, lacking large callus deposits

and collabral parietal callus ridge. Columella with 4

strong, continuous plications, plus a weak to strong pa-

rietal lira or "false 5th" plication, which combined oc-

cupy slightly less than half, but more than one-third, the

aperture length. Internal whorls presumed unmodified.

External anatomy: 2 species studied. Type 2 animal;

tentacles long, slender; siphon moderately long; mantle

smooth or pustulose, covering shell symmetrically; foot

slightly wider than shell, about 1 V2 X shell length; animal

colored with cream, orange, or buff. See summary in

Coovert (1987g:13, 20) as Haloginella.

Internal anatomy: 1 species studied. Same as for "Ser-

rata Group.

Radula (figure 26): 8 species studied. Modified Type 6,

uniserial, ribbon short, broad, of 13-35 plates. Rachidian

plates overlapping, thin, fragile, broad (0.052-0.120 mm

wide), nearly flat, with numerous (22-59) sharp cusps

along sinuous posterior edge. Anterior edge of rachidian

plate generally straight, resulting in elongate, rectangular

"comb-like" plates. Shell length: radular width ratio =

59-117. Radular Index = 0.4-1.6.

Distribution and Habitat: Neozelanic (2 species), S. Aus-

tralian (7 species), Indo-Pacific (3 species). Intertidal to

370 m.

Fossil Record: Middle Oligocene of New Zealand {fide

Powell, 1979:217), Miocene of Australia, to Recent.

Nomenclature, Synonymy: The type of M. serrata is

figured by Kaicher (1981:#2690) and herein (figure 54).

Synonymy of Serrata and Haloginella is based on radular

and conchological similarities. The radula of Australian

S. mustelina, type species of Haloginella, has been stud-

ied (pers. obs. of SEMs, ex. Dean Hewish) and compared

with radulae from S. translata (Redfield, 1870), an un-

doubted congener of S. serrata, type species of Serrata.

These radulae and shells compare very favorably, and

the generic groups they represent are herein considered

synonyms. For discussion of Haloginella, see Coovert

(1987f). Exiginella is synonymized based on concholog-

ical similarity to the other two groups, including a fairly

strong 5th columellar plication, denticulate lip, and an

external varix.

Remarks: Haloginella and Serrata had been considered

distinct until we extracted radulae of S. translata. The

Page 82

THE NAUTILUS, Vol. 109, Nos. 2 & 3

shared possession of a modified Type 6 radula, shell with

a weak "false 5th" plication, lack of a collabral parietal

callus ridge, and the usually denticulate lip are diag-

nostic. (]oan (1965:190) considered Haloginella a sub-

genus of Volvarina and was subsequently followed by

others. Anatomical features described by Fonder (1970)

clearly place this group, along with Mesoginella and

Austroginella, in a separate tribe, based especially on

the presence of a valve of Leiblein, the absence of an

esophageal caecum, and the emptying of the duct from

the gland of Leiblein into the esophagus posterior to the

nerve ring.

Genus Serrataginella Coo vert and Coovert, new genus

(figures 29, 55)

Type species: "Marginella" apryi Clover, 1974; OD (M),

herein (figure 55)

Diagnosis: Shell medium-sized, translucent white to light

orange, unhanded, broadly obovate to subpyriforrn; spire

low; lip strongly thickened, coarsely denticulate; external

varix present; lacking a siphonal notch; with collabral

parietal callus ridge plus parallel callus ridge inside ap-

erture; cohnnella with 4 strong plications occupying about

one-third the aperture length, lacking "false 5th ' pli-

cation. Type 8 radula.

Description: Shell (figure 55) medium in size (adult

length 8.0-9.9 mm). Color translucent white to light or-

ange; surface smooth, glossy. Shape broadly obovate to

subpyriform, moderately strongly shouldered. Spire low.

Aperture narrow, usually wider anteriorly, narrowest

medially due to incurving of lip. Lip moderately to

strongly thickened, especially anterior third, coarsely

denticulate in adults, denticulation absent in subadults,

first developing anteriorly, with a strong external varix.

Siphonal notch absent; posterior notch absent. Shell with

a weak but distinct collabral parietal callus ridge just

outside aperture, plus weak to distinct collabral callus

ridge well within aperture, l)oth most distinct just pos-

terior to plications. Columella with 4 strong, continuous

plications, lacking parietal lira or "false 5th ' plication.

Plications occupying about '/s aperture length. Internal

whorls presumed unmodihed.

External anatomy: 1 species studied. Animal reported

to be light orange in color ((Mover, 1974:215).

Internal anatomy: Unknown.

Radula (figure 29): 1 species studied. Type 8, iniiserial,

ribbon short, broad, of .38 [)lates. Kacliidian plates over-

lapping, broad (0.191 mm wide), nearly llat, with 9-10

sharp cusps along posterior edge. The entire posterior

edge, including all of the edges of the main cusps, with

numerous (total of ca. 70) small, subordinate cusps, giving

the main cusps a serrated ajipearance. Anterior edge of

rachidian plate slightly indented medially, resulting in

a very sliallowly V-shaped plate. Shell length: radular

width ratio = 42. Radular Index = 4.0.

Distribution and Habitat: Indo- Pacific (1 species). Re-

corded from E. African coast. Intertidal to 30 m.

Fossil Record: Known only from the Recent.

Nomenclature, Synonymy: A photograph of the holo-

type of the type species was figured by Clover (1974:pl.

8, fig. 6) and Kaicher (1981:#2629). Name derived from

Serrata, valid marginellid genus to which this new genus

is believed to be allied, and L. serrata, toothed like a

saw, serrated, in reference to the serrated cusps of the

radula, combined with -ginella, adopted from Margi-

nella. Gender feminine.

