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Diseño y maquetación: Gonzalo Rodríguez

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Iberus, 30 (1): 1-14, 2012

© Sociedad Española de Malacología

The genus Pradoxa Fernandes & Rolán, 1993 (Gastropoda:

Muricidae) in SaoTomé, Príncipe and Annobón

El género Pradoxa Fernandes & Rolán, 1993 (Gastropoda:

Muricidae) en SaoTomé, Príncipe y Annobón

Roknd HOUART* and Emilio ROLAN**

Recibido el 15-VI-201 1. Aceptado el 6-X-2011

ABSTRACT

The genus Pradoxa is revised and all the species are described, compared and illustrated.

Two new species are described from Sao Tomé, Pradoxa gorii spec. nov. and P.

urdambideli spec. nov. The radula is illustrated for P. confirmata (Fernandes & Rolán,

1990), P. fhomensis (Fernandes & Rolan, 1990) and P. gorii spec. nov. The subfamilial

classification is discussed and a comparativo table is added in the appendix.

RESUMEN

El género Pradoxa es revisado, con todas las especies descritas, ilustradas y comparadas.

Se describen dos nuevas especies de Sao Tomé, Pradoxa gorii spec. nov. y P. urdambideli

spec. nov. Se ilustra la rádula para P. confirmata (Fernandes y Rolán, 1 990), P. fhomensis

(Fernandes y Rolán, 1 990) y P gorii spec. nov. La clasificación a nivel de subfamilia se

discute y un cuadro comparativo se aporta en un apéndice.

INTRODUCTION

The genus Pradoxa was created to

inelude two strange species, P. confir¬

mata and P. thomensis, originally

described in Buccinidae from Sao Tomé

Island by Fernandes & Rolán (1990).

However, the radula morphology (Figs.

4J-L, 5A=-E) is typical for Muricidae with

a rachidian tooth bearing seven cusps

and denticles and with a sickle-shaped

lateral tooth on each side. Both species

have now been reported from Príncipe

and Annobón, two other islands of the

Gulf of Guinea.

Pradoxa was included by Houart

(1996: 116) in the subfamily Muricopsi-

nae, together with Muricopsis, Pavartia,

Murexiella and Suhpterynotus, because of

their almost similar radula characters. In

any case, the shell morphology presents

some peculiar characteristics which

clearly distinguishes it from any other

muricid genera and which are com-

mented upon in the present work.

The protoconch of the genus Pradoxa

is unusual for the family Muricidae and

differs completely from any other

similar genus, showing some very

strong and high spiral cords separated

by deep interspaces (Figs. 2E-F, 3A“H).

This character is very similar in the four

* Research Associate, Institut royal des Sciences naturelles de Belgique, Rué Vauder, 29, B-1000 Bruxelles,

Belgium. roland.houart@skynet.be

Museo de Historia Natural, Campus Universitario Sur, 15782 Santiago de Compostela,

emiliorolan@gmail.com

1

IberuSy 30 (1), 2012

Table 1. Terminology used to describe the spiral cords and the internal denticles of the outer lip

(based on Merle 1999 and 2001) (Figs 2A-D).

Tabla I. Terminología usada para describir los cordones espirales y los dentículos internos del labio exte¬

rior (basada en Merle 1999 y 2001) (Figs 2A-D).

species included in the genus (Table II).

In some species of Muricopsis, one or

two shallow spiral cords forming a

shoulder may be presen! but never four

to six cords with deep interspaces. The

presence of a very small nucleus, with a

diameter of 120-200 pm is another addi-

tional unusual character.

Abbreviations

IRSNB: Instituí royal des Sciences natu-

relles de Belgique, Bruxelles,

Belgium.

MHNS: Museo de Historia Natural of

the University of Santiago de Com-

póstela, Spain.

MNCN: Museo Nacional de Ciencias

Naturales, Madrid, Spain.

MNHN; Muséum national d'Histoire

naturelle. Taris, Trance.

JCD: Collection Jean-Claude Delemarre.

PR: Collection Peter Ryall.

RH: Collection Roland Houart.

SG: Collection Sandro Gori.

Iv: Collected alive.

dd: Empty shell.

See Table I for additional terminology.

SYSTEMATICS

Genus Pradoxa Fernandes & Rolán, 1993

Paradoxa Fernandes & Rolán, 1990: 342 (júnior homonym of Paradoxa Marshall, 1894).

Paradoxon Fernandes & Rolán, 1990: 67 (replacement ñame for Paradoxa Fernandes & Rolán, 1990

non Marshall, 1894, but júnior homonym of Paradoxon Fleutiaux, 1903).

Pradoxa Fernandes & Rolán, 1993: 61 (replacement ñame for Paradoxon Fernandes & Rolán, 1990

non Paradoxon Fleutiaux, 1903).

Type species by original designation: Paradoxa confirmata Fernandes & Rolán, 1993, Recent, Sao Tomé.

Description: Shell very small, up to 5

mm in height at maturity, narrow, spire

very high. Protoconch large with a small

nucleus and 1.5-2. 2 rounded whorls

sculptured by 4-6 narrow, rounded, high

spiral cords with a deep interspace

between each pair of cords. Teleoconch

of 3. 5-4. 2 whorls. Spiral sculpture of

teleoconch whorls consisting of primary

spiral cords, very rarely with additional

secondary cords. Axial sculpture of

numerous narrow ribs. Aperture small,

width 600-900 pm x height 500-700 pm,

with 4 or 5 narrow, elongate denticles

within the outer lip. Operculum broadly

ovate with subapical nucleus and weak

inner curve on abapical left. Siphonal

canal broadly open, very short, 7-10% of

2

Houart and RoláN: The genus Pradoxa ie SaoTomé, Príncipe and Annobón

total Shell heigth. Radula rachiglossate

with a three-dimensional rachidian

bearing a central cusp with a single

lateral denticle, a single marginal cusp

on each side and a very small marginal

cusp.

Pradoxa confirmata (Fernandes & Rolán, 1990) (Figs. lA-D, 2A, E-F, 4A, C, G, })

Paradoxa confirmata Fernandes & Rolán, 1990: 343, text fig. IB, figs 4=7.

Type material examined: Holotype MNCN 15.05/1041 and several paratypes (see Rolán & Fer-

nandes,1990)

Other material examined: Sao Tomé: Lagoa Azul, 4 m, MHNS, 6 Iv (4 coated for SEM); Lagoa Azul,

0.5-3 m, under stones, RH, 5 Iv & dd; Esprainha, 1-5 m, MHNS, 7 Iv & dd; Esprainha, 2-3 m, RH, 8

dd; Mmerio,10 m, MHNS,12 Iv & dd; 30 m, MHNS, 5 dd; Sao Tomé Town, 1-3 m, MHNS, 21 Iv &

dd, RH, 2 Iv; Sant'Ana, 0.2-3 m, brush of algae, RH, 4 Iv; P. Mutamba, 6 m, MHNS, 73 dd (mostly

fragments and protoconchs). Príncipe: Baía de Santo Antonio, 5 m, MHNS, 17 dd. Annobón: San

Antonio de Palé, 1-8 m, MHNS, 97 dd (mainly fragments).

Type locality: Sao Tomé, Praia Mutamba.

Description: Shell small, up to 5 mm in

height at maturity. Height/ width ratio 2.6-

2.7. Slender, narrowly ovate, heavy,

nodose. Subsutural ramp broad, strongly

sloping, weakly concave. Entirely brown,

chestnut-brown or blackish brown. Inside

of aperture weakly lighter coloured.

Spire very high with 2-2. 1 protoconch

whorls and teleoconch of up to 4-4.2

weakly convex, narrow, weakly shoul-

dered, nodose whorls. Suture weakly

adpressed. Protoconch large, broadly cylin-

drical, width 900 pm x height 800 pm.

Whorls with 4 high, narrow, rounded

spiral cords with a deep channel between

each pair of cords. Each cord giving rise

to a nodose spiral cord on first teleoconch

whorl. Terminal lip not present.

Axial sculpture of teleoconch whorls

consisting of high, narrow, rounded,

nodose ribs. First whorl with 15-17 ribs,

second with 17 or 18, third with 20, fourth

with 20-22 ribs. Four or five last ribs of last

teleoconch whorl fused, giving rise to a

broad, high apertural lip. Spiral sculpture

of high, strong, narrow, nodose, primary

cords. First to tMrd teleoconch whorls with

SP, P1-P3, P3 occasionally partially or com-

pletely covered by next teleoconch whorl;

last whorl with SP, P1-P6, ADP, (MP). SP

broad, P1 and P2 of approximately similar

strength and height, P3 weakly narrower

and lower, P4 lowest cord, occasionally

obsoleto, P5 and P6 broadest and highest,

similar in size, ADP narrow, MP very iow

and narrow or obsolete. Presence of high

rounded nodes at intersection of spiral

cords with axial ribs, giving a nodose

appearance to the whole shell.

Aperture small, 800-900 pm high x 500

pm wide, ovate. Columellar lip narrow,

smooth, occasionally with weak knob

abapically. Rim completely adherent. Anal

notch shallow, broad. Outer lip smooth,

with 5 weak, elongate dentides within:

ID, D1-D4, D4 broadest and strongest,

probably D4-D5 fused. Siphonal canal very

short, 7 or 8% of total shell heigth, narrow,

straight, broadly open.

Operculum light brown, ovate, with

subapical nudeus and weak curve on

abapical left. Radula rachiglossate with a

three-dimensional rachidian tooth bearing

a short, broad central cusp, a narrow, long

lateral denticle, a broad, long lateral cusp

and a very small marginal cusp. Lateral

teeth sickle-shaped with broad base.

Distribution: Sao Tomé, Príncipe and

Annobón, living in 0.5-10 m.

Remarks: Pradoxa confirmata is the

largest species of Pradoxa with specimens

reaching 5 mm in height. It is easily dis-

tinguished from the other species in

having a strongly nodose shell and a very

high spire, a narrow shell with very

weakly convex teleoconch whorls and a

very low or obsolete P4 on the last teleo¬

conch whorl. The protoconch is also dif-

ferent in having 4 broad spiral cords vs 5

or 6 in the other species.

3

Iberus, 30 (1), 2012

Figure 1. A-D: Pradoxa confirmata Fernandes & Rolán, 1990; A, B: Sao Tomé, Lagoa Azul, 2-3 m, 4.9

mm (MHNS); C; Lagoa Azul, 4 m, 4.1 mm (MHNS); D: San Antonio de Palé, Annobon, 3.2 mm

(MHNS). E-G: Pradoxa thommsis Fernandes & Rolán, 1990; E, F: Sao Tomé, Esprainha, 1-5 m, 3.3 mm

(MHNS); G: Sao Tomé, Lagoa Azul, 2-3 m, 2.7 mm (MHNS). H-L: Pradoxa gorü spec. nov. Sao

Tomé, Minerio, 37-43 m; H, I; holotype MNHN 24260,3 mm; J, K: paratype MHNS 100559, 2.9 mm;

L: paratype MHNS 100559, 2.9 mm. M-Q: Pradoxa urdambideli spec. nov. Sao Tomé, Minerio, 41 m;

M, N: paratype SG , 3.6 mm; O, P: holotype MNHN 24264, 3.3 mm; Q: paratype RH, 3.7 mm.

Figura 1. A-D: Pradoxa confirmata Fernandes & Rolán, 1990; A, B: Sao Tomé, Lagoa Azul, 2-3 m, 4,9

mm (MHNS); C: Lagoa Azul, 4 m, 4.1 mm (MHNS); D: San Antonio de Palé, Annobon, 3.2 mm

(MHNS). E-G: Pradoxa thomensis Fernandes & Rolán, 1990; E, F: Sao Tomé, Esprainha, 1-5 m, 3.3

mm (MHNS); G: Sao Tomé, Lagoa Azul, 2-3 m, 2.7 mm (MHNS). H-L: Pradoxa gorü spec. nov. Sao

Tomé, Minerio, 37-43 m; H, I; holotipo MNHN 24260,3 mm;J, K: paratipo MHNS 100559, 2.9 mm;

L: paratipo MHNS 100559, 2.9 mm. M-Q: Pradoxa urdambideli spec. nov. Sao Tomé, Minerio, 41 m;

M, N: paratipo SG , 3.6 mm; O, P: holotipo MNHN 24264, 3.3 mm; Q: paratipo RH, 3.7 mm.

4

Houart and RoláN: The genus Pradoxa in SáoTomé, Príncipe and Annobón

Figure 2. A-D: morphology of the spiral structure and of the apertural denticles. A: Pradoxa confir-

mata Fernandes & Rolán, 1990, Sao Tomé; B: Pradoxa thomensis Fernandes 8¿: Rolán, 1990, Sao

Tomé; C: Pradoxa gorii spec. nov. Sao Tomé (holoíype); D: Pradoxa urdambideli spec. nov. Sao

Tomé. E, F: protoconch of Pradoxa confirmata Fernandes & Rolán, 1990.

Figura 2. A-D: mofología de la estructura espiral y dentículos de la abertura. A: Pradoxa confirmata

Fernandes & Rolán, 1990, Sdo Tomé; B: Pradoxa thomensis Fernandes & Rolán, 1990, Sáo Tomé; C:

Pradoxa gorii spec. nov. Sáo Tomé (holotype); D: Pradoxa urdambideli spec. nov. Sáo Tomé. E, F: pro-

toconcha £7/Pradoxa confirmata Fernandes & Rolán, 1990.

5

Ibems, 30 (1), 2012

Pradoxa thomensis (Femandes & Rolan, 1990) (Figs. lE-F, 2B, 3A-C, 4B, D, H, K-L)

Paradoxa thomensis Femandes & Rolán, 1990: 345, figs 1-3.

Type material examined: Holotype MNCN 15.05/1042 and several paratypes (see Rolán & Fer-

nandes, 1990)

Other material examined: Sao Tomé: Lagoa Azul, MFINS, 2 Iv; Esprainha, 1-5 m, MHNS, 6 Iv &

dd, RH, 2 Iv. Príncipe: Baía de Santo Antonio, 5 m, MHNS, 1 dd (juvenile).

Type locality: Sao Tomé.

Description: Shell small, up to 3.6 mm

in height at maturity. Fieight/ width ratio

2. 1-2.4. Slender, lanceolate, narrowly

convex, lightly built, tuberculate. Subsu¬

tural ramp narrow, weakly sloping on first

teleoconch whorl, more strongly sloping

on adapical whorls, weakly concave.

Entirely brown or blackish brown. Inside

of aperture similarly coloured.

Spire very high with 1.5-1.75 proto-

conch whorls and teleoconch of up to 4

broad, convex, weakly shouldered, nodose

whorls. Suture weakly adpressed. Proto-

conch large, broad, width 600 pm x height

500 pm. Whorls rounded with 4 or 5 high,

narrow, rounded spiral cords with a more

or less deep channel between each pair of

cords. Fifth cord usually covered or par-

tially covered by first teleoconch whorl.

Each cord giving rise to a narrow, weakly

nodose spiral cord on first teleoconch

whorl. Terminal lip not present.

Axial sculpture of teleoconch whorls

consisting of moderately high, broad,

rounded, weakly nodose ribs. First and

second whorls with 10 or 11 ribs, third

with 9-11, fourth with 11 ribs. Apertural

rib of last teleoconch whorl 2 or 3 times

broader than other ribs. Spiral sculpture

of low, roimded, narrow, smooth primary

cords. First and second teleoconch

whorls with SP, P1-P4, P4 usually par-

tially or completely covered by next

teleoconch whorl; third whorl with SP, IP,

P1-P4; last whorl with SP, IP, P1-P6, ADP,

MP, (ABP). IP, P1-P4 narrow, of approxi-

mately similar size, P5 and P6 weakly

broader, ADP and MP broadest; ABP

when present, low and shallow. Spiral

cords weakly broader and higher at inter-

section of spiral and axial sculpture.

Aperture small, 700-900 pm high x 500

pm wide, ovate. Columellar lip narrow,

smooth, occasionally with weak, broad

knob abapically Rim completely adher-

ent. Anal notch shallow, broad. Outer lip

smooth, with 5 weak, elongate denticles

within: ID, D1-D4, D4 broadest and

strongest, probably D4-D5 fused. Siphonal

canal very short, 6 or 7 % of total shell

length, narrow, straight, broadly open.

Operculum light brown, ovate, with

subapical nucleus and weak curve on

abapical left. Radula rachiglossate with a

three-dimensional rachidian tooth bearing

a long broad central cusp, a narrow, long

lateral denticle, a broad, long lateral cusp

and a very small marginal cusp. Lateral

teeth sickle-shaped with broad base.

Distribution: Sao Tomé and Príncipe,

living at 1-5 m.

Remarks: Pradoxa thomensis differs from

P. confirmata in being comparatively

smaller and broader with more strongly

convex teleoconch whorls. In having a

comparatively more rounded and broader

protoconch with narrow, lower spiral

cords, in having less numerous axial ribs

on the teleoconch whorls (9-11 vs 15-22),

in having a different spiral sculpture mor-

phology (Table II) and less obvious nodes

at the intersection of spiral and axial sculp¬

ture, and in having a longer siphonal canal

relativo to its shell height.

Pradoxa gorii spec. nov. (Figs. IH-L, 2C, 3D-F, 4E, I, 5A-E)

Type material: Holotype MNHN 24260. Paratypes: Sao Tomé, Minerio, 37-43 m - 1 IRSNB IG.31

896/ MT2540; 2 MHNS 100559;! MNCN (catalogue number pending). Lagoa Azul, 30-35 m - 1 JCD.

Lagoa Azul, 37 m - 2 RH; 5 SG; 10 PR.

ó

Houart and RoláN: The genus Pradoxa in SaoTomé, Príncipe and Annobón

Figure 3. Protoconchs. A-C: Pradoxa thomensis Fernandes & Rolán, 1990, Sao Tomé; D-F:

Pradoxa gorii spec. nov. Sao Tomé; G, H: Pradoxa urdambideli spec. nov. Sao Tomé.

Figura 3. Protoconchas. A-C: Pradoxa thomensis Fernandes & Rolán, 1990, Sao Tomé; D-F: Pradoxa

gorii spec. nov. Sao Tomé; G, H: Pradoxa urdambideli spec. nov. Sao Tomé.

7

Iberus, 30 (1), 2012

Other material examined: Sao Tomé: Lagoa Azul^ 37 m, 41 Iv, SG; Lagoa Azul, 30-35 m, 13 dd, RH;

9 Iv & dd, MHNS; Minerio, 41 m, 9 Iv, SG; Minerio, 37-43 m. 9 Iv & dd, RH; 7 dd (fragments), MHNS.

Etymology: We are very pleased to ñame this new species in honour of our mutual friend Sandro

Gori (Livomo, Italy), who provided us with the material studied herein.

Type locality: Sao Tomé, Minerio, 37-43 m.

Description: Shell small, up to 33 mm

in height at maturity. Height/width ratio

1.9-2.2. Lanceolate, narro wly convex, tu-

berculate. Subsutural ramp broad,

strongly sloping, almost straight or

weakly convex. Creamy white or light

tan with darker coloured band on subsu¬

tural ramp, on abapical part of last teleo-

conch whorl and on tip of siphonal

canal, P2 and P3 creamy white. Proto-

conch occasionally slightly darker, Aper-

ture creamy white or light tan with

brown band adapically and abapically

on columellar lip and inside of aperture.

D4 darker brown.

Spire high with 13-1.75 protoconch

whorls and teleoconch of up to 3.5 broad,

convex, weakly shouldered, nodose

whorls. Suture weakly adpressed. Proto¬

conch large, broad, width 600 pm x

height 400-500 pm. Whorls rounded with

5 high, narrow, rotmded spiral cords with

a more or less deep channel between

each pair of cords. Last adapical cord nar-

rower. Each cord giving rise to a narrow,

nodose spiral cord on first teleoconch

whorl. Terminal lip not apparent.

Axial sculpture of teleoconch whorls

consisting of high, broad, rounded, no¬

dose ribs. First whorl with 10-12 ribs,

second with 9-11 ribs, last whorl with 10

or 11 ribs. Apertural rib of last teleo¬

conch whorl twice as broad as other

ribs. Spiral sculpture of high, rounded,

strong, broad, smooth, primary cords.

First and second teleoconch whorls with

SP, IP, P1-P3; third and last whorls with

SP, IP, P1-P6, ADP, MP, (ABP). IP, P1 nar¬

row, P2 and P3 broadest cords, P4, P5,

P6, ADP broad, of similar size, MP and

ABP narrow when present. Spiral cords

weakly broader and higher at intersec-

tion of spiral and axial sculpture.

Aperture small, 600-700 pm high x

500 pm wide, broadly ovate. Columellar

lip narrow, smooth, with single, strong,

broad knob abapically. Rim completely

adherent. Anal notch shallow, broad.

Outer lip smooth, with 4 weak, elongate

denticles within: D1~D4, D4 broadest

and strongest, probably D4-D5 fused.

Siphonal canal very short, 7-10 % of

total Shell height, narrow, straight,

broadly open.

Operculum colourless, translucent,

ovate, with subapical nucleus and weak

curve on abapical left. Radula rachiglos-

sate with a three-dimensional rachidian

(Right page) Figure 4. A: Pradoxa confirmata Fernandes & Rolán, 1990, Sao Tomé, holotype

MNCN 15.05/1041, 4.3 mm. B: Pradoxa thomensis Fernandes & Rolan, 1990, Sao Tomé,

holotype MNCN 15.05/1042, 3.4 mm, C-F: Coated SEM; C: Pradoxa confirmata Fernandes &

Rolán, 1990, Sao Tomé, 3.5 mm; D; Pradoxa thomensis Fernandes & Rolán, 1990, 2.9 mm; E:

Pradoxa gorii spec. nov. Sao Tomé, 3.1 mm; F: Pradoxa urdambideli spec. nov. Sao Tomé, 3.4 mm.

G-I: opercula; G: Pradoxa confirmata Fernandes & Rolán, 1990 (SEM A. Warén); H: Pradoxa

thomensis Fernandes & Rolán, 1990 (SEM A, Warén); I: Pradoxa gorii spec. nov. J-L: radulae; J:

Pradoxa confirmata Fernandes & Rolán, 1990 (SEM A. Warén); K-L: Pradoxa thomensis Fernandes

& Rolán, 1990 (SEM A. Warén).

(Página derecha) Figure 4. A: Pradoxa confirmata Fernandes & Rolán, 1990, Sao Tomé, holotype

MNCN 15.05/1041, 43 mm. B: Pradoxa thomensis Fernandes & Rolán, 1990, Sao Tomé, holotype

MNCN 15.05/1042, 3.4 mm. C~F: Coated SEM; C: Pradoxa confirmara Fernandes & Rolán, 1990,

Sao Tomé, 3.5 mm; D: Pradoxa thomensis Fernandes & Rolán, 1990, 2.9 mm; E: Pradoxa gorii spec.

nov. Sao Tomé, 3.1 mm; F: Pradoxa urdambideli spec. nov. SMo Tomé, 3.4 mm, G-I: opercula; G:

Pradoxa confirmata Fernandes & Rolán, 1990 (SEM A. Warén); H: Pradoxa thomensis Fernandes &

Rolán, 1990 (SEMA. Warén); I: Pradoxa gorii spec. nov. J~L: radulae; J: Pradoxa confirmata Fernandes

& Rolán, 1990 (SEM A. Warén); K-L: fméaxA thomcmis Fernandes & Rolán, 1990 (SEM A. Warén).

8

Houart and RoláN: The genus Pradoxa in SaoTomé, Príncipe and Annobón

9

Iberus, 30 (1), 2012

tooth bearing a long broad central cusp,

a narrow, long lateral denticle, a broad,

long lateral cusp and a very small mar-

ginal cusp. Lateral teeth sickle^-shaped

with broad base.

Distribution: Sao Tomé, living at 30^

47 m.

Remarks: Pradoxa gorii spec. nov.

differs in many ways from P. confírmala

(Figs. lA-D) and does not need to be

compared here with this species.

Pradoxa gorii spec. nov. differs from

P. thomensis (Figs. 1E=G) in having a

more strongly shouldered shell with

lower teleoconch whorls, a broader,

weakly convex and less sloping subsu¬

tural ramp, comparatively broader axial

ribs, broader and more closely set spiral

cords with a different rnorphology, a

more obvious apertural D4, and a differ¬

ent microsculpture of the protoconch

(Fig. 3) In all the populations there is an

axial striation between the spiral cords

of the protoconch; this is more irregular

in P. thomensis (Fig, 3C) and more

regular in P gorii spec, nov. (Fig. 3F).

Pradoxa gorii spec. nov. also has a dif¬

ferent shell colour, the shell of P thomen¬

sis being uniformly dark brown (see also

Table II).

Pradoxa urdamhideli spec, nov. (Figs. IM-Q, 2D, 3G-H, 4F)

Type material: Holotype MNHN 24264. Paratypes: Sao Tomé, Minerio, 41 m - 2 MHNS 100560; 2

RH; 1 SG; 1 PR (Iv & dd).

Etymology: The species is named after Carlos Urdambidelus, physitian and recognized researcher

in genealogy, friend and family of the júnior author.

Type locality: Sao Tomé, Minerio, 41 m.

Description: Shell small, up to 3,8

mm in height at maturity. Height/ width

ratio 2. 0-2.2. Lanceolate, narrowly

convex. Subsutural ramp broad, weakly

sloping, weakly convex. White or light

creamy white with a light tan or light

brown band on abapical part of last

teleoconch whorl, covering P5, P6, ADP,

MP and ABP, and an occasional addi-

tional very lightly coloured spiral band

on subsutural area. Aperture white

within, apertural denticle D4, columel-

lar lip and occasionally inside of outer

lip dark brown.

Spire very high with 1.5-1.75 proto¬

conch whorls and teleoconch of up to 4

broad, convex, weakly or strongly

shouldered, lightly nodose whorls.

Suture weakly adpressed. Protoconch

iarge, broad, width 600-700 pm x height

500-600 pm. Whorls rounded with 6

narrow, rounded, moderately high

spiral cords with a moderately deep

channel between each pair of cords,

Sixth cord (adapical) occasionally

covered by first teleoconch whorl. Each

cord giving rise to a narrow nodose

spiral cord on first teleoconch whorl.

Terminal lip not apparent.

Axial sculpture of teleoconch whorls

consisting of moderately high, broad,

rounded ribs. First whorl with 10-12

ribs, second with 10 or 11 ribs, third

with 10-12 ribs, last whorl with 11 ribs,

Apertural rib of last teleoconch whorl

two or three times broader and quite

higher than other ribs, Spiral sculpture

of low, rounded, narrow, smooth, pri-

mary and secondary cords, First and

second teleoconch whorls with SP, IP,

P1-P4, P4 occasionally covered by next

whorl; third whorl with SP, (adis), IP,

P1-P4 or P1-P5, last whorl with SP, adis,

IP, P1-P6, ADP, MP, (ABP). Primary

cords of similar size, P1 occasionally

weakly broader and higher. Spiral cords

very weakly broader and higher at in-

tersection of spiral and axial sculpture.

Aperture small, 800-900 pm high x

600-700 pm wide, broadly ovate. Col-

umellar lip narrow, smooth, with occa¬

sional, single, weak knob abapically.

Rim partially weakly erect abapically,

adherent adapically. Anal notch shallow,

broad. Outer lip smooth, with 5 weak,

elongate denticies within: ID, D1-D4, D4

broadest and strongest, probably D4-D5

fused, DI rarely split. Siphonal canal

10

Houart and RoláN: The genus Pradoxa in SaoTomé, Príncipe and Annobón

Figure 5. A-E: radular morphology of Pradoxa gorii spec. nov. Radula characters (rachidian tooth):

cc= central cusp, ld= lateral denticle, lc= lateral cusp, mc= marginal cusp (Figure 5D).

Figura 5. A-E: morfología radular de Pradoxa gorii spec. nov. Caracteres de la rádula (diente

raquídeo): cc- cúspide central, ld= dentículo lateral, le- cúspide lateral, mc= cúspide marginal (Figura

5D).

Iberus, 30 (1), 2012

Table II. Comparison of the shell morphology in the Pradoxa species.

Tabla 11. Comparación de la morfología de la concha de las especies de Pradoxa.

12

Houart and RoláN: The genus Pradoxa in SaoTomé, Príncipe and Annobón

very short, 8-9 % of total shell heighp

narrow, straight, broadly open.

Operculum and radula unknown.

Distribution: Only known from the

type locality.

Remarks: Pradoxa urdamhideli spec.

nov. differs from P. gorii spec. nov. (Figs.

IH-L) in having a larger shell with the

same number of teleoconch whorls, in

having a comparatively broader and

larger aperture, weakly higher and less

shouldered teleoconch whorls, espe-

cially the last whorl, and in having com-

paratively narrower spiral cords, more

widely spaced from each other and of

approximately similar size vs more vari-

able in P. gorii spec. nov. (see Table II), in

the presence of adis, not observed in P.

gorii spec. nov. (Fig. 2D) and in having a

DISCUSSION

There are still strong doubts about

the correct subfamilial placement of this

genus.

We consider that the following differ-

ences are very important and significant

characters which differentiate Pradoxa

from the other genera in Muricopsinae

and from other subfamilies as well:

“ an almost cylindrical, very small

shell with a very high spire;

” a very short siphonal canal and a

very small aperture vs a very large and

broad protoconch;

- an operculum with a weak curve

on abapical left (Figs. 4G-I);

- the form, the peculiar sculpture

and the large size of the protoconch.

Unfortunately the DNA study of

Pradoxa did not give any information to

date. Further attempts will be made

with new material soon. We will follow

the next steps very closely and will

BIBLIOGRAPHY

Fernandes F. & RoláN E.1989. Nuevo genero

y nuevas especies de la familia Buccinidae

Rafinesque, 1815 (Mollusca, Neogastropoda)

de la Isla de Sao Tomé. Bollettino Malacologico,

25 (9-12); 341-348.

protoconch with 5 or 6 spiral cords,

instead of 4, rarely 5 in P gorii spec. nov.

(Figs. 3D-H)

Pradoxa urdambideli spec. nov. differs

from P thomensis (Figs. lE-G) in having

more strongly shouldered teleoconch

whorls, a comparatively broader proto¬

conch, a broader aperture, narrower

axial ribs, a protoconch with 5 or 6

narrow spiral cords on the last whorl

instead of 4 in P. thomensis, in the pres¬

ence of the secondary cord adis on

penultimate and last teleoconch whorls

and, as in P gorii, in having a different

shell colour. Apparently, P. urdambideli is

also the only species with a lightly

abapically erect columellar lip while

completely adherent in the three other

species. See also Table II.

publish the results of these investiga-

tions and their conclusions.

ACKNOWLEDGEMENTS

Photographs with the scanning elec¬

trón microscope (SEM) were made by

Jesús Méndez and Inés Pazos in the

Centro de Apoyo Científico y Tec¬

nológico a la Investigación (CACTI) of

the University of Vigo and by Anders

Warén (Natural History Museum, Stock-

holm, Sweden). Some optical pho¬

tographs were made in the Servicio de

Xenética (XB2) of the same University.

The authors also thanks Sandro Gori,

Livorno, Italy, for the loan and gift of the

material collected during some trips to

the islands, and John Wolff, Lancaster,

Pennsylvania, USA, for checking the

English text and for his useful comments.

Fernandes F. & Rolán E. 1990. Paradoxon no-

men novum pro Paradoxa Fernandes & Rolán,

1989 (Mollusca, Neogastropoda). Bollettino

Malacologico, 22 (1-4): 67.

13

Iberus, 30 (1), 2012

Fernandes F. & Rolán E. 1993. Pradoxa nomen

novum pro Paradoxon Fernandes y Rolán,

1990. Iherus, 11 (1): 61.

FIouart R. 1996. Les Muricidae d'Afrique Oc-

cidentale. I. Muricinae & Muricopsinae. Apex,

11 (3-4): 95-161.

Merle D. 1999. La mdiation des Muricidae (Gas-

tropoda : Neogastr opoda) au Paléogéne: approche

phylogénétique et évolutive. París. Thése de

doctoral du Muséum National d'Histoire

Naturelle: i-vi, 1-499.

Merle D. 2001. The spiral cords and the inter¬

nal denticles of the outer lip in the Murici¬

dae: terminology and methodological com-

ments. Novapex, 1 (3): 69-91.

14

Iherus, 30 (1); 15-40, 2012

© Sociedad Española de Malacologta

A taxonomic revisión of Oestophora barbula (Rossmássler,

1838) and O. barbella (Servain, 1880), two Iberian ende-

mic land-snail species (Gastropoda: Trissexodontidae)

Revisión taxonómica de Oestophora barbula (Rossmássler, 1838) y

O, barbella (Servain, 1880), dos especies de caracoles terrestres endé¬

micas de la península ibérica (Gastropoda: Trissexodontidae)

David T HOLYOAK and Geraldine A. HOLYOAK*

Recibido el 7-VII-201 1. Aceptado el 17-X-201 1

ABSTRACT

Oestophora barbula and O. barbella are distinct biológica! species that can be reliably

distinguished oniy by the longer and stouter penia! epiphallus of the former taxon. Oesfop-

hora barbula is endemic in western-central Portugal, whereas O. barbella has a much lar-

ger range in western Iberia, including parts of the area occupied by O. barbula. Most

populations of O. barbula have strongly keeied shells and they live mainly in habitats with

exposed (Mesozoic) limestone rock. Oestophora barbella has the shell periphery rounded

to moderately keeied, a small minority of its shells being indistinguishable from some of O.

barbula. Oestophora barbella is found in varied habitats disturbed by man and in more

natural situations, some of which have very acidic or strongly basic rocks. It occurs in

rocky (Mesozoic) limestone habitats in the Aígarve and Serra da Arrábida, but is usuaily

replaced by O. barbula in similar habitats over much of western-central Portugal. The two

species have been found living together oniy at one locality in the Serra da Arrábida,

where they differ widely in shell size. Type material of both species has been located and

lectotypes are designated here.

RESUMEN

Oestophora barbula y O. barbella son especies biológicas diferentes, que pueden distin¬

guirse sólo por el epifalo más largo y más grueso del primer taxón. Oestophora barbula es

un endemismo del oeste y centro de Portugal, mientras que O. barbella tiene una distribu¬

ción mucho más extensa en el oeste de la Península Ibérica, incluyendo partes del área

ocupada por O. barbula. La mayoría de las poblaciones de O. barbula tienen conchas

con una fuerte quilla y viven en hábitats expuestos de roca caliza (Mesozoico). Oestop¬

hora barbella tiene la periferia de la concha redondeada o con una quilla moderada, aun¬

que una pequeña minoría de sus conchas fuera indistinguible de algunos O. barbula. Oes¬

tophora barbella se encuentra en distintos hábitats alterados por el hombre y también en

otros más naturales, algunos de ellos con rocas muy ácidas o ultrabásicas. Se encuentra en

hábitats de roca caliza (Mesozoico) en el Algarve y la Serra da Arrábida, pero suele estar

reemplazado por O. barbula en este tipo de hábitats en la mayor parte del oeste y centro

de Portugal. Las dos especies se han encontrado simpátricas sólo en una localidad de la

Serra da Arrábida, en donde difieren ampliamente en el tamaño de la concha. El material

tipo de ambas especies ha sido localizado y se designan aquí lectotipos.

* Quinta da Cachopa, Barcoila, 6100-014 Cabe9udo, Portugal; holyoak9187@ hotmail.com

15

Iberus, 30 (1), 2012

INTRODUCTION

Oestophora bartula is currently regar-

ded as a widespread and often common

land snail throughout the western half

of the Iberian Península, readily distin-

guished from other species of the genus

by the usual presence of two palatal

teeth in the shell mouth (Puente, 1996).

Although its local populations vary con=

siderably in shell size and development

of a peripheral keel, variation in the

genital anatomy has been regarded as

rather slight and taxonomically unim-

portant in most of the recent literature.

Castillejo, Outeiro and Rodríguez

(1987) reported some differences in the

distal genitalia and in development of a

shell keel within coexisting populations

in the Serra da Estrela of eastern Portu¬

gal, leading them to argüe that O.

barbilla was coexisting with O. barbella at

those localities. However, we agree with

Puente (1996) that the rather small dif¬

ferences in genital anatomy demonstra-

ted in that study can be attributed to

individual variation within a single

species.

The first detailed description of the

genital anatomy of O. barbilla by Hesse

(1931) nevertheless reported different

genital anatomy to that described and

figured by almost all subsequent

authors (cf. Ortiz de Zárate López,

1962; Manga, 1983; Castillejo, 1984;

Castillejo et al., 1987; Puente, 1996),

with a much longer and thicker penial

epiphallus. Ortiz de Zárate López

(1962) attributed this difference to the

epiphallus in Hesse's material being

swollen due to the supposed presence of

a spermatophore inside, a suggestion

later cited by Puente (1996). However,

this seems unlikely as the shape of the

spermatophore in Helicacea is determi-

ned by the internal shape of the epipha¬

llus in which it forms, rather than vice¬

versa. Hesse dissected three individuáis

from near Lisbon and he presented three

drawings (Hesse, 1931: Taf. 8, Pigs. 63 A-

C) each showing similar anatomy with a

large epiphallus, so it anyway seems

unlikely that all three should have the

epiphallus similarly distended by a

spermatophore which Hesse did not

notice.

The present paper reports a study of

Oestophora barbilla sensu lato from Portu¬

gal and neighbouring regions of Spain.

It soon became apparent that many

populations from western-central Portu¬

gal show the large epiphallus figured by

Hesse and these were mainly also cha-

racterised by strongly keeled shells.

Rossmássler's type material of O.

barbilla appears to be of this localised

endemic form. All other populations

studied resembled Spanish material in

having a smaller epiphallus and they

almost always lacked a really strong

keel on the shell; the oldest valid ñame

applicable to this form is O. barbella

(Servain, 1880). Lectotypes are designa-

ted here for both ñames. The two taxa

live cióse together in parts of western

Portugal and living populations of both

with no intermediates were collected

from beneath the same boulders at one

locality in the Serra da Arrábida. We

also discuss variability in shell morpho-

logy and problems of species identifica-

tion from shells, and apparent differen¬

ces in the range of habitat preferences of

each species.

MATERIAL AND METHODS

Pield collections by the authors con-

sisted of all mature shells found, to

avoid possible bias in favour of large

specimens. Localities and altitudes were

recorded mainly using a Carmín Ltrex

High Sensitivity hand-held GPS, accu-

rate to within 10 metres. From 2007

onwards sites were given consecutive

serial numbers (e.g. P37). Habitat notes

(including bedrock type and vegetation)

and associated Mollusca were also

recorded at all sites. Adult snails could

readily be recognised by the thickened,

reflected lip to the shell mouth and

these normally had enlarged distal geni¬

talia. Living adult specimens collected

by the authors for anatomical study

were drowned by immersion in water

overnight, transferred to 80% Industrial

Methylated Spirit, then pulled or part-

ló

Holyoak and HolyoaK: Taxonomic revisión of Oestophora barbula and O. barbella

pulled from the shell when the body

hardened after a few days, "Proximal"

and ''distal" refer to the position in reía-

tion to the ovotestis. Drawings of the

distal genital anatomy were prepared

using a Meiji draw^ing tube on a Meiji

RZ series stereomicroscope.

Measurements of shell breadth and

height, and counts of whorls followed

the methods described by Kerney and

Cameron (1979). These shell naeasure-

ments were made on adult shells to the

nearest 0.1 mm using an eye-piece grath

cule in a stereo-microscope. Áccuracy of

the counts of whorls was slightly

reduced by the protoconch often appea“

ring somewhat triangular when viewed

from above, rather than semicircular.

Measurements of (greatest) width of

umbilicus and the breadth of the same

shell were measured using Infinity

Analyze© software on images of adult

shells taken with an Infinity 1 camera on

a Meiji RZ series stereomicroscope. The

measurements on the images were

reproducidle to ± < 0.01 mm, but una-

voidable slight tilting of the shells

almost certainly caused some additional

loss of precisión.

Full details of specimens studied by

the authors are listed in the Appendix,

which ineludes all those used for anato-

mical study. In addition notes on shells

in the Coimbra, Lisbon and Porto

Museums were kindly supplied by

Alvaro De Oliveira, along with speci¬

mens and data from his personal collec-

tion; this Information has been added to

the distribution maps. Two syntypes of

O. barbula were studied from high

quality photographs supplied by the

Forschungsinstitut Senckenberg. 12

syntypes of O. barbella were studied on

loan from the Bourguignat Collection

housed at the Muséum d'Histoire Natu-

relle, Ville de Genéve. The remainder of

the material studied is retained in the

Collection of G.A. and D.T. Holyoak.

RESULTS AND DISCUSSION

Genital anatomy. The genital anatomy

of O. barbella from Spain was carefully

and accurately described by Puente

(1996) under the ñame O. barbula with

clear drawings of nine individuáis.

Henee the present account concentrates

on the differences between O. barbella

and O. barbula, which appear to be con-

fined to the distal genitalia, especially

the epiphallus and penis (Figs. 1-2).

In the present study, distal genitalia

have been dissected and described in

detail from 142 individual snails repre-

senting 66 populations (listed in Appen¬

dix). O. barbula was represented by 35

individuáis (+ 1 with genitalia too imma-

ture to be useful), representing 16 popu¬

lations; O. barbella was represented by 102

individuáis (+ 4 with genitalia too imma-

ture to be useful), representing 49 popu¬

lations. Only one population (7 indivi¬

duáis, from site P67/P161) could not be

securely assigned to either taxon on the

basis of the epiphallus; the identity of this

unusual population is discussed below.

Figures 1-2 give representative dra¬

wings of the penis and epiphallus in

varied individuáis of each species. Table I

compares measured proportions of epip¬

hallus length relative to penis length and

thickness of epiphallus compared to

thickness of penis. In mature individuáis,

the epiphallus length is generally less

than half the penis length in O. barbella,

more than half the penis length in O.

barbula. The thickness of the epiphallus is

generally less than that of most of the

penis (the epiphallus appearing thin-

walled) in O. barbella, equal to or more

than that of most of the penis (and appea¬

ring muscular and thick-walled) in O.

barbula. Combining the length and thick¬

ness criteria left very few individual

snails that were difficult to assign to one

species or the other. Study of the internal

structures of penis and epiphallus of a

few individuáis did not reveal any clear

interspecific differences.

A single population (P67/P161: ca

2.3 km NE. of Sao Pedro de Moel, Estre-

madura) remained problematical with

seemingly intermediate epiphallus cha-

racteristics based on seven mature speci¬

mens (epiphallus relatively long but

usually no stouter than penis: Fig. 2G).

On the basis of the epiphallus length

17

Iherus, 30 (1), 2012

Figure 1 . Anatomy of distal genitalia in representative specimens of Oestophora from Portugal. A:

O. barbula (Rossmássler, 1838), 2 May 2010, SE. of Moitas Venda, Ribatejo (slightly atypical spe-

cimen with four mucus glands); B: same specimen viewed from opposite side; C: O. barbella

(Servain, 1880), 3 Nov. 2010, NW. of Verride near Rio Mondego, Beira Litoral; D: same speci¬

men viewed from opposite side (specimens in collection of G.A. and D.X Holyoak). Abbrevia-

tions, a: atrium; as: accessory sac on vagina (with small dart sac at distal end); be: bursa copulatrix;

bed: bursa copulatrix duct; e: epiphallus; mg: mucus gland; p: penis; pr: penis retractor; v: vagina;

vd: vas deferens.