Remarks: This monotypic genus is characterized by the

very distinctive radula, possession of a collabral callus

ridge on the parietal area as well as inside the aperture,

plus the 4 strong columellar plications and strong labial

denticulation. Placement is based on conchological sim-

ilarities to Serrata and Hydroginella. See Coomans (1975)

for further information. See Key for further differenti-

ation.

t Genus Stronihoginella Laseron, 1957

Stromhoginella Laseron, 1957:289

Type species: Marginella erassidens Chapman and

C;respin, 1928 f; OD (M)

Diagnosis: Shell medium size, broadly biconic, strongly

narrowed and acute anteriorly; spire medium height;

aperture very narrow; lip thickened, denticulate; exter-

nal varix present; siphonal notch absent; columella with

4 plications crowded anteriorly.

Fossil Record: Pleistocene of Australia.

Nomenclature, Synonymy: Coan (1965:189) placed this

group as a subgenus of Marginella. but the anteriorly

crowded plications indicate little relationship.

Remarks: The broadly biconic shell shape strongly nar-

rowing anteriorly and the very narrow aperture are ap-

parently unique features of this monotypic genus. The

ilenticulate lip and general shape indicate placement in

Austroginellini, "Serrata Group, whereas the 4 anteri-

orly crowded plications indicate close relationshij) to Hy-

droginella.

Genus Hydroginella Laseron, 1957

(figures 30, 56)

llililrofiinrlla La.scroii, 1957:284

Nci>li)M>ncll(i Laseron, 1957:28.'! |'I'S: M . fa.sriciild Laseron, 1957;

ODl

rillargiiiella (laljtiel, 1962:197 (TS: Marginella colurimaria

Hedley & May, 1908; OD (M)]

Type species: //. dispersa La.seron, 1957; OD (M) (fig-

ure 56)

Diagnosis: Shell small to medium, white to orangish-

brown or amber, hyaline or translucent, rarely banded,

G. A. Coovert and H. K. Coovert, 1995

Page 83

usually cylindrical, obovate, or subpyriforni; spire im-

mersed or low to medium; lip thickened, usually finely

denticulate, rarely smootli; external varix present; lack-

ing a siphonal notch; collabral parietal callus ridge usu-

ally present; columella with 3 plications plus a weak

"false 4th' occupying less than 'A tiie aperture length.

Type 2 animal; siphon moderateK long; mantle extension

undetermined. Type 9 radula.

Description: Shell (figure 56) small to medium (adult

length 3.0-10.3 mm). Color white to orangish-brown to

amber, hyaline or translucent, one species with 3 broad

pale orange spiral bands; surface smooth, glossy. Shape

narrowly to broadly cylindrical or obovate to subpyri-

form, usually strongly narrowed anteriorly; weakly to

strongly shouldered. Spire immersed or low to medium

height. Aperture usually narrow throughout, some spe-

cies moderately broad, oiten wider anteriorly or narrow-

est medially due to incurved lip. Lip moderately to

strongly thickened, thickest medially or at anterior third,

usually finely denticulate in adults, denticulation often

weak or absent medially, rarely completely absent, with

weak to strong external varix. Siphonal notch absent;

posterior notch absent. Shell with weak to strong collabral

parietal callus ridge beginning just posterior to plications,

appearing at that point as "false 4th plication, ridge

rarely absent. Columella with 3 continuous plications

crowded anteriorly, occupying 'A or less of aperture

length. Internal whorls unmodiHed.

External anatomy: 2 species studied. Type 2 animal;

tentacles long, slender; siphon moderately long; mantle

extension undetermined; foot about as wide as shell,

slightly longer, some species with opaque white spots.

See Bouchet (1989:79, fig. 2).

Internal anatomy: Unknown.

Radula (figure 30): 4 species studied. Type 9, uniserial,

ribbon very short, narrow, greatly reduced, of 10-30

plates. Rachidiaii plates weak, overlapping, narrow

(0.019-0.032 mm wide), nearly flat, with few (4-7) cusps

along posterior edge. Anterior edge of rachidian plate

straight, resulting in subquadrate plates. Shell length:

radular width ratio = 230-391. Radular Index = 2.3-

7.5.

Distribution and Habitat: S. Australian (4 species), Indo-

Pacific (13 species). Intertidal to 550 m.

Fossil Record: Known only from the Recent.

Nomenclature, Synonymy: Bouchet (1989) discussed

Hydrogi7iella and Neptoginella and figured the holo-

types of both type species (note that his captions of figs.

d and e were reversed). Although conchological similar-

ities were discussed, these genera were not synonymized.

These groups, plus Pillarginella, are here considered con-

generic based on distinctive conchological features, es-

pecially the presence of a collabral parietal callus ridge

and only 3 columellar plications. The very distinctive

radula is known from four species (Bouchet, 1989:fig. 3;

Hewish, 1990:fig. 2A), including H. aihimnaria (pers.

obs. of SEMs, ex. Dean Hewish), the type species of

Pillarginella. The principle of first reviser (ICZN Art.

24) is employed in choosing Hydroginella over Nepto-

ginella. These 3 groups form a very distinctive assem-

blage of species.

Remarks: Bouchet (1989) reported on the parasitism of

sleeping fish by H. caledunica (Jousseaume, 1877). The

very unusual, highly reduced radula, characteristic of

this group, suggests a similar behavior in the other spe-

cies.

"Austroginella Group"

Diagnosis: Shell minute to moderately large, white,

rarely lightly colored or obscurely banded; spire im-

mersed to tall; lip thickened, smooth or denticulate; ex-

ternal varix present; siphonal notch weak or absent; col-

umella with 2-4 plications occupying more than half to

less than half the aperture, some species with a "false

5th ' plication. Type 2 animal; siphon long to very long;

mantle smooth, weakly pustulose, or papillose, extending

over external shell surface. Type 5 radula. Anatomy as

in tribe, single accessory salivary gland absent.

t Genus Mioginella Laseron, 1957

Mioginclla Laseron, 1957:287

Type species: Marginella regiila Cotton, 1949 f; OD

(M)

Diagnosis: Shell medium size, broadly biconic, nar-

rowed anteriorly; spire tall; shoulders angulate or cari-

nate; lip thickened, denticulate; external varix present;

weak siphonal notch present; broad posterior notch pres-

ent; columella with 4 plications, plus a weak 5th, occu-

pying more than half the aperture.

Fossil Record: Eocene of Australia.

Nomenclature, Synonymy: This genus was placed as a

subgenus of Marginella by Coan (1965:189). The fifth

plication is not present in Marginella s. str. and the re-

semblance is superficial.

Remarks: We consider this apparently monotypic genus

to be ancestral to Protoginella based on the presence of

a weak 5th columellar plication, tall spire, and weak

posterior notch. Laseron (1957:287) considered this genus

to be ancestral to Carinaginella based on the carinate

shoulders. Carinaginella is synonymized here with Al-

aginella because we regard carinate shoulders to be con-

vergent and not expressing a close relationship to mem-

bers of Alaginella. which also has 4 columellar plications.

Cotton (1949:218) stated that M. regula was related to

M. muscaroides Tate, 1878, a species we consider to be

clearly in Austroginella based on the distinctively shaped

lip and presence of a strong siphonal notch, and thus not

closely related to Mioginella.

Page 84

THE NAUTILUS, Vol. 109, Nos. 2 & 3

Genus Protoginella Laseron, 1957

(figure 57)

Protoginella Laseron, 1957:285

Type species: Marginella lavigata Brazier, 1877, =

Marginella (Prunum) lavigata Brazier, 1877; OD (figure

57)

Diagnosis: Shell small to medium, white, biconic or

subpyriform, strongly narrowed anteriorly; spire medi-

um height; aperture narrow; lip strongly thickened,

strongly denticulate, alate posteriorly; external varix

present; posterior notch present; siphonal notch absent;

columella with 4 strong plications that are excavated

inside aperture, plus a "false 5th"; plications occupying

more than half the aperture.

Description: Shell (figure 57) small to medium in size

(adult length 4.4-7.0 mm). Color white; surface smooth,

glossy. Shape biconic, subpyriform, strongly narrowed

anteriorly, weakly to strongly shouldered. Spire medium

height. Aperture narrow throughout. Lip strongly thick-

ened, posterior corner strongly alate, strongly denticu-

late, with distinct external varix. Shell without siphonal

notch, but anterior end somewhat truncate; posterior

notch present at posterior corner of lip. Extent of ventral

callusing undetermined. Columella with 4 strong con-

tinuous plications excavated inside aperture, plus weak

"false 5th" plication or parietal lira; plications occupying

4/5 aperture length. Internal whorls presumed unmod-

ified.

External anatomy: LInknown.

Internal anatomy: Unknown.

Radula: 1 species studied. Type 5, uniserial, ribbon

short, broad. Rachidian plates barely overlapping, mod-

erately broad (0.028 mm wide), weakly arched, with 14-

15 strong cusps on posterior edge. Central cusp strongest.

Anterior edge of rachidian plate moderately concave,

resulting in rectangular to chevron-shaped plates. Shell

length: radular width ratio = 160.

Distribution and Habitat: Indo-Pacific (2 species). Re-

corded from 13 to 97 m.

Fossil Record: Eocene and Pliocene of Australia, to

Recent.

Nomenclature, Synonymy: A syntype of the type spe-

cies is figured by Kaicher (1981:#2644) and herein (fig-

ure 57). Hedley (1901:123) emended the name to M.

laevigata, an unjustified emendation. No evidence in the

original publication itself exists of "clear evidence of an

inadvertent error," and incorrect transliteration is not to

be considered an inadvertent error [ICZN Art. 32(c)(ii)].

Thus, there is no homonymy with Marginella laevigata

Eichwald, 1830. The syntype of M. haudinensis Smith,

1899, a synonym, was figured by Kaicher (1981 :#2622),

Some authors have synonymized M. valida Watson, 1886,

which is an Alaginella and very distinct from P. lavigata.

Remarks: Laseron's original concept included M. gem-

inata Hedley, 1912. This was based on the presence of

a denticulate lip, a character variable in Alaginella, in

which this species is now placed. This restricted concept

of Protoginella is based on a strong posterior notch, col-

umellar plications appearing excavated due to callus de-

posits, presence of a weak "false 5th" plication, and a

strongly thickened, denticulate lip.

t Genus Niidifaba Eames, 1952

Marginella. subg. Niidifaba Eames, 1952:122

Type species: Marginella (N.) rakhiensis Eames, 1952

t;OD(M)

Diagnosis: Shell minute, broadly obconic; spire flat; ap-

erture narrow; lip strongly thickened, produced on pos-

terior corner, smooth, lacking denticulation; strong ex-

ternal varix present; strong posterior parietal callus de-

posit present near posterior commissure of lip, forming

weak posterior notch; siphonal notch absent; columella

with 4 strong plications occupying slightly more than

hall the aperture.

Fossil Record: Eocene of Pakistan.

Nomenclature, Synonymy: Currently considered a

monotypic genus.

Remarks: The 4 columellar plications, smooth, strongly

thickened lip that is produced or alate on the posterior

corner, presence of an external varix, and absence of a

siphonal notch all indicate placement in the Austrogi-

nellini and an ancestral relationship to Alaginella. Due

to the early occurrence of this group in the Eocene of

Pakistan, and the presence of a strong posterior parietal

callus deposit, we consider this a separate, valid genus.

Genus Alaginella Laseron, 1957

(figure 58)

Alaginella La.senm, 1957:286

Carinaginclla Laseron, 1957:286 [TS: Marginella carinata E.