Figura 1. Anatomía de los genitales distales en ejemplares representativos de Oestophora de Portugal. A:

O. barbula (Rossmássler, 1838), 2 may. 2010, SE. de Moitas Venda, Ribatejo (ejemplar un poco

atipico, con cuatro glándulas mucosas); B: mismo ejemplar visto desde el lado opuesto; C: O. barbella

(Servain, 1880), 3 nov. 2010, NW. de Verride cerca de Río Mondego, Beira Litoral; D: mismo ejem¬

plar visto desde el lado opuesto (ejemplares en la colección de G.A. y D.T. Holyoak). Abreviaturas, a:

atrio; as: saco accesorio en la vagina (con pequeño saco del dardo en el extremo distal); be: bursa copula¬

trix; bed: conducto de la bursa copulatrix; e: epifalo; mg: glándula mucosa; p: pene; pr: retractor del

pene; v: vagina; vd: vaso deferente.

18

Holyoak and Holyoak: Taxonomic revisión of Oestophora harbula and O. barhella

Tabie I. Relative size of epiphallus and penis in Oestophora b arbula and O. barhella. Data are given

only for individuáis with distal genitalia sufficiently large to be regarded as mature or cióse to

maturity. N = number of individuáis.

Tabla L Tamaño relativo del epifalo y del pene en Oestophora barbula y O. barbella. Los datos se dan

sólo para los individuos con genitales distales suficientemente grandes como para ser considerados

they were certainly closer to O. barbula,

but the small, unkeeled shells (Table II,

Figs. 4G-I) were superficially much

more similar to those of some O. barbe¬

lla. However, the proportionately very

small umbilicus is much more like that

of O. barbula than of populations of O.

barbella from western-central Portugal

(cf. Table III, Figs. 3, 4 and notes below).

Nevertheless, the habitat was more like

that of other material confirmed as O.

barbella (among limestone boulders used

to construct the abutment of a small

bridge over a stream in a Coastal región

with unconsolidated Quaternary sands

supporting a tall Eucalyptus plantation;

see below for additional information).

The possibility that this population

represents an undescribed third spedes

seems unlikely in view of its lack of any

unique characters, the single locality

and the disturbed habitat. Henee it is

tentatively regarded here as a popula-

tiori of O. barbula with shells that lack a

keel and show a relatively narrow umbi¬

licus, although further study is needed

in view of the relatively weak epipha-

ilus and habitat atypical for that species

(see below).

The functional explanation of the

longer, thicker epiphallus of O. barbula

compared to that of O. barbella is uncer-

tain. It is noteworthy that the epiphallus

is approximately similar in length to the

bursa copulatrix duct in O. barbula (ratio

ca 1:1, Figs. lA, B), whereas the epipha¬

llus is much shorter and the bursa copu¬

latrix duct much longer in O, barbella

(ratio ca 1:3, Figs. IC, D). Henee, it might

be surmised that if a spermatophore is

formed inside the epiphallus as usual in

many (if not all) species of Helicacea, it

will be longer in O. barbula than in O. bar-

bella. Indeed, the epiphallus in O. barbella

is so short in comparison to its bursa

copulatrix duct that a spermatophore

formed inside it would apparently be too

short for effective fertilisation in the

manner described for Helix pomatia L. by

Lind (1973) and currently assumed to be

widespread in Helicacea. This fertilisa¬

tion mechanism irivolves the spermatop¬

hore iodging inside the bursa copulatrix

duct, so that when some of the spermato-

zoa it contains escape through its tail

canal they reach the spermatheca by way

of the oviduct. Corresponding to this

widely accepted model of the fertilisation

process, Koene and Schulenberg (2005)

fourid correlations between the length of

the penial flagellum and the spermatop-

hore-receiving organ in helicoid land

19

Iberus, 30 (1), 2012

snails in general, indicating coevolution

which probably results from counter-

adaptation between male and female

reproductive organs that may control fer-

tilisation. Likewise, Sauer and Haus-

DORF (2009) demonstrated a positivo

scaling between male spermatophore

producing organs and female spermatop-

hore=receiving organs in Xerocrassa

species (Hygromiidae sensii lato) from

Crete, indicating sexual coevolution

which they attributed to sexual selection

processes. The relativo lengths of epipha-

llus and bursa copulatrix duct in O.

barbilla and O. barbolla differ so markedly

from results of these studies that further

study is needed. Spermatophores have

apparently not been described in the lite-

rature for Oestophora and they were not

noticed during the present study Henee,

it should be established whether the

spermatozoa are transferred in sperma¬

tophores in both species, and whether or

not self-fertilisation is common. Mating

was seen only once during our fieldwork,

a pair of O. barbolla in coitus being collec-

ted on 3 November 2010, at ca 11.00

hours, on the undersurface of a limestone

boulder, 1 km NW. of Verride near the

Rio Mondego, Boira Litoral.

Seixas (1976) gave a confused

account of a Portuguese Oostophora

under the ñame "Mastigophalus rangi

(De Férussac) Deshayes, 1839", conclu-

ding that Oostophora liisitanica (L. Pfeif-

fer, 1841) may be the same species as

that correctly known as Mastigophalliis

rangianus (Férussac, 1822). The latter is

(Right page) Figure 2. Distal male genitalia in representative specimens of Oestophora from Portu¬

gal. A-F: O. barbula (Rossmássler, 1838), G: O. cf barbula (atypical population with unkeeled

shells) and H-N: O. barbolla (Servain, 1880); drawings show genitalia in situ, partly dissected to

reveal penis and epiphallus, but with only distal ends of penial retractor muscle and vas deferens

drawn. A, B: 9 Jan. 2011, by Paul do Taipal near Montemor-o-Velho, Beira Litoral; C: 13 Jan.

2011, summit area of Serra de Sico, Beira Litoral; D; 12 Feb. 2008, Serró Ventoso, Estremadura;

E: 29 Apr. 2011, just NE. of Ansiáo, Beira Litoral; F: 12 Feb. 2011, Serra da Arrábida, Estrema¬

dura, population coexisting with O. barbolla, cf Fig. 2L; G: 4 June 2011, 2 km NNE. of S. Pedro

de Moel, Estremadura, individual with longest epiphallus and (uniquely) a single minute flage-

llum; H: 20 Oct. 2008, Gerrinho das Neve, Mértola, Baixo Alentejo, Topotype; I: 15 Jan, 2011,

SW. of Proen^a-a-Velha, Beira Baixa; J: 5 Eeb. 201 1, W. slope of Foia, Algarve; K: 28 Jan. 2011, by

N2 NNW. of Alportel, Algarve; L: 12 Feb. 2011, Serra da Arrábida, Estremadura, population coe¬

xisting with O. barbula, cf Fig. 2F; M: 3 May 201 1, S. of Costa Nova do Prado, Beira Litoral; N:

27 Oct. 2010, 1 km NW. of Gafanha do Areáo, Beira Litoral (all specimens in Collection of G.A.

and D.T. Holyoak). Abbreviations as in Figure 1, plus or: ommatophore (with black or grey parts

stippled) and its retractor muscle.

(Página derecha) Figura 2. Genitales masculinos distales en ejemplares representativos de Oestophora

de Portugal. A-F: O. barbula (Rossmássler, 1838), G: O. cf. barbula (población atípica, con conchas

sin quilla) y H-N: O. barbolla (Servain, 1880); los dibujos muestran los genitales in situ, diseccionados

en parte para revelar el pene y el epifalo, pero representando sólo los extremos distales de los músculos

retractores del pene y del vas deferens. A, B: 9 ene. 2011, junto a Paul do Taipal cerca de Montemor-o-

Velho, Beira Litoral; C: 13 ene. 2011, zona de la cumbre de la Serra de Sico, Beira Litoral; D: 12 feb.

2008, Serró Ventoso, Estremadura; E: 29 abr. 2011, inmediatamente al NE. de Ansiáo, Beira Litoral;

F: 12 feb. 2011, Serra da Arrábida, Estremadura, la población conviviendo con O. barbolla, véase Fig.

2L; G: 4 jun. 2011, dos km. al NNE de S. Pedro de Moel, Estremadura, ejemplar con el mayor epifalo

y (singularmente) un solo flagelo diminuto; H: 20 oct. 2008, Gerrinho das Neve, Mértola, Baixo Alen-

tejo, topotipo; L: 15 ene. 2011, SO. de Proenga-a-Velha, Beira Baixa; J: 5 feb. 2011, ladera oeste de

Foia, Algarve; K: 28 ene. 2011, por la N2, NNO. de Alportel, Algarve; L: 12 feb. 2011, Serra da

Arrábida, Estremadura, la población conviviendo con O. barbula, véase Fig. 2F; M: 3 may. 2011, S.

de Costa Nova do Prado, Beira Litoral; N: 27 oct. 2010, 1 km NO. de Gafanha do Areao, Beira

Litoral (todos los ejemplares en la colección de G.A. y D.T. Holyoak). Abreviaturas como en la Figura 1,

además de or: ommatóforo (con partes negras o gris punteadas) y su músculo retractor.

20

Holyoak and HolyoaK: Taxonomic revisión of Oestophora barbula and O. barbella

21

Iberus, 30 (1), 2012

Table II. Measurements of shells of Oestophora barbula and O. barbella. Data are given only for

populations identified anatomically from epiphallus structure by the authors, for which 7 or more

adult shells were available (specimens in authors’ collection). N= number of individuáis; s.d.=

sample standard deviation. Rock types at each site are coded as follows, DS: Coastal sand dunes;

LB: limestone boulders placed to reinforce river banks, and a bridge abutment; Ls: limestone

bedrock; Sh: shale; SI: slate; Ss: sandstone.

Tabla II. Mediciones de conchas de Oestophora barbula y O. barbella. Los datos se dan sólo para las

poblaciones identificadas por los autores a partir de la estructura anatómica del epifialo, de las cuales se

disponía de 7 o más conchas de adultos (ejemplares en la colección de los autores). N= número de indivi¬

duos; s.d. = Desviación estándar de la muestra. Tipos de roca en cada sitio representados como sigue, DS:

dunas de arena costeras; LB: rocas calizas fiormando riberas artificiales o el embasamiento de un puente;

Ls: afloramiento de roca caliza; Sh: esquistos; SI: pizarras; Ss: areniscas.

generally regarded as a distinctive

endemic species restricted to a small

Coastal región in S. France (Depts Pyré-

nées Orientales and Var) and NE. Spain

(Prov. Gerona) (Puente, Altonaga,

Prieto and Ruiz, 1998: 47). Shells of

these taxa have very little in common,

O. lusitanica having a rounded shell

periphery and no apertural teeth, M.

rangianus having a sharp keel and pro-

minent apertural teeth. It is unclear

whether the Portuguese specimens des-

22

Holyoak and HolyoaK; Taxonomic revisión of Oestophora barbula and O. barbella

Table III. Relative width of the umbilicus in shells of Oestophora barbula, O. barbella and a popu-

ladon of O. cf barbula from near S. Pedro de Moel (Estremadura). The data on O. barbula are

based on 20 representative mature shells selected from 6 populations; those on O. barbella on 22

representative mature shells from 7 populations within western-central Portugal (Coimbra to Serra

da Arrábida). N = number of shells.

Tabla 111. Anchura relativa del ombligo en conchas de Oestophora barbula, O. barbella j/ una pobla¬

ción de O. cf. barbula de cerca de S. Pedro de Moel (Estremadura). Los datos de O. barbula se basan en

20 conchas de individuos maduros seleccionadas en 6 poblaciones, los de O. barbella en 22 conchas de

individuos maduros de 7 poblaciones en el oeste y centro de Portugal (Coimbra hasta Serra da Arrá¬

bida). N = número de conchas.

cribed by Seixas were O. lusitanica or O.

barbula sensu lato, since the shell descrip-

tion does not mention teeth. Her

drawing of the genital anatomy (Seixas,

1976: 36, Fig. 8D) is also unclear, with

the organ labelled "p."[enis] apparently

representing the accessory sac on the

vagina and henee no distal male genita¬

lia were shown. In view of these uncer-

tainties the information given by Seixas

seems best disregarded.

Shell characters. Shells of O. barbula

and O. barbella have been compared

closely using samples identified from

epiphallus structure. The most consis¬

ten! difference is that O. barbula has a

Sharp to very sharp peripheral keel on

the body whorl of adult shells, whereas

most O. barbella have the periphery

lacking a keel (slightly angled to bluntly

keeled or rounded) (Figs. 3, 4). Only a

very small minority of shells of O. barbe¬

lla have a keel sufficiently developed to

allow confusión with shells of O. barbula

(e.g. some shells from Verride, Beira

Litoral: Fig. 40).

Castillejo et al. (1987) found that

development of a peripheral keel is quite

variable within some local populations in

the Serra da Estrela (Beira Alta) that we

interpret as being composed entirely of

O. barbella. Matos (2004: 105) claimed

that populations of O. barbula [sensu lato],

both wild and reared under laboratory

conditions, frequently show shells with a

more or less strong equatorial keel. The

proportion of keeled individuáis was

said to vary around 20-28% in a single

population or in the total population,

these valúes corresponding to typical

Mendelian segregation of a recessive

gene, according to results then awaiting

publication. With O. barbella we have not

found any clear demarcation between

keeled, slightly keeled, slightly angled

(commonest type) and rounded shells in

most populations, whereas O. barbula is

consistently keeled. Puente (1996: 88)

mentioned shells as being clearly

rounded in five Spanish populations [of

the taxon we regard as O. barbella] and

clearly angular in one population (from

Las Correderas in Prov. Jaén).

The only other shell character likely

to be helpful in distinguishing the two

species is that most populations of O.

barbula have a proportionately narrower

umbilicus than most populations of O.

barbella, at least within western-central

Portugal (Table III). With careful measu-

rements of adult shells (see above),

umbilicus widths of less than 18% of

shell breadth were found only in O.

barbula, whereas those above 20% were

found mainly in O. barbella. The small

difference in umbilicus width is appa-

23

Iberus, 30 (1), 2012

rently due mainly to the tendency for O.

barbula to have the body whorl wider in

proportion to its height, as reflected in

the tendency for the shell mouth to be

more elliptical in O. barbula, more

rounded in O. barbella. Most shells from

many populations of O. barbella show an

umbilicus that becomes asymmetrical

during growth because of the form of

the body whorl, whereas the smaller

umbilicus of O. barbula is usually more

symmetrical. However, individual shells

often contradict these generalisations.

Detailed data on shell sizes and

number of whorls is presented in Table

II, for populations that have been identi-

fied from the epiphallus structure and

which inelude >6 adult shells, so that

mean sizes, size ranges and standard

deviations can be estimated with at least

modest reliability. Shell size varies

widely from locality to locality in both

species, although O. barbella has popula¬

tions with both smaller and larger mean

shell breadths than O. barbula (Table II).

It is difficult to detect any consistent

regional patterns in the distribution of

mean shell size for either species, with lit-

tle hint of any orderly pattern in size or

shape that might be used to define geo-

graphical subspecies. The most obvious

overall trend is apparently towards small

shell size in drier local habitats or regions

and larger shell size in wetter or more

humid situations. This is most obvious in

O. barbella, where the two samples with

smallest shell breadth (P88, PlOO) are

from areas of eastern Beira Baixa with

low regional rainfall and hot dry sum-

mers, followed by two samples from Co¬

astal sand dunes (P76, P97), then three

samples from open karst areas on Meso-

zoic limestone hills (P128 and P129 on Be¬

rra da Arrábida, P119 from Berra de Bao

Miguel). The largest shells are from sites

on hills of slate and other siliceous rocks,

with relatively high rainfall. A review by

Goodfriend (1986) similarly found that

within a land-snail species larger shell

size is often associated with moist envi-

ronments. Bmaller size was also someti-

mes recorded at higher population densi-

ties, apparently through effeets of phero-

mones on growth rates, but the latter

tendency was not apparent in these Oes-

tophora species. Reduced size of O. bar-

bula living sympatrically with O. barbella

at one of the sites in the Berra da Arrá¬

bida might be a consequence of interspe-

cific competition, as discussed below.

The ratio of shell height (H) to shell

breadth (B) overlaps very widely in O.

barbula and O. barbella, population

means being within an overall range of

H = 0.42-0.49 B in the former species,

0.42-0.50 B in the latter species. The

number of shell whorls tends to be

slightly higher in O. barbula (overall

range 5. 0-6. 3, population means 5.35-

5.92) than in O. barbella (overall range

4. 3-6. 2, population means 4.76-5.76)

(Table II), but there is much overlap and

a strong tendency within populations of

both species for smaller shells to be

comprised of fewer whorls. Develop-

ment of the palatal teeth is rather varia¬

ble, with the inner tooth being absent in

a small percentage of O. barbella, but

apparently always being present in

mature O. barbula.

Although shells of both species seem

at first sight to have rather variable

surface sculpture, cióse study at high

magnification (up to x40) with a stereo-

microscope suggests that both have a

very similar basic pattern that does not

allow species identification. The surface

of the protoconch is virtually smooth,

the same lack of sculpture continuing on

whorl 0-0.3. Whorl 0.3-0. 7 has irregular

low radial ribs, formed by elongate

papillae. Whorls 0.7-1. 4 have about 8-12

raised ridges, which show some ten¬

dency to spiral around the shell whorl

but soon descend into the lower suture;

around whorls 1.4-1. 6 these ridges are

reduced to spiral lines of low scales.

After whorl 1.6 the sculpture of the

mature teleoconch develops, with incre-

asingly prominent, long, low radial ribs,

with small raised papillae on a rectili-

near grid in the valleys between the ribs.

Much of the apparent variability in

sculpture on samples of mature shells

results from differences in the strength

(height) of ribs on later whorls of the

teleoconch and the extent to which these

are developed on the underside of the

24

Holyoak and HolyoaK: Taxonomic revisión of Oestophora barbula and O. barbella

Figure 3. Shells of type material of Oestophora barbula (Rossmássler, 1838) and O. barbella

(Servain, 1880), A-C: lectotype of O. barbula (SMF 7036); D-F: paralectotype of O. barbula

(SMF 7035); G-I: lectotype of O. barbella (MHNG 17059A).

Figura 3. Conchas del material tipo de Oestophora barbula (Rossmássler, 1838) y O. barbella

(Servain, 1880). A-C: lectotipo de O. barbula (SMF 7036); D-F: paralectotipo de O. barbula (SMF

7035); G-I: lectotipo de O. barbella (MHNG 17059A).

25

Iherus, 30 (1), 2012

Shell. The small papillae between the

teleoconch ribs soon disappear in worn

or corroded shells.

Geographical distribution. Records of

O. bartula and O. barbella from Portugal

are mapped in Figure 5. O. bartula con-

firmed by dissection was recorded only

from western-central Portugal, from

near Coimbra (Beira Litoral) south-

wards to the Serra da Arrábida (Estre-

madura), occurring inland only to the

eastern limits of the limestone hills of

SE. Beira Litoral, westernmost Beira

Baixa and N. Ribatejo provinces. O. bar-

bella has a much wider range, appa-

rently throughout Portugal, since it is

likely that the lack of records from much

of Alentejo and parts of northern-central

Portugal merely reflects sparsity of

recent recording of land snails there.

Since O. bartula appears to be endemic

to Portugal, the many records mapped

by Puente (1996: 87) for O. barbilla

[sensu lato] from western and Southern

Spain can probably all be referred to O.

barbella. Her published figures of genital

anatomy (Puente, 1996: 84-85) confirm

O. barbella as occurring in the following

Spanish provinces: Asturias, Cáceres,

Huelva, Jaén, León, Lugo and Zamora.

Many published figures of unkeeled

shells from Spain also imply that O. bar-

bella occurs widely there. Unkeeled

shells figured from the Azores (Back-

HUYS, 1975), two shells seen from Ter-

ceira, Azores and figures of Madeiran

shells (Seddon, 2008) also strongly

suggest it is O. barbella rather than O.

bartula that occurs in these archipelagos,

where it is likely to have been introdu-

ced by man. A population established

near Driebergen, province of Utrecht,

the Netherlands from at least 1994 to

2010 (Dees, 2011) also has unkeeled

shells apparent from published photos.

However, Quaternary fossil specimens

from Mallorca that have been reported

as O. barbilla are now reinterpreted as an

extinct endemic species named as Darde-

ria bellverica by Altaba (2006).

Habitat preferences. Descriptions of

habitats where the present authors co-

llected specimens of known identity

(mainly confirmed anatomically) were

recorded from 2001-2011 at 81 localities

in Portugal or neighbouring provinces of

Spain. Only one locality had both species

living together (see next section). Both

species occur mainly in the lowlands

and on low hills. Altitudinal ranges were

recorded for O. barbilla as ca 13 m (1 km

SW. of Alfarelos, Beira Litoral) to 548 m

(high on Serra de Sico, Beira Litoral); for

O. barbella as 7-10 m (on Coastal sand du-

(Right page) Figure 4. Shells of (A-F) Oestophora bartula (Rossmássler, 1838), (G-I) O. cf. bartula

and (J-O) O. barbella (Servain, 1880) to show variability between populations. A-C: 19 July 2010,

ca 1 km SW. of Serró Ventoso, Estremadura, from restricted type locality; D-F: 12 Feb. 2011,

Serra da Arrábida, Estremadura (29SNC020602), from population coexisting with O. barbella, see

Figures 4J-L; G-I: 4 June 2011, 2 km NNE. of S. Pedro de Moel, Estremadura, from apparently

atypical population of O. bartula with small unkeeled shells; J-L: 12 Eeb. 2011, Serra da Arrábida,

Estremadura (29SNG020602), from population coexisting with O. bartula, see Figures 4D-F; M-

O: 3 Nov. 2010, NW. ofVerride near Rio Mondego, Beira Litoral, from population with shell keel

more developed than in most O. barbella (all specimens in Collection of G.A. and D.T. Holyoak).

(Página derecha) Figura 4. Conchas de (A-F) Oestophora Farbula (Rossmássler, 1838), (G-1) O. cf.