A. Smith, 1891; OD (M)]

t Cassoginella Laseron, 1957:287 [TS: Marginella palla (Cotton,

1949 t; OD (M)]

Triginella Laseron, 1957:280-281 [TS: Marginella malina

Hedley, 1915; OD (M)]

Type species: Marginella oehracea Angas, 1871; OD

(figure 58)

Diagnosis: Shell minute to medium, white, biconic, ob-

ovate, or subpyriform, usually strongly narrowed ante-

riorly; spire low to medium height; aperture narrow to

moderately narrow; lip moilerately thickened, denticu-

late to smooth, alate posteriorly; external varix present;

posterior notch weak or absent; siphonal notch absent;

columella with 4 strong unexcavated plications occu-

pying slightly more than half the aperture, lacking "false

5th" plication. Type 2 animal; siphon long; mantle pa-

pillo.se, extending over external shell surface.

G. A. Coovert and H. K. Coovert, 1995

Page 85

Description: Shell (figure 58) minute to medium in size

(adult length 1.7-13.0 mm). Color white; surface smooth,

glossy. Shape biconic, obovate to subpyriform, usually

strongly narrowed anteriorly; weakly to strongly shoul-

dered, rarely strongly carinate. Spire usually medium

height, rarely very low to low. Aperture narrow to mod-

erately narrow, rarely broad, narrowest medially in some

species. Lip moderately thickened, thickest medially to

posteromedially, posterior corner strongly alate, dentic-

ulate to smooth, with a distinct external varix. Shell with-

out siphonal notch, posterior notch weak, poorly defined

to absent. Shell without obvious ventral callusing. Col-

umella with 4 strong continuous plications occupying

slightly more than half the aperture length. Internal

whorls unmodified.

External anatomy: 2 species studied. Type 2 animal;

tentacles long, slender; siphon long; mantle covering most

or all of external shell surface, papillose with conspicuous,

branched papillae; foot nearly as wide as shell, twice as

long; animal white, dotted with brown. [Based on de-

scription of A. geminata (Hedley, 1912) in Cotton (1944:

16) (as weedingi Cotton, 1944) plus description of A.

ochracea (Angas, 1871), ex. unpublished mss., ex. R. Burn

and D. R. Hewish].

Internal anatomy: Unknown.

Radula: 9 species studied. T\pe 5, uniserial, ribbon

short, broad, of 20-55 plates. Rachidian plates usually

overlapping, moderately broad (0.019-0.040 mm wide),

weakly arched, with 1 1-20 strong cusps on posterior edge.

The central cusp usually strongest. Anterior edge of ra-

chidian plate slightly to strongly concave, resulting in

rectangular to chevron-shaped plates. Shell length: rad-

ular width ratio = 86-174. Radular Index = 1.5-4.4.

Distribution and Habitat: S. Australian (8 species), Indo-

Pacific (8 species). South African (9 species). Intertidal

to 1,650 m.

Fossil Record:

Recent.

Miocene and Pliocene of Australia, to

Nomenclature, Synonymy: The types of M. ochracea,

type species of Alaginella, and M. carinata, type species

of Carinaginella. were figured by Kaicher (1981:#2618,

#2682). The radulae of these two species, plus M. mal-

ina, type species of Triginella, have been studied (pers.

obs. of SEMs, ex. Dean Hewish), and all three species

are here considered to be congeneric. Conchological dif-

ferences are considered to be of specific value only, in-

cluding the carinate shoulders of A. carinata. M. palla,

the type species of Cassoginella, has an axially costate

shell. As in other austroginelline genera, this is considered

to be of specific value only. The alate posterior end of

lip and lack of a siphonal notch place this species in

Alaginella. The principle of first reviser (ICZN Art. 24)

is herein employed in choosing Alaginella over the other,

more restricted names.

Remarks: The alate, flared shoulder of the aperture and

the anteriorly strongly narrowed shell, combined with

the lack of a strong posterior notch and lack of a "false

5th" columellar plication will serve to distinguish this

group. The presence or absence of labial denticulation

and axial costae are considered to be of specific difference

only. The branched mantle papillae may ultimately prove

to be a diagnostic character of this genus. The report on

the radula in the original description of Triginella mal-

inoides Gabriel, 1962, fig. 4 is apparently erroneous. We

have examined SEMs of a radula (ex. Dean Hewish) from

a shell intermediate between A. malina (Hedley, 1915)

and T. malinoides and it is a typical Alaginella radula.

In fact, we consider these two species synonymous. The

radula figured by Gabriel is clearly not a marginellid

radula and is obviouslv in error.

t Genus Hiivia Marwick, 1931

Marginella, subg. Hiwia Marwick, 1931:129

Type species: Marginella (Hiwia) amplificata Mar-

wick, 1931 t; OD (M)

Diagnosis: Shell small, broadly biconic, strongly nar-

rowed anteriorly; spire medium height; strong axial cos-

tae present; shoulders angulate to carinate; aperture nar-

row; lip thickened, smooth, not denticulate; posterior

corner of lip sharply angulate; external varix present;

siphonal notch absent; columella with 4 plications oc-

cupying more than half the aperture.

Fossil Record: Eocene of Australia, to Oligocene of

New Zealand.

Nomenclature, Synonymy: This group was considered

to be a valid genus by Coan (1965:189).

Remarks: The two included species (type species and

M. aldingae Tate, 1878) appear to be closely related, but

otherwise form a distinct, valid genus, apparently extinct

since the Oligocene. A relationship to the "Anstroginella

Group" and especially Anstroginella is indicated by the

4 columellar plications occupying more than half the

aperture, but this group lacks a siphonal notch and has

a narrow aperture.