Farbula y (J-O) O. barbella (Servain, 1880) para mostrar la variabilidad entre poblaciones. A-C: 19

jul. 2010, aproximadamente 1 km. SO de Serró Ventoso, Estremadura, de la localidad-tipo restringida;

D-F: 12 feb. 2011, Serra da Arrábida, Estremadura (29SNC020602), de la población conviviendo

con O. barbella, véase Eiguras 4J-L; G-F. 4 jun. 2011, 2 km. NNE de S. Pedro de Moel, Estremadura,

de la población aparentemente atipica de O. Farbula con pequeñas conchas sin quilla; J-L: 12 feb.

2011, Serra da Arrábida, Estremadura (29SNC020602), de la población conviviendo con O. Farbula,

véase Figuras 4D-F; M-0: 3 nov. 2010, NO. de Verride cerca de Río Mondego, Beira Litoral, de la

población con la quilla de la concha más desarrollada que en la mayoría de O. barbella (todos los ejem¬

plares en la colección de G.A. y D.T. Holyoak).

26

Holyoak and HolyoaK: Taxonomic revisión of Oestophora bartula and O. barbella

27

Iberus, 30 (1), 2012

Figure 5. Distribution of (A) Oestophora barhula (Rossmássler, 1838) and (B) O. barbella (Servain,

1880) in Portugal and adjoining regions of Spain. •: collected since 1999 by authors or Alvaro De

Oliveira, confirmed by dissection; •: collected since 1999 by authors or Alvaro De Oliveira, shells

only; O: collected before 2000, confirmed by dissection (HesSE, 1931 or CASTILLEJO ET AL.,

1987); O: collected before 2000, shells only (mainly confirmed by Alvaro De Oliveira).

Figura 5. Distribución de (A) Oestophora barbula (Rossmássler, 1838) y (B) O. barbella (Servain, 1880)

en Portugal y en las regiones cercanas de España. •; recogidos desde 1999 por los autores o por Alvaro de

Oliveira, confirmados por la disección; %: recogidos desde 1999 por los autores o por Alvaro de Oliveira,

solo eonchas; O.- recogidos antes de 2000, confirmados por la disección (Hesse, 1931 o CASTILLEJO ET

AL., 1987); O: recogidos antes de 2000, sólo conchas (principalmente confirmados por Alvaro de Oliveira).

nes) up to 890 m (Foia, Algarve), 1015 m

(near summit of Pico Sao Mamede, Alto

Alentejo), 1050 m (Sabugeiro, Beira Alta)

and at least 1080 m (N. of Lamas de

Mouro, Minho). Both species were recor-

ded in wooded and open sites, living

specimens most often being found res-

ting beneath substantial boulders.

O. barbilla was found at 21 localities,

20 of which were on bedrock of Cretace-

ous or Jurassic limestone, mainly where

the rock was exposed in crags or as

limestone pavements or large boulders,

but predominantly in dry-stone walls at

a few sites. At a single locality {ca 2 km

NNE. of Santa Cita, Ribatejo) it was

found living on alluvial banks of a

lowland river, albeit within a river

catchment having much limestone

bedrock. At the latter locality limestone

boulders had been imported to provide

protection for parts of the river bank, so

it is possible that O. barbilla reached the

site with these or that it arrived there by

drifting down the river. Possible occu-

rrence of a second population of this

species in a different habitat (at site

P67/P161) is discussed above.

28

Holyoak and HolyoaK: Taxonomic revisión of Oestophora barbula and O. barbella

O. barbella was recorded at 60 locali-

ties, over a much wider range of rock

types than those where O. barbilla was

found, including acidic lithologies. Sedi-

mentary rock types characterised less

than half of the localities, with records

from sites with exposed limestone rock

(10 sites; 16.7%), man-made banks of

limestone boulders (2 sites; 3.3%),

sandstone (3 sites; 5.0%), and unconsoli-

dated sediments of calcareous sand in

Coastal dunes (8 sites; 13.3%) and sands

and gravel capping limestone near the

coast (1 site; 1.7%). A majority of records

were from localities with various meta-

morphic rocks, recorded as slates

(mainly Paleozoic) (11 sites; 18.3%), slate

and shale (2 sites; 3.3%), slate and schist

(3 sites; 5.0 %), slate and quartzite (1

site; 1.7%), quartzite (5 sites; 8.3%),

granite (11 sites; 18.3%) and Foiaite (2

sites; 3.3%). The single remaining site

(1.7%) was along a concrete roadside

Wall with no rock exposed nearby. Some

sites with O. barbella were more distur-

bed by man than those tolerated by O.

barbula, including edges of cultivated

land and managed road verges.

It is noteworthy that although O.

barbella was recorded at 12 sites with

limestone rocks, as noted above, the

species was normally absent from places

with exposed Cretaceous or Jurassic

limestone within the range of O. barbula

(i.e. from Lisbon north to around

Coimbra: Fig. 5). Nevertheless, over that

extensivo región O. barbella was found

on man-made banks of limestone boul¬

ders near rivers (2 sites) and on Coastal

dunes (3 sites), so it appears to either

avoid the rocky sites with natural Meso-

zoic limestones or to be excluded from

them by the presence of the relatively

common O. barbula. Further evidence

that competitivo exclusión might occur

is that outside the range of O. barbula, O.

barbella was found to occur widely on

outcropping Mesozoic limestone. In the

Algarve it was usually found (7 sites) at

much lower density than the accompan-

ying Gittenbergeria turríplana (Morelet,

1845) (Trissexodontidae), an Algarve

endemic with a largor sharply keeled

shell; it was also found among hard

limestone rocks in the Sierra de la

Encina (Prov. Ourense) (1 site). Where

O. barbella meets the Southern edge of

the range of O. barbula in the Serra da

Arrábida (Estremadura), O. barbella was

found alone at one site (in plenty), O.

barbula was found alone (in plenty) at

two sites and both species coexisted at a

fourth site, as follows.

Coexistence of O. barbula and O. bar-

bella. Although the ranges of O. barbula

and O. barbella approach closely and pro-

bably interdigitate around Coimbra and

Figueira da Foz (Boira Litoral), they were

not found living together and their re¬

cords were from different habitats, as

described above. Flowever, at one of four

sites studied on the Serra da Arrábida

(P129 at 29SNC020602) the two species

were found living together, at 380-388 m

altitude on N.-, NW. and S.-facing rocky

limestone slopes with patches of scrub.

Visits on 17 May 2007 and 12 February

2011 amassed 22 adult O. barbula (4 li¬

ving, 18 shells) and 16 adult O. barbella (8

living, 8 shells) from within a 100 metre

radius, both species twice being found

together alive under a single boulder.

The same two visits to the Serra da Arrá¬

bida revealed O. barbella occurring alone

at one site (P128 at 29SNC032604, 45

adults, from rocky limestone hilltop with

low scrub) and O. barbula occurring alo¬

ne at two other sites (P130 at

29SMC999576 to 29SNC000577, 68

adults, limestone road-cutting, scree and

slope with tall scrub locally; P131 at

29SNC01175874, 12 adults, roadside at

base of cutting in limestone rock). The

identity of specimens from all four sites

was confirmed by study of the epipha-

llus (in several individuáis from three si¬

tes, one from the fourth) and all adult

shells were easily separated by the pre¬

sence of a Sharp peripheral keel in O.

barbula alone (cf. Figs. 4D-F and 4J-L).

It is noteworthy that at these four

sites in the Serra da Arrábida shell size

of both species is rather similar where

each occurs alone, but it diverges mar-

kedly where the two species coexist

(Table II, Figs. 4D-F and 4J-L). Thus,

mean shell breadth for O. barbella occu¬

rring alone (site P128) is 9.58 mm, and

29

Iberus, 30 (1), 2012

for O. bartula occurring alone it is 9.65

mm (site P130) and 10.34 mm (site

P131). At site P129 with both species

living together, O. barbella is larger at

10.63 mm whereas O. bartula is smaller

at 8.51 mm.

Based on data listed in Pable II, sta-

tistical significance of the differences

among sample means for shell breadth

was calculated pair-wise for sites P128-

P131. After it was established with Sne-

decor's F that there is no significant diP

ference of the respective variance ratio

between each of the pair of samples

compared (p >0.05), valúes of Student's

t were calculated applying BessePs

correction. These results confirm that at

site P129 O. bartula is smaller than at

neighbouring sites (P130: t = 8.94, 86

d.f., p <0.001; P131: t = 8.47, 31 d.f., p

<0.001) whereas O. barbella is larger than

at a neighbouring site (P128: t = 6.18, 42

d.f., p <0.001); they also confirm the size

difference between the two species at

site P129 {t = 3.02, 36 d.f., p <0.01).

Furthermore, the small size of O.

bartula at site P129 appears to be a

special effect of sympatry with O, barbe¬

lla rather than due to chance alone, since

this is much the smallest of 30 popula-

tions of O. barbilla encountered anyw-

here in this study (cf. Pable II), with the

possible exception of specimens from

P67/P161 (see above) which have unke-

eled shells and somewhat ambiguous

genital anatomy so some doubt remains

about the species identification.

Phese differences in shell size

between the two species are suggestive of

the phenomenon of "character displace-

ment" (Brown and Wilson, 1956;

PiANKA, 1974), which refers to increased

differences between species where they

occur together. It is argued that in such

cases selective pressures from interspeci-

fic competition have tended to maximise

the differences in size and henee in

resource utilisation among coexisting

species (Brown, 1975; Diamond, 1975).

Phis presumes that differences present in

sympatry are inherited and allow an evo-

lutionary response. Fenchel (1975) pro-

posed this explanation for size differen¬

ces developed between some mud-snail

species (Hydrobiidae) only when they

coexist, but Saloniemi (1993) proposed

an environmental explanation whereby

only those habitat types able to support

two coexisting mud-snail species allow

one of the species to grow large. Subse-

quently, experimental studies by Gor-

BUSHIN (1996) pointed to the size diffe¬

rences in sympatry being an effect of

direct competition for food, affecting one

of the species more than the other, not an

inherited adaptive response. In fact, most

examples of "character displacement"

involve birds or other vertebrates where

size differences in adults of two species

can allow partitioning of resources. As

pointed out by Hylleberg (1976) and

stressed by Gorbushin (1996) it is unli-

kely that coexisting species of snail can

partition resources in the same manner

because a large part of their populations

will be comprised of immatures that

show extensivo size overlap between

species. Henee, 'Aharacter divergence" in

sympatric snails may be evidence of

interspecific competition, but need not

involve inherited size differences.

TAXONOMIC DISCUSSION

Authorship, lectotypification and

type-locality of O. bartula

Hidalgo (1875: 42) cited the ñame

Helix bitiiberculata A. Férussac, 1837,

from the catalogue published in Paris

for the sale of the Férussac Collection

(on p. 4, from Portugal). This ñame was

evidently a nomen nudum that Hidalgo

interpreted as representing [Helix]

bartula Charpentier.

Rossmássler (1838: 11, Taf. 32, Fig.

451) gave the first description and

figures of the species, as Helix bartula v.

Charp., with "Syn. Helix bartula v. Charp:

in litt/' also listed. Subsequently, Hesse

(1931: 52) listed the authorship of the

ñame as "Oestophora bartula (Chrp.)

Rssm." and Ortiz de Zárate López

(1962: 91), Castillejo (1984: 128) and

Castillejo et al. (1987) all adopted

''Oestophora (Oestophora) bartula (de

Charpentier)". However, Puente (1996:

82) accepted Rossmássler as author and

30

Holyoak and HolyoaK: Taxonomic revisión of Oestophora barbula and O. barbella

this appears to be correct on the basis

that Charpentier presumably provided

the ñame whereas Rossmássler was res-

ponsible for the description, so only the

latter can be regarded as the author res-

ponsible for its naming.

The collection of the Forschungsins-

titut Senckenberg contains two shells

labelled as those of Helix barbula figured

by Rossmássler {loe. cit.), one more

sharply keeled (SMF 7036, shell diame-

ter 11.1 mm: Figs. 3A-C) than the other

(SMF 7035, diameter 11.9 mm: Figs. 3D-

F). SMF 7035 had been labelled as "lec-

totype" by the late Dr. Adolf Zilch, but

the designation is invalid since it was

never published. Cióse study of

Rossmássler 's fig. 451 {loe. eit.; espe-

cially the first of the four figures) sug-

gests the figured specimen was actually

the more sharply keeled SMF 7036.

Since SMF 7036 also shows the clearest

development of the shell keel characte-

ristic of the taxon with a large epipha-

llus from western-central Portugal this

shell is selected here as lectotype,

leaving SMF 7035 as a paralectotype.

Rossmássler (1838: 11) recorded the

provenance of the type material as "von

Holl auf seiner Aktien-reise in Portugal

gesammelt". Searches have produced no

additional Information to indicate a more

precise locality. This is unfortunate

because a small proportion of shells of O.

barbella appear indistinguishable from the

lectotype of O. barbula. We cannot desíg¬

nate a neotype for O. barbula based on a

specimen with large penial epiphallus

because undoubted syntype shells still

exist (ICZN Art. 75(a)). Although the

ICZN could itself set aside the existing

type material and desígnate a neotype

under its plenary powers if a detailed

application were to be made (Recommen-

dation 75E), it would need to be demons-

trated that such action "is essential for

solving a complex zoological problem"

(Art. 75). Mayr (1969: 375, 56.B.3) conclu-

ded that "in most cases [of this type] no

such neotype designation will be

needed", adding (Mayr, 1969: 374-375,

56.B.3) that the ICZN Code "contains no

provisions regarding what to do when

the type lacks all diagnostic characters".

A convenient solution to this pro¬

blem lies in restricting the type-locality,

which was given only as "Portugal" in

the original description of O. barbula

and has never before been restricted, eit-

her directly, or by our designation of a

lectotype. Following ICZN Recommen-

dation 72H on restricting type localities,

only its section (4) can be applied: "as a

last resort, and without prejudice to ot¬

her clarification, localities within the

known range of the taxon or from which

specimens referred to the taxon had

been taken". Henee we restrict the type-

locality of O. barbula to "limestone crags

ea 1 km SW. of Serró Ventoso, S. of Porto

de Mós, Estremadura, in central Portu¬

gal". This locality exeludes any like-

lihood of O. barbella being present, whe¬

reas anatomically verified O. barbula

with large keeled shells (Figs. 4A-C) re-

sembling that of the lectotype are abun-

dant there. Although this action is arbi-

trary with respect to Rossmássler 's ma¬

terial, a first reviser may designate a

restricted type locality. Such a restriction

may later be set aside if it confliets with

the available evidence, but only if the

case is unequivocal (Mayr, 1969: 377,

note 2).

Authorship and lectotypification of

O. barbella

Type specimens of Helix barbella

(Servain, 1880: 66) were studied on loan

from Muséum d'Histoire Naturelle,

Ville de Genéve (MHNG). These notes

describe this material, set out reasons

for believing that it forms all or part of

the original "type series" and provide a

background to selection of a lectotype

for the species.

The type material studied comprised

two similar corked glass tubes with

round bases, each of them labelled on

the outside with a blue edged label

annotated as follows in ink:

Helix barbella Servain Mertola Portu¬

gal [11 shells; subsequently numbered at

MHNG as 17059; cork with 51. written

in ink].

Helix barbella Servain Lisbonne [1

shell; subsequently numbered as 17060;

cork with 34. written in ink];

31

Iberus, 30 (1), 2012

These two tubes are contained in a

small card box that is glued to a larger

wooden base. A label pasted on the

bottom of the inside of the box States:

Helix Barhella, Bourguignat in: Servain,

Cat. Molí. Esp. p. 1880. A label pasted

outside the box on the top of the

wooden base has printed "Muséum de

Genéve Collection Bourguignat" and

hand-written Helix barhella Bgt. [Loca-

lité] Portugal.

The box and its two labels evidently

date from the curation of the material as

part of Bourguignat's collection at

MHNG. Their attribution of authorship

to Bourguignat is probably erroneous

because the labels on the two tubes give

only Servain as author of the ñame, and

this is in agreement with the original

publication having Servain as solé

author and also the original description

making no mention of Bourguignat.

It can be inferred that these speci-

mens from Bourguignat's collection

form all or part of Servain's "type

series" of Helix barhella for the following

reasons, which taken together, leave no

real doubts:

(1) The original collection of G.

Servain has apparently not survived

intact. At least, the list of collections by

Dance (1986: Appendix IV) makes no

mention of it.

(2) There is clear evidence that

Servain worked closely with Bourguig¬

nat and apparently made at least some

of his Spanish and Portuguese speci-

mens available to him. The book by

Servain (1880: 6) makes a lavish ack-

nowledgement of Bourguignat (and no

other collaborater) stating: "á notre

excellent ami M. J. R. Bourguignat, nos

remerciements pour son extréme oblige-

ance, qui ne s'est jamais démentie á

notre égard, et pour la complaisance

avec laquelle il a bien voulu nous éclai-

rer sur la valeur de nos espéces et mettre

á notre disposition sa riche bibliothéque

et son immense collection."

(3) Types of other Portuguese snails

described in Servain (1880) appear to be

in the Bourguignat collection. Thus, in

dealing with Candidula, Gittenberger

(1993: 283) points out that "Topotypes of

nominal taxa described by G, Servain,

found in the collection of J.R. Bourguig¬

nat, might in fact be syntypes donated

by their author."

(4) The two localities given in the

original description of H. barhella

(Servain, 1880: 67) and those on tubes in

the Bourguignat collection correspond

exactly and the number of specimens

from each of them also corresponds

well. Thus, Servain wrote "Alluvions du

Tage prés de Lisbonne, oü elle est rare. -

Elle est assez ahondante, á ce qu'il

parait, aux alentours de Mertola en Por¬

tugal." The material of H. barhella in

Bourguignat's collection comprises just

two tubes, respectively from "Lisbonne"

[1 Shell] and "Mertola" [11 shells].

(5) The characters of the shells in the

Bourguignat collection show unusual

features which correspond exactly to

parts of Servain's original description.

Thus, Servain (1880: 67) States that there

is only a single palatal tooth ("orné seule-

ment d'iine grosse dent vers la région

médiane du borde externe": italics as in

original). The Mertola material at MHNG

(11 shells) has one shell with only a single

tooth (outer palatal only), four shells

giving a superficial impression of having

only a single tooth (with a very weak or

weak and rather inconspicuous inner

tooth), one shell with the inner tooth

almost obscured by the dried body;

henee only the remaining five shells have

a conspicuous and strong inner palatal

tooth. The single shell at MHNG from

"Lisbonne" also has only a small, weak

inner palatal tooth. Occurrence of only a

single palatal tooth is recurrent but rare

in populations of the O. barbula/0. barbe-

lla group of taxa (Ortiz de Zárate

López, 1962; Castillejo et al, 1987;

Puente, 1996).

In the original description, Servain

(1880: 67) also stated that the shell has

"son dernier tour aussi fortement lamellé en

dessous qu'en dessus" (italics as in origi¬

nal), i.e., the body whorl has ribbing on

the underside as strong as that on the

upperside. Ribbing in the MHNG mate¬

rial is present on the underside of all 12

shells and strong in some of them; five of

the shells from Mertola have the ribbing

32

Holyoak and HolyoaK: Taxonomic revisión of Oestophora bartula and O. barbella

on the underside as strong or nearly as

strong as that on the upperside. Gene-

rally, ribbing of shells in this species-

group is rather variable, but it is infre-

quently as strong on the underside of the

body whorl as it is on the upperside.

(6) The condition of the shell at

MHNG from 'Tisbonne" corresponds to

the "Alluvions du Tage" of the original

description: it is encrusted with silt on

the outside of the spire and inside the

shell mouth.

Overall, it seems there can be no cer-

tainty that all of Servain's original "type

series" is now in the Bourguignat collec-

tion. However, the evidence that at least

part of it is present there is compelling.

Since the original Servain collection of

shells no longer appears to exist, there is

thus ampie justification for selecting a

lectotype from the Bourguignat collec¬

tion at MHNG.

Therefore, a lectotype is designated

here to fix application of the ñame Helix

barbella Servain, 1880. The lectotype

selected is from Mertola (i.e. Mértola)

and is placed in a sepárate small tube

and labelled as MHNG 17059 A (Figs.

3G-I); it measures 11.2 mm in diameter,

4.5 mm in height and has 5.6 whorls, a

weak inner palatal tooth and strong

ribbing on the underside of the shell;

traces of the body and dark staining

suggest the animal was alive when

collected. Ten paralectotypes from

Mértola remain in the original tube (as

17059B); these have shell diameter 9.2-

11.3 mm (mean 10.33, s.d. 0.79 mm),

shell height 3.8-4.9 mm (mean 4.29, s.d.

0.39 mm), 4.8-5. 7 whorls (mean 5.27, s.d.

0.24 whorls). One paralectotype from

"Lisbonne" (17060) remains in its origi¬

nal tube; it has shell diameter 10.1 mm,

shell height 4.7 mm and 5.4 whorls. All

the shells of the "type series" of O, bar-

bella have a rounded keel on the body

whorl, less sharp than in either of the

two syntypes of O. barbula Rossmássler.

The genital anatomy has been studied

from two specimens of O. barbella collec¬

ted recently at Mértola (20 Oct. 2008,

Cerrinho das Neves, Mértola

(29SPB1767), Mértola, Baixo Alentejo, leg,

Alvaro De Oliveira; Collection of G.A.

and D.T. Holyoak); both show a relati-

vely short and weak penial epiphallus

(Fig. 2H). The shells from these two spe¬

cimens have been compared with the lec¬

totype and ten paralectotypes from

Mértola, which they resemble in the

rounded keel on the body whorl and

general form of the shell. These two

recently collected shells are within the

range of variation of the paralectotypes

in their shell dimensions, weak/ médium

strength of ribbing on underside of body

whorl and the médium /strong inner

palatal tooth.

ACKNOWLEDGEMENTS

This study originated in a proposal

from Alvaro De Oliveira to collaborate

in studying the remarkable variability

within shells of Oestophora barbula from

Portugal. It eventually became mainly a

study of characters of the genital

anatomy and identity of type material

undertaken by the present authors, but

we were greatly assisted in this by pro¬

visión of numerous specimens collected

by Alvaro, use of his data from shell

collections in Portuguese museums and

many other helpful comments and dis-

cussions. Thanks are also due to Dr

Ronald Janssen of the Forschungsinsti-

tut Senckenberg (SMF) for information

on Rossmássler 's type material of Oes¬

tophora barbula and for permitting use of

images of these specimens prepared by

Mrs S. Hof. Dr Yves Finet of the

Muséum d'Histoire Naturelle, Ville de

Genéve kindly assisted with enquiries

about Servain's type material of O. bar-

bella and allowed study of it on loan.

The curators of Museu de Historia

Natural de Coimbra, Museu Nacional

de Historia Natural (Universidade de

Lisboa) and Museu de Historia Natural

da Universidade do Porto kindly

allowed Alvaro De Oliveira the access to

specimens in their care that contributed

to this study. Ruud Bank provided

helpful comments on authorship of

ñames. Comments from an anonymous

referee significantly improved this

paper.

33

IberuSy 30 (1), 2012

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35

Iberus, 30 (1), 2012

APPENDIX. MATERIAL STUDIED.

Data are listed in sequence as country, province, locality ñame, habitat description,

altitude, U.T.M. grid reference, date (YYYY.MM.DD), collector(s) followed by - then

collector's field number, number of specimens, number of shells, collection, museum

registration number, comments; successive entries are separated by semi-colons.

Abbreviations: ÁDO = Alvaro De Oliveira, CGAH = Collection of G.A. and D.T.

Holyoak, DTH = D.T. Holyoak, GAH = G.A. Holyoak, leg. = collected by, MHNG =

Muséum d'Histore Naturelle, Ville de Genéve, Switzerland, SMF = Forschungsinstitut

Senckenberg, Germany, sh = number of intact adult shells (without preserved bodies),

sp = total number of adult specimens with bodies preserved in alcohol (number of

dissected adult specimens is given in parentheses).

O. harhella, Portugal, Algarve: ca 0.5 km S. of Bensafrim, base of N.-facing limes-

tone crag, 76 m, 29S 05237/41116, 2011.02.06, GAH and DTH - P121, 2 sh, CGAH; W.

slope of Foia, hillslopes with low metamorphic crags (Foyaite) and patchy cover of

Erica and Cistus, grassland locally, 810 m, 29S 05347/41292, 2011.02.05, GAH and DTH

- P120, (1) sp, CGAH; ca 0.5 km E. of summit of Foia (W. of Monchique), hilltop with

blocks of metamorphic rock amongst low scrub, ca 890 m, 29SNB33, 2001.06.03, GAH,

5 sh, CGAH; E. end of Rocha da Pena, rocky limestone slope and platean with patches

of bushes, 460 m, 29S 05803/41234, 2011.01.28, GAH and DTH - P104, (1) sp, 4 sh,

CGAH; E. end of Rocha da Pena, rocky limestone slope with patches of tall bushes,

462 m, 29S 058033/412344, 2011.01.31, GAH and DTH - P112, (1) sp, CGAH; by N2 at

4.5 km NW. along road from Ameixial, roadside cutting with steep slaty/shaley rock

and patchy low herbs and grasses, 265 m, 29S 05898/413885, 2011.01.30, GAH and

DTH - Pilo, (1) sp, 7 sh, CGAH; ca 2 km E. of Santa Barbara de Nexe, rocky limestone

hill-slopes with patchy low scrub, 261 m, 29S 059428/410732, 2011.02.03, GAH and

DTH - P117, 1 sh, not kept; by N2 NNW. of Alportel, rocky (slate and sandstone)

slopes in open Piniis, Quercus and Eiicalyptus woodland by road, 365 m, 29S

059569/411731, 2011.01.28, GAH and DTH - P106, (2) sp, 9 sh, CGAH; by N124 W. of

Quinta, banks and steep bases of road cuttings in slaty/ schist rock, with sparse Cistiis,

grasses and herbs, 211 m, 29S 059037/412121, 2011.01.31, GAH and DTH - P113, 1 sh,

CGAH; by N2 S. of Ameixial, steep slaty/ shaley roadside cuttings with sparse patches

of grasses and herbs, 406 m, 29S 059164/413483, 2011.01.30, GAH and DTH - Pili, (1)

sp, CGAH; by N398 ca 2 km N. of Moncarapacho, limestone slope with low rocks and

patches of scrub, ca 121 m, 29S 06068/41078, 2011.01.29, GAH and DTH - P108, (1) sp,

1 sh, CGAH; Serra de Sao Miguel {ca 4 km NW. of Moncarapacho), rocky limestone

slopes with patchy scrub and grassland, 340 m, 29S 06043/41067, 2011.02.04, GAH

and DTH - P119, (1) sp, 7 sh, CGAH; ca 3 km NNW. of Moncarapacho (just S. of A22),

rocky limestone hillslope with patchy bushes, 131 m, 29S 060695/410771, 2011.02.04,

GAH and DTH - P118, (1) sp, 1 sh, CGAH; just NW. of Barroqueira {ca 7 km NW. of

Moncarapacho), rocky limestone slopes and quarries with patchy bushes, 183 m, 29S

060291/411092, 2011.01.29, GAH and DTH - P107, 2 sh, CGAH; Alto Alentejo: Belver

castle, Gaviáo, 29SND8972, 2008.11.21, ÁDO, (3) sp, CGAH; by Rib. de Sor ca 4 km

NW. of Monte da Pedra (NW. of Grato), granite rocks with sparse scrub near river, ca

183 m, 29SPD05, 2001.05.30, GAH, 7 sh, CGAH; N. end of Barragem da Povoa, granite

walls and rocks in deciduous grove on reservoir margin, 320 m, 29SPD27, 2001.05.31,

GAH, 1 sh, CGAH; Pico Sao Mamede, beneath boulders on sparsely vegetated N.-

facing hillside near crags, ca 1015 m, 29S 06413/43527, 2007.05.15, GAH - 2007.5, (1)

sp, 3 sh, CGAH; Azores: Duque da Terceira Garden, Angra do Heroismo, Ilha Terceira,

26NMH87, 2005.11.01, ÁDO, 2 sh, CGAH; Baixo Alentejo: by N2 at ca 1.5 km SSE. of

Dogueno, unshaded road cuttings with exposed sandstone, sparse low vegetation,

338 m, 29S 059045/414242, 2011.01.30, GAH and DTH - P109, 2 sh, CGAH; Cerrinho

36

Holyoak and Holyoak: Taxonomic revisión of Oestophora barbula and O. barbella

das Neves, Mértola, 29SPB1767, 2008.10.20, ÁDO, (2) sp, CGAH; Noudar Castle,

Barrancos, 29SPC6927, 2008.10.19, ÁDO, (3) sp, CGAH; Mertola, G. Servain, 1 sh,

MHNG 17059 A, Lectotype, 10 sh, MHNG 17059B, Paralectotypes; Beira Alta: Sabu-

geiro, bases of low granite crags and walls near river, 1050 m, 29T 06149/44733,

2010.08.31, GAH and DTH - P71, 3 sh, CGAH; by N16 on W. edge of Gongalbocas,

granitic slopes with much bare rock, patchy herbs and grasses, bushes locally, 814 m,

29T 065385/449325, 2011.10.14, GAH and DTH - P209, (3) sp, 7 sh, not kept; W. edge

of Moinhos de Aveia (W. of Alverca da Beira), rocky granitic slopes with patchy low

scrub, 554 m, 29T 06518/45071, 2011.06.26, GAH - P165, 1 sh, not kept; Beira Baixa: ca

l. 5 km W. of Lousa (S. of Vila de Rei), rocky (quarztite and slaty) slopes with patchy

scrub and few trees, 210 m, 29S 05747/43862, 2011.07.27, GAH and DTH - P174, 1 sh,

not kept; by Rio Zézere and N238 ENE. of Domes, rocky (slate) cuttings and slopes by

road and reservoir, building ruins, 143 m, 29S 05644/44037, 2011.08.05, GAH and

DTH - P177, (2) sp, 28 sh, CGAH; Barcoila, near Serta, garden walls, ca 410 m, 29S

05741/44096, 2010.09.11, GAH ^ P72, (1) sp, 6 sh, CGAH; W. edge of Serta, under

cardboard on roadside verge with patchy herbs, ca 240 m, 29S 05769/44065,

2010.04.22, GAH - P37, (3) sp, 2 sh, CGAH; Rio Zézere valley W. of Pedrógáo

Pequeño, rocky granitic slopes of valley side with open Piniis and Qiiercus woodland,

ca 250 m, 29S 05737/44176, 2011.04.27, GAH and DTH - P140, (2) sp, CGAH; by Rio

Ocreza and N241 E. of Vale da Mua, grassy banks near river, 29S 060554/439514,

2008.08.21, GAH -- P6, 1 sh, not kept; Portas de Almoráo, rocky (quartzite) slopes with

patchy scrub and herbs, 275 m, 29S 060681/439902, 2010.12.19, GAH and DTH - P86,

15 sh, CGAH; ca 2 km W. of Vila Velha de Ródao, rocky (quartzite) ridge and slopes

with patchy low scrub and herbs, 321 m, 29S 061177/439016, 2010.12.19, GAH and

DTH - P87, 5 sh, CGAH; by N240 on W. bank of Rio Ponsul (W. of Ladoeiro), S.-facing

rocky (sandstone) banks/cutting with patchy scrub and grasses, 183 m, 29S

064235/0441332, 2010.12.23, GAH and DTH - P88, (3) sp, 12 sh, CGAH; by N233 ca 3

km W. of Oledo, rocky (granite) slopes by road and river, with patches of trees and

scrub, 260 m, 29S 06413/44263, 2011.01.15, GAH and DTH - P99, 1 sh, CGAH; by

N353 just SE. of Idanha-a-Nova, rocky (granite) hillside with patches of scrub and low

trees, 288 m, 29S 065144/442069, 2010.12.27, GAH and DTH - P95, (1) sp, 4 sh, CGAH;

by N239 just SW. of Proenga-a-Velha, rocky (granite) slopes by road and river, with

patches of trees and scrub, 375 m, 29T 064964/443134, 2011.01.15, GAH and DTH -

PlOO, (3) sp, 5 sh, CGAH; Beira Litoral: Casal de Sao Simao, NE. of Aguda, quartzite

crags and slopes, grassland, walls of chapel, 385 m, 29S 05562/44188, 2011.09.06, GAH

and DTH - P192, 5 sh, not kept; ca 2 km NE. of Favacal, quartzite rocks and slopes of

hilltop, with grassland, bushes and concrete walls, 755 m, 29S 05576744272, 2011.09.06,

GAH and DTH P191, (1) sp, not kept; ca 1 km N. of Pedrogáo Grande, deciduous

groves by stream, roadsides, rocky banks, walls, 330 m, 29S 057223/442063,

2011.08.11, GAH and DTH - P179, 3 sh, not kept; just S. of S. Pedro (S. of Figueira da

Foz), Coastal sand dunes with patchy low scrub, ca 10 m, 29T 05117/44412, 2011.01.13,

GAH -- P97, (3) sp, 11 sh, CGAH; Praia da Tocha, sand dunes with patches of low

bushes, 14 m, 29T 05134/44647, 2010.11.18, GAH and DTH - P83, 2 sh, CGAH; ca 1

km NW. of Gafanha do Areáo (SW. of Aveiro), sand dunes, with much Acacia scrub, 8

m, 29T 05188/44862, 2010.10.26, GAH - P76, (3) sp, 28 sh, CGAH; ca 1 km NW. of

Verride near Rio Mondego, grassy banks with limestone boulders locally, 11 m, 29T

05244/44437, 2010.11.03, GAH and DTH - P78, 10 (4) sp, CGAH; just S. of Costa Nova

do Prado, sand-dunes with patchy herbs and low shrubs, 7 m, 29T 05209/44948,

2011.05.03, GAH and DTH - P144, (3) sp, 4 sh, CGAH; Vista Alegre, ílhavo, 29T

NE2793, 2009.10.26, ÁDO, (1) sp, CGAH; N. of Ameal, grassland and limestone boul¬

ders on bank of river, ca 12 m, 29T 05393/44502, 2010.11.03, GAH and DTH - P80, 11

(4) sp, 12 sh, CGAH; Quinta das Lágrimas, Coimbra, 29TNE4849, 2010.05.11, ÁDO, (1)

sp, CGAH; Botanical Garden, Coimbra, 29TNE4950, 2009.03.10, ÁDO, (3) sp, 1 sh,

CGAH; Instituto Geofisico, Coimbra, 29TNE5051, 2009.11.20, ÁDO, (1) sp, CGAH;

37

IberuSy 30 (1), 2012

Coja, Arganil, 29TNE8658, 2009.11.02, ÁDO, (1) sp, CGAH; S. Jacinto, Aveiro,

29TNF2303, 2008.08.30, ÁDO, (1) sp, CGAH; Cais do Puchadouro, Ovar, 29TNF3120,

2009.11.20, ÁDO, (1) sp, CGAH; Feira castle, Santa María da Feira, 29TNF3830,

2008.12.12, ÁDO, (3) sp, CGAH; Douro Litoral: just E. of Sao Félix da Marinha, walls

and plantation with Eiicalyptus, ca 105 m, 29T 05330/45424, 2007.05.20, GAH -

2007.17, 29 sh, CGAH; Botanical Garden, Porto, 29TNF3056, 2008.10.07, ÁDO, (3) sp,

CGAH; Scarp over Douro River, Porto, 29TNF3354, 2008.01.14, ÁDO, (3) sp, CGAH;

Estremadura: ca 1 km W. of Cela Velha {ca 6 km S. of Nazaré), roadsides, grassland

and grove of Populas, ca 9 m, 29S 049419/437824, 2011.06.04, GAH and DTH - P164, 1

sh, not kept; just E. of lighthouse at Nazaré, sandy and gravelly Coastal slope with

short patchy vegetation, 52 m, 29S 049282/438393, 2010.12.25, GAH and DTH - P91, 3

sh, CGAH; S. Bartolomeu, 2 km SE. of Nazaré, granitic hill with crags, wooded slopes

and oíd buildings, 155 m, 29S 04955/43825, 2011.06.04, GAH and DTH - P162, 7 sh,

CGAH; just SW. of Vale de Parades, narrow strip of dune vegetation at base of cliff

slope, 12 m, 29S 049561/439426, 2010.12.25, GAH and DTH - P92, 4 sh, CGAH; Pedras

Negras, ca 3 km N. of S. Pedro de Moel, dunes with patchy low grasses and herbs, 20

m, 29S 049813/440313, 2010.08.12, GAH and DTH - P68, 1 sh, not kept; Serra da Arrá¬

bida, N.- to NW.-facing limestone crags and rocky slopes, 380 m, 29S 05021/42601,

2007.05.17, GAH - 2007.10, 5 sh, CGAH, mixed with O. barbilla; Serra de Arrábida, N.-

and S.-facing rocky limestone slopes with patches of scrub, 388 m, 29S 05020/42602,

2011.02.12, GAH and DTH - P129, 8 (5) sp, 3 sh, CGAH, mixed with O. barbilla; Serra

da Arrábida, S.-facing limestone crags and rocky slope, 300-350 m, 29S 05032/42604,

2007.05.17, GAH - 2007.11, 15 sh, CGAH; Serra de Arrábida, high on rocky S.-facing

limestone slope with patchy low scrub, 351 m, 29S 05032/42604, 2011.02.12, GAH and

DTH - P128, 20 (3) sp, 10 sh, CGAH; Lisbonne, G. Servain, 1 sh, MHNG 17060, from

river floodline debris; Minho: Ofir (SE. of Esposende), low sand dunes with patchy

bushes, herbs and grassland, 9 m, 29T05175/45972, 2011.08.30, GAH and DTH - P190,

2 sh, not kept; Valenga fortress, Valenga, 29TNG2953, 2009.04.08, ÁDO, (2) sp, CGAH;

Mongáo fortress, Mongáo, 29TNG4258, 2009.01.30, ÁDO, (3) sp, CGAH; N. edge of

Peneda village, rocky granitic slope with oíd walls, patchy grasses and herbs, 688 m,

29T 056447/464734, 2011.08.29, GAH and DTH - P187, (1) sp, not kept; ca 2 km N. of

Lamas de Mouro, rocky mountain slope, facing S., sparse low vegetation, 1080-1100

m, 29T 05665/46570, 2011.08.29, GAH and DTH, 1 sh, not kept, oíd shell only; Trás-os-

Montes: Penas Roías Castle, Mogadouro, 29TPF9685, 2009.11.10, ÁDO, (1) sp, CGAH;

by IP2/N103-7 ca 2 km NNE. of Rabal, beside irrigation channels in Fraxinus grove

with few Quercus, 618 m, 29T 06873/46388, 2011.06.29, GAH - P170, 2 sh, not kept;

Braganga castle and Fervenga River, Braganga, 29TPG 8630-8730, 2009.11.11, ÁDO, (3)

sp, CGAH; by N218 ca 4 km due NW. of Vimioso, slopes of river gorge with crags

(slate and schist), screes and patchy scrub, 447 m, 29T 070334/460939, 2011.06.30,

GAH and DTH - P171, 2 sh, not kept; ca 1 km S. of Vale de Águia (NE. of Miranda do

Douro), exposed low and flat granitic rocks with boulders, patchy scrub and low ever-

green Quercus trees, 672 m, 29T 0729638/4600096, 2011.07.01, DTH - P172, (3) sp, not

kept; Spain, Prov. Cáceres: by EX117 at Rio Salor, NE. of Membrio, slopes of slaty rock

above river banks with patchy low vegetation, 159 m, 29S 067198/438245, 2010.12.23,

GAH and DTH - E89, (1) sp, 4 sh, CGAH; by EX302 at Rio Salor, NE. of Herreruela,

slopes of slaty rock above river banks with patchy low herbs and scrub, 209 m, 29S

068226/437263, 2010.12.23, GAH and DTH - E90, (2) sp, 1 sh, CGAH; Prov. La

Coruña: E. edge of P. N. de Corrubedo, exposures of low granite rocks with moss

locally and very sparse grasses, at edge of Pinas wood, 48 m, 29T 049808/471162,

2011.08.27, GAH and DTH - E183, 1 sh, not kept; Prov. León: by N536 ca 1 km S. of

Salas de la Ribera, low roadside cutting, crag and scree of slaty rock with sparse low

grasses and herbs (bushes above), 393 m, 29T 06796/46993, 2011.10.16, GAH and DTH

- E213, 5 sh, not kept; NW. of Almázcara, low banks between grassland and road,

above shallow ditch, with patchy grasses and herbs, bushes locally, 672 m, 29T

38

Holyoak and HolyoaK: Taxonomic revisión of Oestophora barbula and O. barbella

07035/47203, 2011.10.16, GAH and DTH - E212, 3 sh, not kept; Prov. Ourense: by

OU533 0.5 km S. of Santa Cruz, low granitic rocks of road cutting with short patchy

herbs and grasses, bushes locally, 621 m, 29T 065415/468485, 2011.10,17, GAH and

DTH - E214, 2 sh, not kept; ca 1 km W, of San Tirso (Sierra de la Encina), N.-facing

crags and slopes of hard limestone, ca 870 m, 29T 06701/47089, 2007.05.21, GAH ~

2007.19, 1 sh, CGAH; by OU533 near Cemiterio ca 2 km S. of Santa Cruz, granitic

rocks and slopes by road, with sparse herbs and few bushes, 648 m, 29T

065406/468446, 2011.10.17, GAH and DTH - E215, 5 sh, not kept; Prov. Pontevedra: la

Lanzada (S. of O Grove), sand dones with low vegetation, 8 m, 29T 05107/46992,

2011.08.26, GAH and DTH ” E180, 2 sh, not kept; ca 3 km NW. of Sanxenxo, top of

slaty sea-cliffs with low scrub and patches of grasses and herbs, ca 15 m, 29T

051032/469590, 2011.08.26, GAH and DTH - E181, (3) sp, CGAH; just NW. of San Vin-

cente do Grove, open slopes above beach with blown sand over granite, patchy herbs

and grasses, 15 m, 29T 050522/470284, 2011.08.27, GAH and DTH - E182, (2) sp, not

kept; Prov. Zamora: by N631 at E. S^ de Argavanzal (S. of Rionegro del Puente),

slopes above reservoir edge with outcrops of metamorphic rock, patchy grassland,

bushes and saplings, 797 m, 29T 072914/465153, 2011.10.17, GAH and DTH - E216, 2

sh, not kept.