Genus Anstroginella Laseron, 1957

(figures 25, 59, 80)

Anstroginella Laseron, 1957:285

Plicaginclla Laseron, 1957:285 [TS: Marginella formicula La-

marck, 1822; OD]

Type species: Marginella muscaria Lamarck, 1822; OD

(figure 59)

Diagnosis: Shell medium to moderately large, white,

yellowish-white, or pale yellowish-orange, biconic, ob-

ovate, or subpyriform, usually strongly narrowed ante-

riorly; spire low to medium height with shouldered whorls;

aperture broad; lip smooth, rapidly thickening posteri-

orly with sharpl) defined posterior notch; external varix

Page 86

THE NAUTILUS, Vol. 109, Nos. 2 & 3

present; siphonal notch present; heavy ventral callusing

usually present; columella with 4 strong plications oc-

cupying more than half the aperture. Type 2 animal;

siphon long to very long; mantle smooth or weakly pus-

tulose, extending over external shell surface.

Description: Shell (figure 59) medium to moderately

large (adult length 6.2-16.3 mm). Color white, yellowish-

white, or pale yellowish-orange; surface smooth, glossy,

some species with weak to distinct axial costae. Shape

biconic, narrowly to broadly obovate, or subpyriform,

usually strongly narrowed anteriorly; weakly to strongly

shouldered. Spire low to medium height; spire whorls

convex or shouldered, giving uneven profile. Aperture

broad, especially anteriorly. Lip moderately to strongly

thickened, narrow anteriorly, rapidly thickening poste-

riorly but slightly narrowed at posterior fourth, smooth,

lacking labial denticulation, with distinct external varix.

Shell with distinct siphonal notch, and strong, grooved

posterior notch at the junction of body whorl and pos-

terior commissure of lip. Shell usually with heavy ventral

callusing. Columella with 4 strong continuous plications

occupying slightly more than half the aperture length.

Internal whorls unmodified.

External anatomy: 4 species studied. Type 2 animal;

tentacles long, slender; siphon long to very long; mantle

smooth or weakly pustulose, covering most or all of ex-

ternal shell surface; foot very broad, 2-3 X as wide as

shell, about 2 X as long; animal spotted or blotched with

white, dark brown, orange, yellow, red, or green. See

summary in Ponder and Taylor (1992).

Internal anatomy (figure 80): 2 species studied. Same

as for "Austroginella Group."

Radula (figure 25): 5 species studied. Type 5, uniserial,

ribbon short, broad, of 29-61 plates. Rachidian plates

overlapping, moderately broad (0.029-0.114 mm wide),

weakly arched, with 13-20 strong cusps on posterior edge.

Central cusp strongest. Anterior edge of rachidian plate

slightly to moderately concave, resulting in chevron-

shaped plates. Shell length: radular width ratio = 118-

162. Radular Index = 1.7-3.5.

Distribution and Habitat: S. Australian (6 species). In-

tertidal to 27 m.

Fossil Record: Miocene to Pleistocene of Australia, to

Recent.

Nomenclature, Synonymy: The nomenclature of these

two genus-group names and their type species were dis-

cussed in Coovert (1988b), in which the presence of axial

costae are considered to be of specific value only.

Remarks: This well-characterized group is distin-

guished by the presence of 4 strong cohiniellar plications

occupying more than half the aperture, combined with

the presence of a siphonal and a posterior notch, along

with the distinctive lip shape. For an account of pred-

atorv shell drilling and anatomv, see Ponder and Tavlor

(1992).

Genus Mesoginella Laseron, 1957

(figure 60)

Mesoginella Laseron, 1957:282

Deviginella Laseron, 1957:283-284 [TS: Marginclla brachia

Watson, 1886, = Marginella {Glabella} brachia Watson,

1886; OD]

t Hianoginetla Laseron, 1957:288 [TS: Marginella physa Cot-

ton, 1949 t: OD (M)]

Sinuginella Laseron, 1957:282 [TS: Marginella inconspicua G.

B. Sowerby II, 1846; OD]

Spiroginella Laseron, 1957283 [TS: Marginella leia Cotton,

1944, = M. turbinata G B. Sowerby 11, 1846; OD (M)]

t Urniginella Laseron, 1957:287 [TS: Marginella cassidijormis

Tate, 1878 t; OD (M)]

Type species: Marginella turbinata G. B. Sowerby II,

1846; OD (M) (figure 60)

Diagnosis: Shell small to medium, white or yellowish-

white, rarely brownish-orange or with pale bands, bi-

conic, obconic, obovate, or broadly cylindrical, not

strongly narrowed anteriorly; spire low to medium height

with evenly contoured whorls; aperture moderately nar-

row; lip smooth to denticulate, moderately to strongly

thickened, thickest medially; external varix present; weak

siphonal notch usually present; posterior notch weak to

absent; ventral callusing usually absent; columella with

4 strong plications occupying slightly less to slightly more

than half the aperture, 4th plication often remote. Tvpe

2 animal; siphon long; mantle smooth or weakly pustu-

lose, extending over external shell surface.

Description: Shell (figure 60) small to medium in size

(adult length 2.5-11.0 mm). Color usually white to yel-

lowish-white, semi-opaque to translucent, rarely brown-

ish-orange or with pale yellow bands; surface smooth,

glossy, some species with weak to distinct axial costae.

Shape narrowly to broadly obovate, obconic, biconic, or

broadly cylindrical, not strongly narrowed anteriorly;

weakly to strongly shouldered. Spire low to medium

height with evenly contoured whorls. Aperture moder-

ately narrow, usually wider anteriorly. Lip moderately

to strongly thickened, thickest medially, usually smooth,

some species with weak to moderately strong labial den-

ticulation, with distinct external varix. Siphonal notch

nearly absent to weak, rarely strong, distinct or com-

pletely absent. Posterior notch weak, poorly defined or

absent. Shell usually without evident ventral callusing.

Columella with 4 strong continuous plications, 4th often

remote, rarely with parietal tubercle appearing as "false

5th ' plication. Plications occupying slightly less to slight-

ly more than half the aperture length. Internal whorls

unmodified.