O. barbula, Portugal, Beira Baixa: by IC8 just NE. of Ansiáo, woodland and grass¬

land on limestone, 210 m, 29S 054915/441926, 2011.04.29, GAH and DTH ~ P141, (2)

sp, CGAH; Beira Litoral: Serra de Sico (summit area), rocky limestone slopes with

patchy low scrub and grassland, 548 m, 29S 053909/441909, 2011.01.13, GAH - P98,

(3) sp, 10 sh, CGAH; Pombal Castle, 29SNE3218, 2008.04.24, ÁDO, 6 (3) sp, CGAH; ca

0.5 km S. of IC8 at ca 1.5 km WNW. of Ansiáo, limestone slopes with scrub, crags and

grassland, ca 194 m, 29S 05466/44185, 2010.09.26, GAH and DTH - P73d, (1) sp, 23 sh,

CGAH; 0.5 km S. of IC8 at ca 2 km WNW. of Ansiáo, rocky limestone slopes with

scrub above, ca 190 m, 29S 054642/441843, 2010.09.26, GAH and DTH - P75, 2 sh, not

kept; Vale da Couda (SE. of Almoster), limestone valley with low crags and patchy

Quercus woodland, ca 263 m, 29S 05487/44103, 2010.05.29, GAH and DTH - P45, (1)

sp, CGAH; ca 1.5 km W. of Ansiáo, low Quercus woodland on slope with limestone

rocks, ca 239 m, 29S 05466/44179, 2010.09.26, GAH and DTH - P74, 3 sh, not kept; ca 1

km SW. of Ateanha village (E. of Alvorge), S.-facing limestone hillside with grassland,

patchy scrub and rocks, 310 m, 29S 05498/44262, 2010.05.29, GAH and DTH - P46, (1)

sp, CGAH; S. of IC8 at ca 3 km W. of Avelar, limestone slopes with short grasses and

herbs, scrub locally, 288 m, 29S 05520/44192, 2011.02.24, GAH and DTH - P133, 9 sh,

CGAH; ca 1 km SW. of Alfarelos, limestone crags and grassland, 13 m, 29T

05283/44439, 2010.11.03, GAH and DTH - P79, (4) sp, 8 sh, CGAH; by Paul do Taipal

near Montemor-o-Velho, disused limestone quarry with patches of short vegetation,

45 m, 29T 05271/44481, 2011.01.09, GAH - P96, (3) sp, 6 sh, CGAH; by Ruinas de

Conimbriga, open Olea grove with grassland and herbs beneath, ca 100 m, 29T

054352/443878, 2010.04.27, GAH and DTH - P39, 1 sh, CGAH, oíd shell only; Conim-

briga, Condeixa-a-Nova, 29TNE4339, 2008.10.26, ÁDO, (3) sp, CGAH; by Rio de

Mouros, S. of Condeixa-a-Velha, deciduous woodland with rocky limestone banks

beside stream, ca 117 m, 29T 05436/44386, 2010.04.27, GAH and DTH - P38, 2 sh,

CGAH; Gástelo do Rabanal, rocky limestone slopes, grassland and scrub, ca 315 m,

29T 05489/44309, 2010.04.27, GAH and DTH - P40, 1 sh, CGAH; Penela Castle,

29TNE5231, 2008.10.13, ÁDO, (2) sp, CGAH; Estremadura: Serra da Arrábida, limes¬

tone crags and rocky slopes (S.-facing), 250 m, 29S 04999/42576, 2007.05.16 and 18,

GAH - 2007.9 and 15, 68 sh, CGAH; Serra de Arrábida, rocky limestone slope and

road cutting facing S. and E., with patchy scrub, 263 m, 29S 05000/42577, 2011.02.12,

GAH and DTH - P130, 4 (3) sp, 9 sh, CGAH; Serra de Arrábida, herbs and grasses at

base of S.-facing limestone road cutting, 131 m, 29S 050117/425874, 2011.02.12, GAH

and DTH - P131, (1) sp, 12 sh, CGAH; Serra da Arrábida, N.- to NW.-facing limestone

39

Iberus, 30 (1), 2012

crags and rocky slopes, 380 m, 29S 05021/42601, 2007.05.17, GAH - 2007.10, 12 sh,

CGAH, mixed with O. barbella; Serra de Arrábida, N.~ and S.-facing rocky limestone

slopes with patches of scrub, 388 m, 29S 05020/42602, 2011.02.12, GAH and DTH -

P129, 4 sp, 6 sh, CGAH, mixed with O. barbella; Serra de Montejunto, rocky limestone

slopes and crags with low scrub and varied herbs, ca 400 m, 29S 04950/43382,

2010.04.19, GAH - P34, 2 sh, not kept; ca 2.3 km NE. of Sao Pedro de Moel, crevices

amongst4imestone boulders at base of wall of road bridge by stream, 35 m, 29S

049877/440268, 2010.08.12, GAH and DTH - P67, (3) sp, 12 sh, CGAH, atypical shells

(see text); ca 2 km NNE. of Sao Pedro de Moel, bank 2 m above stream in Eucalyptus

woodland on Coastal sands, 20 m, 29S 049872/440265, 2011.06.04, GAH and DTH -

P161, (4) sp, 4 sh, CGAH, atypical shells (see text); Fórnea (SE. of Alearla), limestone

crags, screes and slopes, patchy scrub, ca 275 m, 29S 05170/43789, 2010.05.02, GAH

and DTH - P43, 10 sh, CGAH; ca 1 km SW. of Serró Ventoso, rocky limestone slopes

and road cutting, scrub and wooded slopes, ca 380 m, 29S 05140/43784, 2010.07.19,

GAH and DTH - P59, 13 sh, CGAH; Serró Ventoso, Porto de Mós, 29SND1479,

2008.02.12, ÁDO, 3 (2) sp, CGAH; ca 2 km NW. of Serró Ventoso, rocky limestone

slope with low crags and scrub, 29S 05129/43809, 2010.07.30, GAH - P62, 9 sh,

CGAH; valley 0.5 km E. of Livramento, limestone valley with patchy scrub and low

rocks, 174 m, 29S 05168/43814, 2010.08.06, GAH and DTH - P65, 21 sh, CGAH; ca 1.5

km SE. of Livramento, rocky limestone slopes facing E. below crag, with evergreen

Quercus woodland, 290 m, 29S 05170/43806, 2010.08.06, GAH and DTH - P64, 19 sh,

CGAH; ca 0.5 km SE. of Livramento, rocky limestone slope with grassland and scrub,

238 m, 29S 05170/43809, 2011.03.22, GAH - P136, 2 sh, not kept; Ribatejo: SE. of

Moitas Venda, limestone crags and rocky slopes with scrub, ca 195 m, 29S

05311/43717, 2010.05.02, GAH and DTH - P41, (2) sp, 27 sh, CGAH; Convento de

Cristo, just W. of Tomar, convent and castle walls and ruins, oíd gardens, 126 m, 29S

05499/43839, 2011.08.05, GAH and DTH - P175, 6 sh, not kept; ca 2 km NNE. of Santa

Cita, river banks with mature trees, dumped masonry, Ulmus grove, roadside verge

and grassy banks, 65 m, 29S 05526/43795, 2011.08.05, GAH and DTH - P176, 9 (4) sp,

CGAH; locality unknown: leg. Holl, 1 sh, SMF 7035, Paralectotype, 1 sh, SMF 7036,

Lectotype.

40

Iberus, 30 (I): 41-52, 2012

© Sociedad Española de Malacologia

A revisión of the Mediterranean Raphitomidae 1 : on the

sibling species Raphitoma contigua Monterosato, 1884 and

Raphitoma spadiana n. sp. (Gastropoda, Conoidea)

Revisión de los Raphitomidae del Mediteráneo 1: las especies her¬

manas Raphitoma contigua Monterosato, 1884 y Raphitoma spadi¬

ana n. sp. (Gastropoda, Conoidea)

Francesco PUSATERI*, Riccardo GIANNUZZI-SAVELLI** and Marco

OLIVERIO***

Recibido el 28-V-201 1. Aceptado el 26-X-20 11

ABSTRACT

A new raphitomid toxoglossan, Raphitoma spadiana n. sp., is described from the Mediter¬

ranean Sea. !t is the sister species to R. contigua Monterosato, 1 884, from which it differs

in the different protoconch (paucispiral vs. multispiral), adding to a long list of pairs of

caenogastropod species differing in their larval development.

RESUMEN

Una nueva especie de toxogloso rafitómido, Raphitoma spadiana n. sp., se describe del

Mediterráneo. Es especie hermana de R. contigua Monterosato, 1 884, de la cual se diferen¬

cia por su protoconcha (pauciespiral vs. multiespiral), y se añade a una larga lista de parejas

de especies que difieren en su desarrollo larvario (planctotrofico vs. no-planctotrofico).

INTRODUCTION

The superfamily Conoidea, with

over 300 genera and 4.000 recognised

species, but probably over 12,000 extant

species (Bouchet, 1990; Taylor,

Kantor & Sysoev, 1993; Tucker, 2004),

represents the largest radiation of the

entire phylum Mollusca. In a work on

the phylogeny of the group based on a

cladistic analysis of foregut morphology,

Taylor et al. (1993) have highlighted

the rampant homoplasy in the charac-

ters of Shell and radula in conoideans.

Accordingly, they have rearranged most

of the conoideans into two families:

Conidae, comprising Coninae and 4

subfamilies traditionally considered as

Turridsb and Turridae s.s. including

some of the traditional Turrids'. More

recently, Puillandre, Samadi, Boisse-

LiER, Sysoev, Kantor, Cruaud,

CouLOUx & Bouchet (2008) and

Bouchet, Kantor, Sysoev &

Puillandre (2011), based on DNA phy-

logeny, have provided a major update of

conoidean classification. Although a

larger taxonomic coverage would be

desirable to further stabilize the molecu¬

lar phylogeny, however, the position of

*Via Castellana, 64 - 90135 Palermo. E-mail: francesco@pusateri.it

** Via Mater Dolorosa, 54 - 90146 Palermo. E-mail: malakos@tin.it

*** Dept. Biology and Biotechnologies “Charles Darwin”. Zoology, Viale deH’Universitá 32, 1-00185 Roma. E-

mail: marco.oliverio@uniromal.it

41

Iberus, 30 (1), 2012

the Raphitomidae as a clade of the

Conoidea is sufficiently supported.

Raphitomidae are based on the

genus Raphitoma, which was introduced

by Bellardi, (1847: 85), as including 34

fossil and extant species. During the

revisión of the Mediterranean Raphito¬

midae we are currently carrying out, we

have found several pairs of species, dif-

fering only or mostly in the size and

shape of the protoconch. After the

pioneer works of Thorson (1946, 1950),

addressing the relationships between

protoconch morphology and type of

larval development in the caenogas-

tropods, the dichotomy multispiral pro-

toconch/planktrotrophic development

vs. paucispiral protoconch/ lecitho-

trophic development has been largely

accepted (Jablonski & Lutz, 1980; Rex

& Warén, 1982). Although some

authors have used this dichotomy to

allocate species into planktotrophic vs.

non-planktotrophic genera (e.g.:

PowELL, 1966; Aartsen 1988), Bouchet

(1990) has clearly demonstrated that

such abuse of protoconch morphology

would produce polyphyletic taxa, and

artificially sepárate closely related

species among distinct genera. In raphit-

omids, the separation of Raphitoma

(multispiral protoconch) and Philbertia

(paucispiral protoconch) is inconsistent

and must be rejected.

On the other hand, differences in

protoconch morphology, which reflect

differences in larval development

(planktotrophic vs. lecithotrophic) are a

good base to recognise distinct species

in caenogastropods. In fact, poecilogony

(the ability of a species to use different

developmental strategies, e.g. plank-

totrophy and lecithotrophy) is not

observed within the prosobranchs

(Bouchet, 1989), whilst it has been

sometimos recorded among opistho-

branchs [e.g.: Haminoea callidegenita

SYSTEMATICS

Gibson & Chia, 1989, Elysia chlorotica

Gould, 1870, Alderia luillowi Krug,

Ellingson, Burton & Valdés, 2007 (West,

Harrigan & PiERCE 1984, Gibson &

Chía, 1995, Krug, Ellingson, Burton

& Valdés, 2007)].

The presence of pairs of caenogastro-

pod species with multispiral vs. pau¬

cispiral protoconchs has been particu-

larly reviewed in the Mediterranean Sea

(see Oliverio, 1996a, b, with references

therein), but it is a phenomenon

observed on a global scale (Oliverio,

1997). It has been related to a speciation

model, where the very shift in larval

development with loss of planktotrophy

has been the driving mechanism of spe¬

ciation (Oliverio 1996a).

In the genus Raphitoma, we have

scored several such pairs of species with

different protoconchs (multispiral vs.

paucispiral). A thorough revisión of all

Mediterranean pairs (which we have

currently in progress) will be the subject

of a forthcoming papen In the present

Work we present the case of the rather

obscure taxon Raphitoma contigua (Mon-

terosato, 1884), of which we have exam-

ined the type material and selected a

lectotype with multispiral protoconch to

stabilize the usage of the ñame, and its

unnamed sister species with paucispiral

protoconch.

Abbreviations:

HUJ = Hebrew University of Jerusalem

(Israel)

MCZR = Museo Civico di Zoologia di

Roma (Italy)

MNHN = Muséum National d'Histoire

Naturelle (Paris, Erance)

NMW = National Museum of Wales

(Cardiff, United Kingdom)

SMNH = Swedish Museum of Natural

History (Stockholm, Sweden)

sh = shell(s)

(Citation of unpublished ñames is not intended for taxonomic purposes)

Family Raphitomidae Bellardi, 1875

42

PUSATERI ET AL.: Mediterranean Raphitomidae 1: Raphitoma contigua and R. spadiana

Genus Raphitoma Bellardi, 1847

Type species: Pleurotoma hystrix Cristofori and Jan, 1832 {nomen niidiim, validated by Bellardi,

1847 as "Pleurotoma histrix Jan") by subsequent designation (Monterosato, 1872: 16).

Synonyms:

Homotoma Bellardi, 1875 non Guérin-Ménéville, 1844 - type species: Murex textile Brocchi, 1814

by subsequent designation (Fischer, 1883).

Cirillia Monterosato, 1884, non Rondani, 1856 - type species: Murex linearis Montagu, 1803, by

subsequent designation (Crosse, 1885).

Cordieria Monterosato, 1884 non Roualt, 1848 - type species: Murex reticulatus Brochi, 1814, by

subsequent designation (Crosse, 1885).

Leufroyia Monterosato, 1884 - Type species: Pleurotoma leufroyi Michaud, 1827, by subsequent

designation (Crosse, 1885)

Philbertia Monterosato, 1884 - type species: Pleurotoma philberti Michaud, 1829 by subsequent

designation (Crosse, 1885)

Peratoma Harris & Burrows, 1891 - nomen novum pro Homotoma Bellardi, 1875, non Guérin

Ménéville, 1844

Cenodagreutes E.H. Smith, 1967 - type species: Cenodagreutes aethus E. H. Smith, 1967, by original

designation.

Cyrtoides Nordsieck, 1968 - type species: Pleurotoma ruáis Scacchi, 1836, by original designation.

Lineotoma Nordsieck, 1977 - nomen novum pro Cirillia Monterosato, 1884, non Rondani, 1856

Diagnosis: Shell of small to médium^

small size (5-25 mm), turreted, fusiform

or ovoid-fusiform, with slender spire

and uniformly convex whorls. Relien-

late-cancellate sculpture on the teleo-

conch. Protoconch of 3-4 whorls when

multispiral, with a protoconch I (embry-

onic shell) of 0.5-0. 7 whorl and a proto¬

conch II (larval shell) of 2-3 whorls, with

a diagonally cancellate sculpture. Pau-

cispiral protoconch of 2 whorls, with

large nucleus and reticulate sculpture.

Remarks: For the complex nomen¬

clatura! issues, including the type

species designations, see Dall (1918),

VAN Aartsen, Menkhorst & Gitten-

BERGER (1984: 89-90) and Rolán,

Otero-Schmitt & Fernandes (1998:

105).

In the Mediterranean Sea, species of

Raphitoma are usually found alive on

soft bottoms, ranging from Coastal bio-

clastic sands to muddy bioclastic sands

(DC and DE, respectively: Pérés &

Picaro, 1964) although it is not infre-

quent to find specimens hiding under

stones and in crevices, especially at

daytime.

Raphitoma contigua (Monterosato, 1884) (Figs. 1-3, 14)

Philbertia contigua Monterosato, 1884:133

Type material: Lectotype (here designated, 9.05 x 4.16 mm) and one paralectotype (9.30 x 4.10 mm)

with handwritten label by Monterosato: "Ph. contigua typ. Monts! Palermo" (MCZR, 16702/1); 3

paralectotypes with handwritten labels by Monterosato: "Ph. contigua Monts. / Napoli Staz. ZooL"

(MCZR, 16702/2); 2 paralectotypes labelled "contigua Napoli" (MCZR, 16702/3); 3 paralectotypes

with handwritten labels by Monterosato: "R. contigua - Isole Baleari" (MCZR, 16702/4); 16 para¬

lectotypes with handwritten label by Monterosato "Ph. contigua M. / Pal!", coll. Coen in HUJ, 8072

(including 7 specimens of R. bicolor); 2 paralectotypes with handwritten Dautzenberg label: "ClathureUa

purpurea/ Montagu /var. Philberti Michaud/St Lunaire/Types Molí. Rouss./ 1. 1 pl. 14 fig. 13,14"

(two specimens of R. oblonga) (MNHN, Paris); 1 paralectotype with handwritten Dautzenberg label:

"ClathureUa purpurea/Montagu/var. Philberti Michaud /Roussillon/ Type Molí. Rouss./ t. I pl.

14 fig. 15" (a specimen of R. atropurpúrea) (MNHN, Paris). Type locality: Palermo.

43

Iberus, 30 (1), 2012

Figures 1-4. Shells of Raphitoma spp. 1-3. Raphitoma contigua (Monterosato, 1884). la-c: lecto-

type (MCZR, 16702/1, h: 9.05 mm) with original label; 2: Ficarazzi, Palermo (coll. Pusateri, h: 10

mm); 3: Roquebrune, Les Issambres (coll. Hoarau, h: 9.7 mm), subadult shell with 8 spiral

cordlets above the aperture. 4. Raphitoma lineolata B.D.D., 1884, St, Raphael, France (coll.

Hoarau, h: 7.2 mm).

Figuras 1-4. Conchas de Raphitoma spp. 1-3. Raphitoma contigua (Monterosato, 1884). la-c; lec-

totipo (MCZR, 16702 ! 1, h: 9,05 mm) con la etiqueta original; 2: Ficarazzi, Palermo (col Pusateri,

h: 10 mm); 3: Roquebrune, Les Issambres (col. Hoarau, h: 9,7 mm), concha subadulto con 8 cordonci¬

llos espirales encima de la abertura. 4. Raphitoma lineolata B.D.D., 1884, Saint Raphael, Francia

( col. Hoarau, h: 7,2 mm).

44

PUSATERI ET AL.'. Mediterranean Raphitomidae 1: Raphitoma contigua and R. spadiana

Figures 5-7. Shells of Raphitoma spp. figured in Bucquoy, Dautzenberg & Dollfus (1883), labelled

as “Clathurella purpurea philberti Michaud”. 5, 6: Raphitoma oblonga (Jeffreys, 1867), St.

Lunaire (BDD, 1883: pl. 14, figs 13, 14) with original label (MNHN, Paris); 7: Raphitoma atrop¬

urpúrea (Locard & Caziot, 1899), Roussillon (BDD, 1883: pl. 14 fig. 15) with original label

(MNHN, Paris).

Figuras 5-7. Conchas de Raphitoma spp. figuradas en Bucquoy, Dautzenberg y Dollfius (1883), etique¬

tadas como “Clathurella purpurea var. philberti Michaud”. 5, 6: Raphitoma oblonga (Jejfireys,

1867), Saint Lunaire (BDD, 1883: Idm. 14, fiig. 13, 14) con la etiqueta original (MNHN, Paris); 7:

Raphitoma atropurpúrea (Locard y Caziot, 1899), Rosellón (BDD, 1883: lám. 14, fiig. 15) con la eti¬

queta original (MNHN, París).

Material examined: The type material and: France - Marseille (MCZR, coll. Monterosato, inv. 16811,

4 sh.: labelled by Monterosato “H. conformis” , an unpublished manuscript ñame). Roquebrune, Les

Issambres (Var), (coll. Hoarau, 1 sh.). St. Raphael, Cote d'Azur (MCZR, coll. Monterosato, 16696,

1 sh.: labelled "P. tomentosa")- Corsíca - Bastía (coll. Pusateri, 1 sh.; coll. Margelli, 1 sh.). Bastía,

Alístro (coll. Margelli, 2 sh.). Sardínía - Ennío Falco Cave, Porto Conte, Sassari (coll. Oliverio, 3 sh.).

La Maddalena Is. (coll. Rufini, 1 sh.; coll. Spanu, 1 sh.). Oristano, (coll. Rufini, 2 sh.). S'Archittu,

Oristano (coll. Sossu, 2 sh.). Stintino, Sassari (coll. Rufini, 1 sh.). Torre del Bollo, Alghero (coll. Spanu,

1 es.). Liguria - Genova (coll. Pusateri 2 sh.). Tuscany - Calambrone (Pisa), coll. Bartolini, 1 sh. Cas-

tiglioncello, Livorno (coll. Coppini, 1 sh.), (coll. Margelli, 2 sh.). Punta Ala, Grosseto (coll. Coppini,

1 sh.). Giannutri Is. (coll. Smriglio, 1 sh). Elba Is., Marina di Campo (coll. Bartolini, 7 sh.). Elba Is.

(coll. Bartolini, 4 sh.). Tuscan Archipelago (coll. Bartolini, 4 sh.). Latium - Tor Paterno Marine Pro-

tected Area, Roma (coll. Oliverio, 1 sh.), (coll. Rufini ex Ruggeri, 1 sh.). Campania - Trombetta Cave,

-35 m. Palinuro, Salerno (coll. Oliverio, 1 sh. ). Punta Campanella, Napoli (coll. Bartolini, 2 sh.).

Sicily - Acitrezza, Catania (SMNH, lot 73198F, 10 sh.). Cannizzaro, Catania (coll. Micali, 3 sh.).

Palermo (coll. Pusateri, 18 sh., 1 albinistic). Palermo (HUJ, In. 8080 with Coen's label Philbertia pur¬

purea lineolata BDD, 1 sh). Aspra, Palermo (coll. Girgenti, 3 sh.). Isola delle Femmine, Palermo (coll.

Pusateri, 3 sh. legit Bagnera). Palermo (MCZR, coll. Monterosato, inv. N, 16702, 2 sh.). Gulf of Carini,

Palermo (coll. Palmeri, 3 sh.). Capo Passero (coll. Margelli, 1 sh.). Taormina, Messina (coll. Villari,

1 sh.). Pantelleria Is. (coll. Bartolini, 1 sh.). Apulia - Taranto (MCZR, coll. Monterosato, sine numero

45

Iberus, 30 (1), 2012

3 sh.). Croazía - Korcula Is. (coll. Micali, 1 sh.). Krk Is. (coll. Bartolini, 10+ sh.). Velirat, Dugi Otok

(Isola Lunga) (coll. Pusateri, 1 sh.). Algeria - Algiers (MCZR, coll. Monterosato, 16815, 1 sh). "C.

d'Afr." [Coasts of Africa], (MCZR, coll. Monterosato, unnumbered lot, sine nomine, 4 sh.).

Description: Shell sub-fusiform (mean

h/d 2.2, std 0.13), height 9-15 mm (mean

10.9, std 1.70), width 4.5-6 mm (mean 4.9,

std 0.49). Protoconch multispiral of 2.7

convex whorls, height 325 pm, width 300

pm; protoconch I of 0.6 whorls, width 170

pm, with irregularly placed small tuber-

cles; protoconch II of 2.1 whorls, with a

diagonally cancellate sculpture, and two

suprasutural small spiral threads.

Teleoconch of 7 convex whorls. Axial

sculpture of 16-18 orthocline or slightly

opisthocline ribs (sometimes more in

larger specimens), and interspaces

wider (xl.5) than the ribs. Spiral sculp¬

ture on the last whorl of 17-20 cordlets,

of which 6-7 (rarely 8) above the aper-

ture, slightly narrower than the axial

ribs. Cancellation rectangular, with

small and elongated tubercles at the

intersection of axials and spirals. Tuber¬

cles on the two adapical cordlets narrow

and spinulose (especially in fresh or

subadult specimens). Sculpture visible

in transparency throughout the internal

Shell Wall.

Subsutural ramp narrow, with small

tubercles in correspondence with the

axial ribs tip.

Columella simple, slightly sinuous

anteriorly, gently angled posteriorly.

Outer lip with 8-9 strong inner denticles

(rarely 10-11), the most anterior delimit-

ing the siphonal canal, the most poste¬

rior delimiting the anal sinus.

Colour uniformly tawny, rarely

darker, sometimes with lighter spots.

Suprasutural cordlet occasionally white.

Occasional comma-shaped white spots

on the subsutural ramp.

Distribution: Only known from

examined material, probably the entire

Mediterranean Sea.

Remarks: Monterosato (1884: 133)

introduced Philbertia contigua without

any description, but with reference to

BDD (1883) as follows:

"P. contigua, Monts. (nov. forma)

= Clathurella purpurea (non Mtg.) var.

Philberti (non Mich.), B.,D. e D. - l.c.

[Molí. Mar. Roussillon] p. 40, t. 14, f. 13-

15 (C. di Prov.).

=? Anna Massena Risso - 1826, p. 214,

f. 68 (foss. Alpi Marit.),

Isole Baleari (Monjo); Alger (Joly);

Bona (Hagenmüller); Palermo e Messina

(Monts.); Napoli, Lipari (Tiberi) ecc."

Evidently, Monterosato did not

check de visu the original material used

for BDD's plates (now stored at MNHN,

Paris: Figs. 5-7), relying only on BDD's

figures. In fact, Monterosato cites

"C.[oste] di Prov.[enza]" (coasts of

Provence), evidently presuming that the

figured specimens were all from

Provence. Actually, the specimens in

BDD's figs 13 and 14 (MNHN, Paris) are

labelled as coming from St. Lunaire

(Bretagne), and only the shell of fig. 15

is from Roussillon. However, the lots in

the Monterosato collection labelled as

Philbertia contigua are not conspecific

with the specimens figured in BDD's

figs 13-14 (Figs 5, 6), which are referable

to Raphitoma oblonga (Jeffreys, 1867) a

valid species from the Charmel area, ñor

with the specimen figured in BDD's fig.

15 (Fig. 7), which is referable to Raphit¬

oma atropurpúrea (Locard & Caziot,

1899), Monterosato's concept of contigua

remained constan!, as we have verified

in several collections hosting specimens

ex Monterosato (Melvill-Tomlin,

National Museum of Wales, lot 12908;

Coen, Hebrew University, lot 8072). Fvi-

dently, also Dautzenberg & Durou-

CHOUX, 1900: 14 and Locard & Caziot,

1899:58, while citing R. contigua realized

that the material sent to them by Mon¬

terosato did not fit BDD's figures 13-14,

and kept contigua as a distinct entity.

Nevertheless this does not invalidate

that the specific ñame contigua is made

nomenclaturally available "by indica-

tion" (ICZN art. 12.2), with a type mate¬

rial that ineludes all the localities cited

in Monterosato (1884).

R. contigua (h/d 2.2) is less slender

than the most similar species with mul¬

tispiral protoconch: R. lineolata (BDD,

46

PUSATERI ET AL,: Mediterranean Raphitomidae 1: Raphitoma contigua and R. spadiana

Figures 8-13. Shells of Raphitoma spadiana spec. nov. 8a-c: holotype, Lipari Is. (MNHN, h: 9.6

mm); 9, 12: paratype A, Lipari Is, (MNHN, Paris, ex coll. Pusateri h: 8 mm); 10: North African

coasts, “C.[oste] d’Afr.pca]” (MCZR, coll. Monterosato, unnumbered lot, h: 6.8 mm); 11, 13: La

Ciotat, France (MCZR, 16773, coll. Monterosato, sine nomine, h: .8.6 mm).

Figuras 8-13. Conchas de Raphitoma spadiana spec. nov. 8a-c: holotipo, islas Lipari (MNHN, h: 9,6

mm); 9, 12: paratipo, islas Lipari (MNHN, París, ex coll Pusateri h: 8 mm); 10: costas de África del

Norte “C.[oste] dÁfr.[ica].” (MCZR, coll Monterosato, lote sin número, h: 6,8 mm); 11, 13: La

Ciotat, Francia (MCZR, 16773, col Monterosato, sine nomine, H: 8,6 mm).

47

Iberus, 30 (1), 2012

Figure 14. Raphitoma contigua (Monterosato, 1884). I4a,b: protoconch; I4c: subsutural ramp.

Figure 15a-c. Protoconch of Raphitoma spadiana n. sp.

Figura 14. Raphitoma contigua (Monterosato, 1884). I4a,b: protoconcha; l4c: rampa subsutural.

Figura 1 5a-c. Protoconcha de Raphitoma spadiana n. sp.

1882) (h/d 2.8), R. atropurpúrea (Locard

& Caziot, 1899) (h/d 2.8), R. densa

(Monterosato, 1884) (h/d 2.8), R.

oblonga (Jeffreys, 1867) (h/d 2.5). Raphit¬

oma lineolata (Fig. 4) is very similar in

general aspect, but has a less robust

Shell, lacks the narrow subsutural

ramp, has a narrower aperture, and

shows always few and scattered white

tubercles on the last whorl as well as a

subsutural white cord, both lacking in

contigua. R. atropurpúrea has a different

colour (dark brown-purplish vs.

tawny). R. densa (Fig. 17) is more

densely sculptured, and has a colour

pattern of ash-grey spots over a dark

chestnut background. R. oblonga is a

less known, exclusively Atlantic

species, differing from R. contigua by its

narrower aperture and the constant

presence of a white spiral line as long

as the space of four axial ribs, starting

on the outer lip.

R. alternans (Monterosato, 1884) (see

Fig. 16) has a paucispiral protoconch.

and differs also from R. contigua in

being more slender (h/d 2.6 vs. 2.2), and

in its colour pattern of white spots over

a dark chestnut background.

Shells with identical teleoconchs of

the contigua type, but with two distinct

protoconch types (multispiral vs. pau-

cispiral) are known, and we consider

them as distinct species. Therefore, we

have examined the protoconchs in the

type material of Philbertia contigua Mon-

terosato, which turned out to consist as

follows: MCZR 16702/1, one shell has a

multispiral protoconch, the other lacks

protoconch; MCZR 16702/2, one lacks

protoconch, two with parts of multispi¬

ral protoconch; MCZR 16702/3, both

with parts of multispiral protoconch;

MCZR 16702/4, with parts of multispi¬

ral protoconch. To stabilize the use of

the ñame R. contigua we have selected

the shell in the lot MCZR, 16702/1 with

intact (multispiral) protoconch, as the

lectotype of Philbertia contigua Mon¬

terosato.

48

PUSATERI ET AL.'. Mediterranean Raphitomidae 1: Raphitoma contigua and R. spadiana

Figures 16, 17. Shells of Raphitoma spp. 16: Raphitoma alternans (Monterosato, 1884), syntype

(MCZR, 16676, h; 12.00 mm), Mondello (Palermo); 17: Raphitoma densa (Monterosato, 1884),

syntype (MCZR, 16807, h: 10.15 mm), Palermo.

Figuras 16, 17. Conchas de Raphitoma spp. 16: Raphitoma alternans (Monterosato, 1884), sintipo

(MCZR, 16676, h: 12,00 mm), Mondello (Palermo); 17: Raphitoma densa (Monterosato, 1884),

sintipo (MCZR, 16807, h: 10,15 mm), Palermo.

Raphitoma spadiana Pusateri & Giannuzzi-Savelli spec. nov. (Figs. 8-13, 15)

Type material: Holotype (MNHN, París) height 9.6 mm, width 4.1 mm, from Lipari Is. (Sicily) (legií

Claudio Ebreo). Paratype A (coll. Pusateri, height 8.0 mm, width 3.7), from Lipari Is.; Paratype B

(MNHN, París) height 8.1 mm, width 3.6 mm, from Scilla, Calabria, -50 m (legit Angelo Vazzana)

Other material examined: France - Marseille (coll. Locard, MNHN París, 1 shell). La Ciotat, Cote

d'Azur (coll. Monterosato, MCZR, 16773, sine nomine 8.6 x 3.8 mm). Sardinia - Alghero (coll. Spanu,

1 sh. juv.). Calabria - Scilla, Reggio Calabria (coll. Vazzana, 20 shells). Sicily - Mondello (Palermo)

(coll. Pusateri, 2 shells, juv.). Marettimo Is. (coll. Agamennone 1 shell, juv.). Marettimo Isl. (coll.

Agamennone, 1 sh.). Apulia - Taranto (coll. Giannuzzi Savelli, 1 shell). Egypt - Alexandria (coll.

Pusateri, legit Claudio Ebreo, 2 shells). Tunisia- Djerba Is., (coll. Agamennone, 1 shell). "C. á'Afr.”

[Coasts of Africa], (MCZR, coll. Monterosato, unnumbered lot, sine nomine 1 sh. subadult, height

6.8 mm, width 3.2 mm; stored with 4 shells of R. contigua). Greece - Crete Is. (coll. Alfinito, 1 shell).

Cyprus - (coll. Bartolini, 1 sh.)

Etymology: after our dear friend Gianni Spada, a specialist of turrid taxonomy.

Type locality; Lipari Is.

Description: Shell subfusiform 9.5

mm high and 4.1 mm wide.

Protoconch paucispiral, only proto=

conch I of 1.25 convex whorls, height

425 pm, width 450 pm; sculpture irregu-

larly cancellate.

Teleoconch of 6 convex whorls. Axial

sculpture of 18 orthocline ribs, ínter-

spaces wider (xl.5) than the ribs.

Spiral sculpture on the last whorl of

23 cordlets, of which 8 above the aper-

ture, slightly narrower than the axial

ribs, interspaces wider (x2) than the

ribs. Cancellation rectangular, with

small and elongated tubercles at the

intersection of axials and spirals. Tuber¬

cles on the two subsutural cordlets

narrow and spinulose. Sculpture visible

in transparency throughout the very

thin internal shell wall.

Subsutural ramp narrow, with small

tubercles in correspondence with the tip

of the axial ribs. Two cordlets on the

subsutural ramp, the adapical smaller

with smaller tubercles, the second being

the largest of the spirals. The third

cordlet, as small as the first one.

Columella simple, slightly sinuous

anteriorly, gently angled posteriorly.

Outer lip with 12 strong inner denticles,

the most anterior delimiting the

siphonal canal, the most posterior

delimiting the anal sinus. Denticles 1-2

and 11-12 closer than the others.

49

Iberus, 30 (1), 2012

Siphonal canal short, open.

Colour uniform light tawny, with

rare lighter tubercles. Some axial ribs

entirely white on the first three teleo-

conch whorls; on the last whorl tenth

spiral cordlet and last two axial ribs

partly white.

Distrihution: Only known from

examined material, probably entire

Mediterranean Sea.

Remarks: Variation of the examined

material: height 6.8=11.7 mm (mean 9.4

std 1.76), width 3. 2-4. 6 mm (mean 4.1

std 0.56); mean h/ d: 2.3 std 0.15,

Axial ribs 16-18, sometimes slightly

opisthocline in the last whorl. Occa-

sional comma-shaped white spots on

the subsutural ramp. Colour ranging

from light to very light tawny. White

axial ribs from presen! in all whorls to

very rare or absent. The spiral cordlet 10

may be white spotted.

Raphitoma spadiana differs from the

closely related R. contigua in its pau-

cispiral protoconch and thus an inferred

lecithotrophic development (vs. multi-

spiral protoconch and an inferred plank-

trotrophic development in contigua).

Additionally, R. spadiana is of slightly

smaller size, and lighter in colour. R.

alternans (Monterosato, 1884) has also a

paucispiral protoconch, but differs also

from R. spadiana by being more slender

(h/d 2.6 vs. 2.3), and by its colour

pattern of irregular white spots over a

dark chestnut background.

R. spadiana (h/d 2.3) is also less

slender than the most similar species

ACKNOWLEDGEMENTS

Franco Agamennone, Silvia Alfinito,

Giuseppe Bagnera, Manrico Coppini,

Antonio Girgenti, André Hoarau, Fiera

lacopelli, Alessandro Margelli, Paolo

Mariottini, Pasquale Micali, Italo

Nofroni, Alberto Palmeri, Stefano Rufini,

Maria Scaperrotta, Cario Smriglio, Maur-

izio Sosso, Maria Teresa Spanu, Angelo

Vazzana, Alberto Villar!, are thanked for

having placed their materials at our dis-

posai, Gianni Repetto for his valuable

bibliographic help.

with multispiral protoconch: R. lineolata

(BDD, 1882) (h/d 2.8), R. atropurpúrea

(Locard & Caziot, 1899) (h/d 2.8), R.

densa (Monterosato, 1884) (h/d 2.8), R.

oblonga (Jeffreys, 1867) (h/d 2.5).

Raphitoma lineolata (Fig. 4) is very

similar in general aspect, but differs in

having a less robus! shell and a nar-

rower aperture, lacking the narrow sub¬

sutural ramp, and showing always few

and scattered white tubercles on the last

whorl as well as a subsutural white

cord, both lacking in spadiana. Raphitoma

atropurpúrea has a different colour (dark

brown-purplish vs. tawny). Raphitoma

densa is more densely sculptured, and

has a colour pattern of irregular ash-

grey spots over a dark chestnut back¬

ground. Raphitoma oblonga is exclusively

Atlantic, and differs by its narrower

aperture and the constan! presence of a

white spiral line as long as the space of

four axial ribs, starting on the outer lip.

According to the admittedly limited

material examined, R. spadiana seems to

show some interpopulational variation,

even over a short geographic scale

(which would be congruent with a non-

planktotrophic larval development),

with at least one extreme case. A lot of

20 adult shells from Scilla (Calabria)

range between 5. 2-7.8 mm (mean 6.67)

in height, and 2.5“3.9 mm (mean 3.16) in

width, with h/d 2.11, and are darker

that all other examined shells. Given the

small number of shells available overall

for comparison, we keep this lot within

the putative range of R. spadiana.

Claudio Manicastri allowed the

study of the materials in the Mon¬

terosato collection (MCZR, Rome) and

Massimo Appolloni and Angela Mastro-

giacomo assisted during the visit at

MCZR. Philippe Bouchet allowed the

study of the materials at MNHN (Paris),

and Virginie Fieros and Fierre Lozouet

assisted during the visit at MNHN. H.

Mienis loaned materials from the Coen

collection (HUJ, Jerusalem). Anders

Warén loaned materials from SMNH

50

PUSATERI ETAL.: Mediterranean Raphitomidae 1: Raphitoma contigua and R. spadiana

(Stokholm). Stefano Bartolini did the

light photographs, SEM photographs

were done at the "LIME" (Interdepart-

mental Laboratory of Electron Micros-

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52

Iberus, 30 (1); 53-66, 2012

© Sociedad Española de Malacologia

New species of neogastropods from the islands of the Gulf

of Guinea, West Africa

Nuevas especies de neogasterópodos de las islas del golfo de Guinea,

África occidental

Emilio ROLAN* and Sandro GORI**

Recibido el 26-V-201 1. Aceptado el 28-X-201 1

ABSTRACT

Materia! collected from a recent expedition by the second author (SG) to Principe Island in

the Guiñean Gulf is studied. Material from nearby islands that was waiting for publication

is aiso included. The new species proposed here belong to the genera Anachis (1 species)

and Mifrella (1 species) in the family Columbellidae, and Mitromorpha (3 species) in the

famiíy Mitromorphidae, superfamily Conoidea.

RESUMEN

Se estudia el materia! recogido en una reciente expedición del segundo autor (SG) a la

isla de Príncipe en el Golfo de Guinea. Al tiempo se incorpora material de otras islas pró¬

ximas que estaba pendiente de publicación. Los géneros estudiados en los que se inclu¬

yen las nuevas especies encontradas han sido Anachis (una especie) y Mifrella (una espe¬

cie) de la familia Columbellidae, y Mitromorpha (tres especies) de la familia

Mitromorphidae, superfamilia Conoidea.

INTRODUCTION

In recent years the second author

(SG) made several trips to the islands of

the Guiñean Gulf, during which a num-

ber of samples were collected. When

these were examined, a few new species

were found. In the present paper we de¬

scribe the ones belonging to the families

Columbellidae Swainson, 1840 (genera

Anachis and Mifrella) and Mitromorphi¬

dae Casey, 1904 (genus Mitromorpha) af-

ter the recent systematic changes in

BOUCHET, KANTOR, SYSOEV & PuiLLAN-

DRE (2011). On the other hand, material

collected on a previous trip by the first

author (ER), was mentioned in a previ¬

ous paper (Rolán & Boyer, 2001), but

has now been examined again and com¬

pared with the new material collected,

enabling us to reach new conclusions re-

garding its status, which leads us to de¬

scribe here a further two new species.

Abbreviations used:

MHNS Museo de Historia Natural de la

Universidad, Santiago de Compostela

MNCN Museo Nacional de Ciencias

Naturales, Madrid, Spain

MNHN Muséum National d'Histoire

Naturelle, Paris, France

* Museo de Historia Natural, Universidad de Santiago, Campus Universitario Sur, 15782 Santiago de

Compostela, Spain

** Via Sernesi, 7, 57123 Livorno, Italia

53

Iberus, 30 (1), 2012

MCZ Museum of Comparative Zoology,

Cambridge, USA

NHMUK Natural History Museum,

London, England

CFS collection of Francisco Sicilia

Guillen, Arrecife, Spain

CMS collection of Manuel Suárez,

Ferrol, Spain

CPR collection of Peter Ryall, Maria

Rain, Austria

CSG collection of Sandro Gori, Livorno,

Italia

SYSTEMATIC PART

Family Columbellidae Swaison, 1840

Genus Anachis H. & A. Adams, 1853

H. & A. Adams (1858) defined the

genus Anachis in the following terms:

"Shell ovately-fusiform; spire elevate,

whorls longitudinally ribbed; aperture

narrow; columella straight; outer lip

somewhat rectilinear, crenate inter-

nally". Thirteen species are known in

the Fastern Atlantic región (including

the Macaronesian Archipelagos, the

Mediterranean and the West African

coast).

Some species of this genus in West

Africa were described long ago, the first

being Anachis cuspidata Marrat, 1877.

Not many more species were found

over the years and by the end of the

XXth century, only five more had been

described, the latest being Anachis

avaroides Nordsieck, 1975. All of these

species were described singly and they

have mostly a limited geographical

range. As the result of the frequent col-

lecting trips made by many malacolo-

gists to several countries in the area, it is

curious to point out that in the revisions

and other papers published over the last

10 years (Rolán & Ruque, 2002;

Hernández & Boyer, 2005; Pelorce &

Boyer, 2005; Rolán, 2005; Rolán &

Boyer, 2006; Pelorce & Rolán, 2007;

Rolán & de Oliveira, 2008; Horro &

Rolán, 2010; Segers, Swinnen & de

Prins, 2009), seven new species were

described or studied: Only in one case

(Pelorce & Boyer, 2005), however, are

we to find two species together in this

genus, the others having been described

always individually.

Since most species have a paucisph

ral protoconch, presumably their geo¬

graphical range is small. Only Anachis

avaroides seems to have a larger distribu-

tion area in the Macaronesian archipela¬

gos.

Anachis xani spec. nov. (Figs. 1-6)

Type material: Holotype (Figs. 1-3) in MNHN (24819); Paratypes in the following collections:

MNCN (15.05/60002, 1 s, Fig. 4); MHNS (100561, 2 s); NHMUK (1); CPR (1); CSG (9).

Other material examined: 10 shells eroded or without protoconch, and 17 fragments from the

type locality.

Type locality: Tinhosa Pequeña, 01°22.52'N, 07°16.53'E, under rocks, 25 m. Principe Island.

Etymology: The species is named after Juan Manuel "Xan" Guerra, who is in charge of the shell

collections of the Museo de Historia Natural "Luis Iglesias", University of Santiago de Com-

postela.

Description: Shell (Figs. 1-4) solid,

broadly fusiform with a modérate high

spire. Protoconch (Fig. 5) with 1 i/i

whorls, sharply pointed and about 0.5

mm in máximum diameter, apparently

smooth but with spiral micro striae (Fig.