External anatomy: 5 species studied. Type 2 animal;

tentacles long, slender; siphon long; mantle smooth or

weakly pustulose, asymmetrically extending over exter-

nal shell surface; foot narrow to broad, slightly narrower

to 1 '/2 .\ as wide as shell, about I'/s X as long; animal

variously spotted, blotched, or with lines of various colors.

G. A. Coovert and H. K. Coovert, 1995

Page 87

including white, yellow, orange, brown, and black. See

summary in Coovert (1987g).

Internal anatomy: 1 species studied. Same as for "Aus-

trogiiiella Group.

Radula: 16 species studied. Type 5, uniserial, ribbon

short, broad, of 19-75 plates. Rachidian plates usually

overlapping, moderately broad (0.018-0.050 mm wide),

weakly arched, with 9-22 strong cusps along posterior

edge. The central cusp (or two subcentrals) is the strong-

est. Anterior edge of rachidian plate slightly to strongly

concave, resulting in rectangular to chevron-shaped plates.

Shell length: radular width ratio = 101-221. Radular

Index = 1 5-7.1.

Distribution and Habitat: Neozelanic (13 species), S

Australian (13 species), Indo- Pacific (7 species). Intertidal

to 640 m.

Fossil Record: Miocene, Pliocene, and Pleistocene of

Australia, to Recent.

INomenclature, Synonymy: The nomenclature of Me-

soginella and Sinuginella and their type species were

discussed by Coovert (1988b). The type of M. inconspi-

cua was figured by Kaicher (1981;#2702). M. brachia,

type species of Deviginella, falls well within the limits

of Mesoginella. The principle of first reviser (ICZN Art.

24) is here employed in choosing the better known Me-

soginella over Deviginella. The holotype of M. leia was

figured in Hewish and Gowlett-Holmes (1991;64, figs.

E-F) and synonymized with M. turbinata, thus placing

Spiroginella in synonymy. The two fossil genera Hian-

oginella and Urniginella are here synonymized, based

on conchological characters falling within the limits of

Mesoginella. Laseron (1957:284) used several nomina

nuda that are apparently manuscript names for genera

synonymized here. These should not be further consid-

ered.

Remarks: The presence or absence of axial costae are

not considered to be a genus-level character in this group.

The 4th columellar plication is often remote, i.e. sepa-

rated farther from the other 3 plications. This is a rather

diverse genus but all species possess the same type of

radula and conchological characters.

Genus Closia Gray, 1857

(figure 61)

Closia Gray. 1857:36

Type species: Closia sarda (Kiener, 1834), = Margi-

nella sarda Kiener, 1834; M (figure 61)

Diagnosis: Shell medium to moderately large, white or

colored, obovate; spire immersed; lip thickened, dentic-

ulate; external varix present; siphonal notch and posterior

notch absent; columella with 4 thin, sharp plications

crowded anteriorly.

Description: Shell (figure 61) medium to moderately

large (adult length 9.9-48.0 mm). Color grayish-white,

without other markings; pale rose, pink, or orangish-

yellow, with lip and apical spot darker brownish-rose or

yellowish-orange; or pale yellowish- to light orange with

paler mid-body band and lip darker with tiny white

specks. Shell surface smooth, glossy. Shape obovate,

weakly to strongly narrowed anteriorly, with rounded to

angulate shoulders. Spire immersed. Aperture moder-

ately narrow, wider anteriorly. Lip moderately to strong-

ly thickened, strongly to very strongly denticulate in

adults, with a distinct external varix. Shell without si-

phonal notch or posterior notch. Shell usually with evi-

dent ventral callus deposits anteriorly and especially pos-

teriorly. Some with heavy parietal callus wash abruptly

ending at entrance to aperture, creating appearance of

collabral parietal callus ridge. Columella with 4 thin,

sharp, continuous plications, crowded anteriorly, occu-

pying slightly less than Va to Vi aperture length. Internal

whorls presumed unmodified.

External anatomy: 1 species studied. Animal white,

semi-transparent. [From description of C. limpida Boz-

zetti, 1992:11, = C. majuscula (Martens, 1880)]. Pre-

sumed Type 2 animal.

Internal anatomy: Unknown.

Radula: Unknown.

Distribution and Habitat: Indo-Pacific (3 species), re-

stricted to western Indian Ocean. Recorded from 20 to

140 m.

Fossil Record: Known only from the Recent.

Nomenclature, Synonymy: Coan (1965:189), followed

by others, considered this group to be a subgenus of

Bullata, apparently based on the similarity to B. lilacina

(G. B Sowerby II, 1846). Bullata, which is included in

Prunini based on its radula, anatomy, and fossil ancestry,

has a tendency toward heavy ventral callusing and

stronger columellar plications that are not nearly as

crowded anteriorly as in Closia. Although the radula and

animal are unknown, we feel that Closia is not closely

related to Bullata and has a different ancestry.

Remarks: Closia has been variously associated with Bul-

lata and Ovaginella. Conchologically, it comes closest to

Ovaginella, with which it shares shell shape, and thin,

sharp, anteriorly crowded columellar plications. The

coarser labial denticulation, distinct shell pigmentation,

heavier posteroventral callus deposit, and Indian Ocean

distribution serve to distinguish it. Until the radula and

animal are known, these two groups are provisionally

considered distinct.

Genus Ovaginella Laseron, 1957

(figure 62)

Ovaginella Laseron, 1957:280

Type species: Marginella ovulum G. B. Sowerby II,

1846; OD (figure 62)

Page 88

THE NAUTILUS. Vol. 109, Nos. 2 & 3

69

FiKures 61-69. Shells of tvpe .species of marginellid genera, ventral views. 61. Closia ^ania (Kiener, l*^*^-!* GAG M1341 off

Anibanja, N.E. Madagascar.' dredged. Length 17.6 mm. 62. Ovaginella ovulum (G. B. Sowerby II, 1846). Type, BM(NH) 80.9.^5,

locality unknown. After photograph in Kaicher (1981:#2647). Length ca. 9 mm. 63. Balanetta baylei ^"'f'^^'^'^lJ^'^ ^-.^^

M1607 Margaret River, Western Australia, on beach. Length 10.5 mm. 64. Voharina mitrella (Risso, 1826). OAC MZbM, balina

Bav Malta, dredged on sandy bottom at 6 m. Length 9.3 mm 65. Prunum prurium (Cimelin, 1791). GAG M1393, Venezuela^

Length 26 9 mm. 66. Bullata Imllaia iBorn, 1778). GAC: M1787, Praia de Forte Gragoata, Niteroi, Rio de Janeiro, Brazil. Length

G. A. Coovert and H. K. Coovert, 1995

Page 89

Diagnosis: Shell small to medium, white, elliptic to ob-

ovate; spire immersed; lip thickened, usually smooth,

rarely denticulate; external varix present; siphonal notch

and posterior notch absent; columella with 4 anteriorly

crowded plications.

Description: Shell (figure 62) small to medium in size

(adult length 4.2-10.1 mm). Color translucent to semi-

opaque white; surface smooth, glossy. Shape narrowly to

broadly elliptic to obovate, weakly to strongly shoul-

dered. Spire immersed. Aperture moderately narrow,

slightly wider anteriorly. Lip moderately thickened, usu-

ally smooth, rarely weakly denticulate, with distinct ex-

ternal varix. Siphonal notch weak or absent; posterior

notch absent. Shell usually without evident ventral cal-

lusing. Columella with 4 continuous plications crowded

anteriorly, occupying distinctly less than half aperture

length. Internal whorls unmodified.

External anatomy: Unknown.

Internal anatomy: Unknown.

Radula: 2 species studied. Type 5, uniserial, ribbon

short, broad, of 20-31 plates. Rachidian plates overlap-

ping, moderately broad (0.025-0.035 mm wide), weakly

arched, with 8-14 strong cusps along posterior edge. The

central cusp is the strongest. Anterior edge of rachidian

plate slightly concave, resulting in rectangular to chev-

ron-shaped plates. Shell length: radular width ratio =

170-285. Radular Index = 1.8-3.5.

Distribution and Habitat: Neozelanic (2 species), S Aus-

tralian (2 species). Recorded from 3 to 370 m.

Fossil Record: Known only from the Recent.

Nomenclature, Synonymy: The type species of Ova-

ginella, M. ovulum, is figured by Kaicher (1981:#2674)

and herein (figure 62). Coan (1965:189) placed Ovagi-

nella as a subgenus of Balanetta. but the latter has only

3 columellar plications and a narrower shape.

Remarks: Although Balanetta and Ovaginella both have

a Type 5 radula, they are considered distinct based on

conchological differences.

Genus Balanetta Jousseaume, 1875

(figure 63)

Balanetta Jousseaume, 187.5:168, 269

Type species: B. baylei Jousseaume, 1875; M (figure 63)

Diagnosis: Shell minute to medium, white, cylindrical

to narrowly obovate; spire immersed; lip thickened,

smooth; external varix present; with weak or absent si-

phonal notch; posterior notch absent; columella with 2

or 3 plications crowded anteriorly.

Description: Shell (figure 63) minute to medium in size

(adult length 1.5-12.0 mm). Color translucent to opaque

white or yellowish-white; surface smooth, glossy. Shape

cylindrical to narrowly obovate, weakly to strongly shoul-

dered. Spire immersed. Aperture moderately narrow,

distinctly wider anteriorly. Lip moderately thickened,

smooth, not denticulate, with a distinct external varix.

Siphonal notch weak or absent, posterior notch absent.

Shell without evident ventral callusing. Columella with

3 continuous plications (one species with 2), crowded

anteriorly, occupying less than Vi aperture length. In-

ternal whorls presumed unmodified.

External anatomy: Unknown

Internal anatomy: L'nknown.

Radula: 1 species studied. Type 5, uniserial, ribbon

relatively short, broad, of 29-37 plates. Rachidian plates

overlapping, moderately broad (0.027-0.031 mm wide),

weakly arched, with 9-14 strong cusps on posterior edge.

Central cusp strongest. Anterior edge of rachidian plate

slightly concave, resulting in rectangular to chevron-

shaped plates. Shell length: radular width ratio = 185-

239. Radular Index = 2.6-3.1.

Distribution and Habitat: S. Australian (3 species), Indo-

Pacific (1 species). Intertidal to 183 m.

Fossil Record: Known only from the Recent.

Nomenclature, Synonymy: The type species was ded-

icated to M. Bayle in the original description by Jous-

seaume (1875:269, 274) but spelled "baylii" in the text

and "baylei" in the figure caption (op. cit., pi. 8, fig. 5).

This is to be considered a case of multiple original spell-

ings, which is dealt with in the ICZN [Art. 32(b)(i)]. This

section, then, is subject to Section 32(c)(ii), which con-

siders an incorrect original spelling one in which the

original publication itself shows clear evidence of an

inadvertent error. Because Jousseaume intended the spe-

cies to be dedicated to M. Bayle, it is clear that the

spelling "baylei" is the correct one. Tomlin (1917:252)

considered "baylei" an error.

Remarks: The immersed spire, lack of labial denticu-

lation, and presence of an external varix and only 3

columellar plications, distinguishes this group.

Tribe PRUNINI Coovert and Coovert, new tribe

Diagnosis: Shell small to very large, white, uniformly

colored, patterned, or banded; spire immersed, or low to

tall; lip thickened, smooth to denticulate; external varix

present or absent; siphonal notch usually absent; posterior

notch absent; columella with 2-6 plications occupying

half or less of the aperture. Type 2 animal; siphon long

57.6 mm. 67. Rivomarginella morrisoni Brandt, 1968. GAC Ace. # 11-89, Prachin River, Kabinburi. Thailand. Length 10 0 mm.

68. Cryptospira tricincta (Hinds, 1844). GAC M1253, Formosa Strait, S.W. Taiwan, dredged at .37 m. Length 23.8 mm. 69.

Hyalina pallida (Linne, 1758). GAC M2068, St. Croix, U.S. Virgin Islands. Length 14.3 mm.

Page 90

THE NAUTILUS, Vol. 109, Nos. 2 & 3

to very long; mantle smooth, pustulose, rarely distinctly

papillose, usually extending over external shell surface.

Type 6 radula or non-radulate. Marginellid buccal pouch

present, absent in non-radulate species; odontophoral

cartilages present in radulate species, fused anteriorly or

both anteriorly and posteriorly; valve of Leiblein absent;

esophageal caecum present; gland of Leiblein with long,

convoluted duct and a terminal bulb, passing through

nerve ring and emptying into anterior end of proboscis;

paired salivary glands ascinous or tubular, either con-

tained within proboscis or free, ducts either attached to

walls of esophagus or free; single accessory salivary gland

present or absent, ascinous or tubular.

Genus Voharina Hinds, 1844

(figures 38, 64)

Marginella, section Volvarina Hinds, 1844:75

Type species: Marginella nitida Hinds, 1844, = Mar-

ginella (Volvarina) nitida Hinds, 1844, = Valuta mi-

irella Risso, 1826; SD Redfield, 1870:221 (figure 64)

Diagnosis: Shell small to moderately large, color trans-

lucent white, amber, or brown, usually with darker bands;

spire immersed, or low to tall; lip thickened, smooth, not

denticulate; external varix usually absent, but weak to

strong in some species; siphonal notch absent or weak;

columella with 3 or 4 plications, some species with weak-

er 5th or 6th, combined occupying half or less of the

aperture. Type 2 animal; siphon long to very long; mantle

smooth or usually pustulose, rarely distinctly papillose,

usually extending over external shell surface. Type 6

radula.

Description: Shell (figure 64) small to moderately large

(adult length 2.6-22.4 mm). Color translucent to opaque

white, amber, or brown, usually with yellow, orange,

brown, or reddish spiral bands, rarely axially streaked;

surface smooth, glossy. Shape elongate to moderately

broadly cylindrical, elliptic, narrowly to broadly obovate,

or oblong, rarely elliptic-cylindrical with immersed spire;

weakly to strongly shouldered. Spire rarely immersed,

usually low, medium, or tall. Aperture narrow to mod-

erately broad, rarely broader, usually wider anteriorly.

Lip narrowly to strongly thickened, smooth, not dentic-

ulate, usually lacking an external varix, some species with

weak to strong external varix. Siphonal notch rarely weak,

usually absent; posterior notch absent. Ventral callusing

usually not evident, present in one species group. Col-

umella usually with 4 continuous plications; rarely with

3 plications and "false 4th" plication that joins short

collabral parietal callus ridge; or 5 plications, the 5th not

continuous past V2 whorl internally, plus often "false 6th"

plication or parietal lira posteriorly. Plications usually

occupying less than half, but some species up to half the

aperture length. Internal whorls umnodified.

External anat<imy: 26 species studied. Type 2 animal;

tentacles long, slender; siphon long to very long; mantle

smooth or usually pustulose, rarely distinctly papillose.

extending over external shell surface, often nearly com-

pletely covering shell; foot moderately broad, about 1%

X shell width, 1 '/2-2 X length; animal variously spotted

or mottled in white, black, or various colors.

Internal anatomy (figure 38): 2 species studied. As in

tribe. Marginellid buccal pouch present; odontophoral

cartilages fused both anteriorly and posteriorly; paired

salivary glands ascinous, either contained within pro-

boscis or free, ducts either attached to walls of esophagus

or free; single accessory salivary gland present, ascinous

or tubular.

Radula: 43 species studied. Type 6, uniserial, ribbon

short, broad, of 31-75 plates. Rachidian plates overlap-

ping, broad (0.034-0.562 mm wide), nearly flat, with 10-

33 (exceptionally 5) sharp cusps on posterior edge. An-

terior edge of rachidian plate generally straight, resulting

in elongate, rectangular "comb-like" plates. Shell length:

radular width ratio = 25-96. Radular Index = 1.0-4.5.

Distribution and Habitat: Neozelanic ( 1 species), S. Aus-

tralian (2 species), Indo-Pacific (33 species), E. Pacific (3

species), W. Atlantic (24 species), Arctic (1 species), Mag-

ellanic / Antarctic (3 species), Mediterranean (1 species),

W. African (29 species). South African (10 species). In-

tertidal to 1,780 m.

Fossil Record: Eocene of France, Oligocene to Pleis-

tocene of W. Atlantic, early Miocene of W. Pacific, Mio-

cene to Pliocene of Italy, Pleistocene of California, to

Recent.

Nomenclature, Synonymy: As discussed in Gofas (1989b:

160), the correct type species designation, by subsequent

designation, is Redfield (1870:221). Mention in Hinds

(1844:75) of M. avena as a "typical species" does not

constitute a type designation, and the Cossmann (1899:

92) designation was preceded by that of Redfield.

Remarks: Apparently the smooth mantle does not readi-

ly cover the external shell surface in the narrow, non-

varicose species. The broader, varicose West African and

Mediterranean species all have a pustulose mantle that

more readily covers the external shell surface (see Gofas,

1989b, and Gofas & Fernandes, 1992). Several unde-

scribed Claribbean species, discussed in Coovert and

Coovert (1990) (as Pnimim sp., red-papillose mantle; and

Volvarina sp. SW), have a distinctly papillose mantle

that readily covers the shell. This is the most widely

distributed genus as currently conceived, being found in