6) and usually coloured with brown and

whitish bands. Teleoconch of 4 Vi

slightly convex whorls, with a distinct

suture and a very small subsutural

54

RoláN and Gori: New species of neogastropods from the islands of the Gulf of Guinea

Figures 1-5. Anachis xani spec. nov. 1-3: holotype, 5.0 mm (MNHN); 4: paratype, 5.5 mm

(MNCN); 5: apex and protoconch of the holotype; 6: detall of the micro sculpture.

Figuras 1-5. Anachis xani spec. nov. 1-3: holotipo, 5,0 mm (MNHN); 4: paratipo, 5,5 mm (MNCN);

5: ápice y protoconcha del holotipo; 6: detalle de la microescultura.

55

Iberus, 30 (1), 2012

shelf. Sculpture formed by axial ribs,

badly defined subsuturally, a little irreg¬

ular, with variable interspaces, about 20

in the two first whorls; in the last whorl,

about 15-19 can be observed, but on the

last quarter of the whorls, they are

almost entirely faded. There are no clear

spiral threads, but under high magnifi-

cation numerous micro striae can be

seen: about 10 on first whorl, 13-15 on

the second and about 24-28 on the third.

Towards the base about 12-13 well

marked striae form evident cords. Aper-

ture narrow and axially elongate,

usually with 6 teeth on the inner part of

the external lip, the upper ones being

larger. Columella curved, S-shaped,

with 2 denticles usually almost unno-

ticeable. Shell colour brownish with a

pattern of numerous ocellae which are a

little larger below the suture, mainly on

the last quarter of the shell.

Soft parts, operculum and radula

unknown.

Dimensions: The holotype is 5.0 mm;

the largest paratype reaches 6.0 mm.

Distribiition: Only known from

Principe Island.

Remarks: The differences with previ-

ously known species are as follows:

Anachis ciispidata (Marrat, 1877) has a

larger (about 7-8 mm) shell, the coloura-

tion is very variable, the ocellated

pattern very uncommon. Known from

Senegal.

Anachis avaroides Nordsieck, 1975 has

a larger and wider shell, variable in

colouration but usually lacking an ocel¬

lated pattern; the axial ribs are not

numerous, and it lacks spiral sculpture.

It is known from the Azores, Madeira,

Selvagem Islands and Canaries.

Anachis valledori Rolán & Tuque,

2002 has a smaller and narrower shell,

narrow and elongate axial ribs, variable

but not ocellated colouration. Endemic

to Cape Verde Islands.

Anachis ryalli Rolán, 2005 has a

larger and narrower shell, of uniform

tan color, the protoconch is more ele-

vated, spiral threads only noticeable in

the intervals. Collected in Ghana.

Anachis delaniarrei Rolán & Boyer,

2006 has a smaller shell, protoconch

with more whorls, fewer axial ribs and

it lacks spiral sculpture. Colour uni-

formly dark brown. Known from

Gabon.

Anachis delineata Rolán & de

Oliveira, 2008 has a larger shell, with

few axial ribs; the colouration comprises

bands with an ocellated pattern alternat-

ing with others with white blotches.

Known from the Cape Verde Islands.

Anachis jnani Horro & Rolán, 2010

has a shell with strong axial sculpture

but fewer ribs; the spiral cords are

strong, about 20 on the last whorl; it

lacks an ocellated pattern and has a

light band on the last whorl. It is from

Angola.

Mitrella annobonensis Rolán, 2005 is

rather similar, having less axial sculp¬

ture, less white spots, 14 shorter proto¬

conch with different microsculpture (see

Rolán, 2005: fig. 138-139). Perhaps this

species, due to its axial ribs, could be

placed in Anachis better than in Mitrella.

Genus Mitrella Risso, 1826

Remarks: In the same previously

mentioned works in which Anachis

species were described, species of the

genus Mitrella also appeared. This genus

ineludes a larger number of species, and

about 30 are presently known for the

area that ineludes Mediterranean, Mac-

aronesian and West African shores. In

this genus, there are species with large

distribution ranges, such as Mitrella ocel-

lata (Gmelin, 1791) which reaches the

West Atlantic coast, or Mitrella melvilli

Kudsen, 1956 from the West African

coast, which have a multispiral proto¬

conch. Most of the species, however, like

Mitrella broderipi (Sowerby, 1844) from

the Mediterranean and the Macarone-

sian archipelagos, have a short paucispi-

ral protoconch and several of these are

endemic to small areas.

56

RoláN and Gorí: New species of neogastropods from the islán ds of the Gulf of Guinea

Mitrella siciliai spec. nov. (Figs. 7-9)

Type material: Holotype (Fig. 7) in MNHN (24820). Paratypes in the following collections: MNCN

(15.05/60003, 1 s, Fig. 8), MHNS (100562 , 1 s), NHMUK (1 s), CFS (1); CPR (1); CSG (3 s, Fig. 9).

Type locality: Tinhosa Pequeña, 01° 20.33' N, 07° 17.35' E, under rocks, 25 m. Principe Island.

Etymology: Named after Francisco Sicilia Guillén, from Arrecife, Lanzarote, mollusc expert, friend

and companion of many diving trips.

Description: Shell (Figs. 7-9) solid,

broadly fusiform with a moderately high

spire. Protoconch with about 1^/2 whorls,

sharply pointed and about 0.65 mm in

máximum diameter, apparently smooth,

usually coloured with cream and a brown

line within. Teleoconch of between 5 - 5 V2

slightly convex whorls, with distinct su¬

ture and a very small subsutural shelf.

There is no any axial or spiral sculpture,

only growth lines. Towards the base about

10 well marked striae form evident cords,

the upper ones being wider. Aperture nar-

row and axially elongate, usually 6 teeth

on the inner part of external lip, the up¬

per one being larger, with a larger space

before the following two, which are closed.

Columella curved, S-shaped, with about

3 very small denticles, usually almost un-

noticeable. Shell colour brownish with a

pattern of numerous white ocellae, very

variable and more or less apparent from

one shell to another, and smaller white

blotches which are a little larger below

the suture. In the middle of the last whorl

there is a dark irregular band interrupted

by white circles and below it, these circles

are smaller and more numerous.

Dimensions: Fiolotype is 8,3 mm.

The paratypes are similar in size.

Distrihution: Only known from

Principe Island

Remarks: This species is slightly

similar to the following:

Mitrella psilla (Duelos, 1846) is a smaller

species present on the continental coast of

West Africa, the teleoconch has one whorl

less, the colour is lighter and more

uniform, and the aperture has 4-5 denti¬

cles on the inner part of the external lip.

Besides other morphological differ-

ences, Mitrella melvilli Knudsen, 1956

and M. africana Rolán, 2005, both have a

planktotrophic multispiral protoconch.

Mitrella inesitae Rolán, 2005 is larger,

more elongate, the second tooth on the

inner part of the external lip is larger,

the base is not concave, the colour is

lighter and the pattern different with a

light band in the middle of the last

whorl and brown blotches on the suture.

Mitrella aemulata Rolán, 2005 and M.

saotomensis Rolán, 2005 have smaller

and more fragüe shells, lighter in colour,

wider protoconch, and the small tuber-

cles on the columella more evident.

Mitrella annobonensis Rolán, 2005 is

smaller in size with a lighter colour, rel-

atively wider shell, with spiral striae

and 5 teeth on the inner part of the

external lip.

Mitrella noel Rolán & Gori, 2009 has

larger shells (up to 9 mm) being also

slender, more evident small tubercles on

the columella, protoconch lighter, colour

of the shell lighter presenting a different

pattern with a brown irregular area on

the upper part of the whorls.

Other Mitrella shells (Figs. 10-13)

In the material collected some other

shells with different patterns appeared.

The shells of the species of this genus are

sometimes very similar at specific level,

with an analogous colour and pattern.

Others however may have a different

pattern and variable colour. For this rea-

son, when the collected sample consists

of just a few specirnens, only very clear

differences can assure us that they are

different, whereas small differences may

only hint at specific separation, without

allowing US to decide whether we are

dealing with one or two sepárate species.

Mitrella cf. tenebrosa Rolán, 2005 (Fig.

10); collected in Minerio, in tangle nets.

57

Iberus, 30 (1), 2012

Figures 7-9. Mitrella siciliai spec. nov. 7: holotype, 8.3 mm (MNFíN); 8: paratype, 8.8 mm

(MNCN); 9: paratype, 8.3 mm (CSG). Figures 10-13. Other Mitrella. 10: Mitrella cf. tenebrosa

Rolán, 2005, 8.9 mm (CSG); 11, 12: Mitrella sp,, 8.3, 8.2 mm (CSG); 13: Mitrella cf. ;2í?^/Rolán

& Gori, 2009, 9.4 mm (CSG).

Figuras 7-9. Mitrella siciliai spec. nov. 7: holotipo, 8,3 mm (MNHN); 8: paratipo, 8,8 mm (MNCN);

9: paratipo, 8,3 mm (CSG). Figuras 10-13. Otras Mitrella. i 0.- Mitrella cf. tenebrosa Rolán, 2005,

8,9 mm (CSG); 11, 12: Mitrella sp., 8,3, 8,2 mm (GSG); 13: Mitrella cf. noel Rolán & Gori, 2009,

9,4 mm (CSG).

58

Rolan and GorI: New spedes of neogastropods from the islands of the Gulf of Guinea

Figures 14-17. Mitromorpha hernandezi spec. nov. A: 14, 15: holotype, 4.5 mm (MNCN); 16:

paratype, 4.4 mm (MNHN); 17: protoconch.

Figuras 14-17. Mitromorpha hernandezi spec. nov. A: 14, 15: holotipo, 4,5 mm (MNCN); 16: para¬

tipo, 4,4 mm (MNHN); 17: protoconcha.

48 m, Sao Tome, which is very similar to

the holotype.

Mitrella sp. (Figs. 11, 12). Two light

coloured shells, one of them with some

ocellae similar to the previous shell.

They were collected in Lagoa Azul, 00°

24' 49" N, 06° 36' 43" E, at 24 m, Sao

Tomé. They could be the same spedes as

the latter, but we do not really know the

full variability of this spedes.

Mitrella cf. noel Rolán & Gori, 2009.

These shells (Fig. 13), from Lagoa Azul,

00° 24' 49" N, 06° 36' 43" E, at 24 m, Sao

Tome, have a certain similarity to M. noel,

with a more minute pattern, but the vari¬

ability of this species is not well known.

59

Iberus, 30 (1), 2012

Family Mitromorphidae Casey, 1904

Genus Mitromorpha Carpenter, 1865

The species placed in this genus

were previously included in Mitrolumna

Bucquoy, Dautzenberg & Dollfus, 1883.

More recently, Mifsud (2001) made a

revisión of the species of the Mediter-

ranean and the Macaronesian archipela-

gos, employing for them the genus

Mitromorpha. CLEMAM also keeps this

genus as valid for these species.

West African species had been men-

tioned in the literature in scarce

numbers. Only four species were known

in the last century: M. monodi (Knudsen,

1956) from Senegal and M. crenipicta

(Dautzenberg, 1889), M. smithi Dautzen¬

berg & Fischer, 1896 from Azores. More

recently, a fourth species from deep

water was described by Bouchet &

Warén (1980). In the last ten years

Mifsud (2001) described several species

from the Mediterranean and the Mac¬

aronesian archipelagos. Rolán & Boyer

(2001) described two new species, one

from Senegal and another from Sao

Tome and presented also some morphs

considered as variations.

Mitromorpha hernandezi spec. nov. (Figs, 14-17)

Type material: Holotype (Figs. 14-15) in MNCN (15.05/60004). Paratypes in the following col-

lections: MNHN (24821, 1 s, Fig. 16); NHMUK (1 s); MHNS (100563, 7 s); MCZ (1 s), CPR (1 s);

CSG (2 s).

Other material examined: 10 s (in bad condition), 1 j, 21 f (MHNS), from type locality; 7 s, and f,

from San Antonio de Palé, 2-8 m.

Type locality: Isla Tortuga, Annobón Island, 12 m.

Etymology: The specific ñame is after José María Hernández, malacologist of Gran Canaria, recently

passed away, a frequent companion of the authors on trips and dives.

Description: Shell (Figs. 14-16) small,

solid, fusiform, almost biconical. Proto-

conch (Fig. 17) smooth and shiny, chest-

nut colored, with a nucleus of about 120

pm and a diameter of about 450 pm.

Teleoconch usually with 4-4 V2 whorls

bearing a coarse sculpture of axial ribs,

numbering 12-15 on the first whorl, 18-22

on the second and about 28 - 35 on the

last one. These ribs are crossed by three

spiral cords on the first whorls and 18-20

on the last. The Crossing of ribs and cords

gives the shell a reticulated aspect. Aper-

ture elongate axially, columella opistho-

cline, with 2-3 small denticles; on the

inner part of the external lip, there are

about 10-12 small teeth, some of them

more prominent. Colour brown and

white: brown on the subsutural area and

in the lower part of the last whorl below

the periphery. A band just on the separa-

tion of these two parts is formed by

blotches of brown and white. Towards

the base, the brown colour is reduced to

small rectangles on the spiral cords. In

some shells a similar line can also be seen

on the upper part of the shell.

Dimensions: Flolotype is 4.5 mm;

specimens may perhaps grow to be a

little larger, but most of them are

smaller, even between 3.3-4.0 mm

Distribution: Only known from

Annobón, from where it is supposed to

be endemic.

Remarks: In Rolán & Boyer (2001:

figs. 28-29, 42, 45) the present species

was figured as a morph of M. saotomen-

sis. At the time, not enough material

had been examined. Today, after having

the opportunity to study more shells,

we can point out the constancy of mor-

phological differences with M. sao-

tomensis. This and the isolation of both

populations on two islands many kilo-

meters apart, convinced us that M. her¬

nandezi is a valid different species.

Mitromorpha saotomensis has a shell

which is larger in size, darker, almost

ÓO

RoláN and GorI: New species of neogastropods from the islands of the Gulf of Guinea

black, very dense sculpture of axial ribs

and spiral cords.

Mitromorpha monodi (Knudsen, 1956)

is larger, whitish on the upper part, the

protoconch is white, and it is endemic to

Senegal.

Mitromorpha wilheminae Aartsen,

Menkhorst and Gittenberger, 1984 is

larger and wider, with a similar colour,

but the axial sculpture is faded; more-

over, it is endemic to the Mediter-

ranean.

Mitromorpha annobonensis spec. nov. (Figs. 18-23)

Type material: Holotype (Figs. 18-19) in MNCN (15.05/ 60005). Paratypes in the following collec-

tions: MNHN (24822, 1 s, Fig. 20); NHMUK (1 s); MHNS (100564, 7 s, Fig. 21); CPR (1 s); CSG (1 s).

Other material examined: 10 s (in eroded conditions) and 5 f, from the type locality.

Type locality: San Antonio de Palé, Armobon Isiand, 5-8 m.

Etymology: The specific ñame refers to the isiand where it was collected.

Description: Shell (Figs. 18-21) small,

solid, fusiform, almost biconical. Proto¬

conch (Figs. 22-23) smooth and shiny,

pink coloured, with a nucleus of about

150 pm and a diameter of about 410 pm.

There is a kind of blunt angle on the

subsutural part. Teleoconch usually

with prominent nodulous spiral cords

crossed by axial ribs forming nodules at

the intersection points. There are three

of these spiral cords on the first whorls

and on the last whorl, below the end of

the spire, there are 9-13 additional cords,

with slightly smaller nodules. There are

13- 16 axial ribs on the first whorl, and

14- 20 on the subsequent ones. Aperture

narrow and elongate, columella opistho-

diñe, almost straight and with very

small teeth in the middle. On the inner

part of the external lip there are 7-9

small teeth, of which the second on the

upper part is larger and more promi¬

nent. Colour pink with lighter nodules

on the upper part of the last whorl and

darker on band which continúes the

suture, and in some points more.

Towards the base, the colour is lighter.

Dimensions: Holotype is 4.3 mm,

like most of the paratypes.

Distribution: Only known from

Annobón, from where it is supposed to

be endemic.

Remarks: Mitromorpha annobonensis

spec. nov. was figured in Rolán &

Boyer (2001: figs. 26-27, 41 and 43) as a

form of M. saotomensis Rolán & Boyer,

2001. At present, after a careful compari-

son, we can State that they are different

species. It must be distinguished from

the closest species by the spiral angle of

the protoconch which was not found in

any other species. Other morphological

differences are:

M. saotomensis is darker, almost

black; the number of spiral cords on the

whorls is higher (21 to 30 on the last

one). Also the axial ribs are denser; the

protoconch is rounded, lacking any

angle and presenting micro tubercles.

M. monodi (Knudsen, 1956) is larger

and the upper middle part of the shell

and the protoconch are whitish.

M. hernandezi spec. nov. lives in sym-

patry and is smaller, more densely

sculptured in axial ribs and spiral cords,

the upper part of the last whorls is

lighter with separation from the lowest

dark one.

Mitromorpha suarezi spec. nov. (Figs. 24-25)

Type material: Holotype (Figs. 24) in MNHN (24823). Paratypes in the following collections: MNCN

(15.05/60006, 1 s, Fig. 24); NHMUK (1 s); MHNS (100565, 1 s); CMS (1 s), CPR (1 s), CSC (4 s),

Other material examined: 24 s eroded and f, from the type locality.

Type locality: Tinhosa Pequeña, 01° 20.33' N, 07° 17.35' E, under rocks, 25 m. Principe Isiand.

Etymology: The specific ñame is after Manuel Suarez Bustabad, enthusiastic malacologist from

Ferrol (A Coruña), Spain.

Ó1

Iberus, 30 (1), 2012

Figures 18-22. Mitromorpha annobonensis spec. nov. 18, 19: holotype, 4.3 mm (MNCN); 20:

paratype, 4.3 mm (MNHN); 21: paratype, 4.0 mm (MNHN); 22: apex with protoconch; 23:

detail of the protoconch.

Figuras 18-22. Mitromorpha annobonensis spec. nov. 18, 19: holotipo, 4,3 mm (MNCN); 20: para¬

tipo, 4,3 mm (MNHN); 21: paratipo, 4,0 mm (MNHN); 22: ápice con protoconcha; 23: detalle de la

protoconcha.

62

RoláN and GorI: New species of neogastropods from the islands of the Gulf of Guinea

Figures 24, 25. Mitromorpha suarezi spec. nov. 24: holotype, 4.7 mm (MNHN); 25: paratype, 4.6

mm (MNCN). Figures 26-29. Mitromorpha saotomensis (Rolán & Boyer, 2001); 26: holotype, 5

mm (MNCN); 27: paratype, 5.3 mm (MNHN). Both from Esprainha, Sao Tomé; 28, 29: shells,

4.3 and 4.9 mm, Sete Brazas, Principe ísland, 01° 39.13’ N, 07° 28.13’ E, 20 m.

Figuras 24, 25. Mitromorpha suarezi spec. nov. 24: holotipo, 4,7 mm (MNHN); 25: paratipo, 4,6

mm (MNCN). Figuras 26-29. Mitromorpha saotomensis (Rolán & Boyer, 2001); 26: holotipo, 5

mm (MNCN); 27: paratipo, 5,3 mm (MNHN). Ambas de Esprainha, Sao Tomé; 28, 29: conchas, 4,3

y 4,9 mm, Sete Brazas, Isla de Príncipe, OH 39. 13’ N, 07° 28. 13’ E, 20 m.

63

Iberus, 30 (1), 2012

Figures 30-34. Protoconchs oí Mitromorpha. 30: Mitromorpha suarezi spec. nov. 31-34: Mitromor-

pha saotomensis (Rolán & Boyer, 2001). 31: shell from Sete Brazas, Principe Is. (CSG); 32: detail

of the microsculpture; 33: shell from Principe, Tinhosa Pequeña; 34: shell from Sao Tomé, Baía

das Agulhas (from ROLÁN & BOYER, 2001).

Figuras 30-34. Protoconchas de Mitromorpha. 30: Mitromorpha suarezi spec. nov. 31-34: Mitro¬

morpha saotomensis (Rolán & Boyer, 2001). 31: concha de Sete Brazas, Isla de Príncipe (CSG); 32:

detalle de la microescultura; 33: concha de Príncipe, Tinhosa Pequeña; 34: concha de Sao Tomé, Baía

das Agulhas (de Rolán & Boyer, 2001).

64

RoláN and GorI: New species of neogastropods from the islands of the Gulf of Guinea

Description: Shell (Figs. 24-25) small,

solid, fusiform, almost biconical. Proto-

conch (Fig. 30) smooth and shiny,

coloured from pink to brown, with a

nucleus of about 110 pm and a diameter

of about 480 pm. Teleoconch usually

with 4-4 V2 whorls bearing a coarse

sculpture of axial ribs and spiral cords.

There are about 12 ribs on the first

whorl, 14-17 on the second and 20-28 on

the last one. There are three ribs on the

first whorls and on the last, about 14-16

more from the end of the spire. Aperture

narrow and elongate axially, columella a

little undulating with two small teeth;

on the inner part of the external lip,

there are about 10 teeth, the second one

larger and prominent. The colour is

cream on the upper part of the shell,

with an isolated brown rectangle on the

suture and darker below; towards the

base it is light again with small brown

rectangles on the spiral cords.

Dimensions: Holotype is 4.7 mm.

Distribution: Only known from the

type locality.

ACKNOWLEDGEMENTS

The authors thank Antonio A. Mon-

teiro of Lisbon, for the correction of the

English text. The optical photographs

were made in the Departamento de

BIBLIOGRAPHY

Bouchet P., Kantor Y.I., Sysoev A. & Puil-

LANDRE N. 2011. A new operational classifi-

cation of the Conoidea. Journal ofMolluscan

Studies, 77; 273-308.

Bouchet P. & Warén A. 1980. Revisión of the

North-East Atlantic bathyal and abyssal Tur-

ridae (Mollusca, Gastropoda). The Journal of

Molluscan Studies, suppl. 8 : 1-119.

Horro J. & Rolán E. 2010. A new species of

Anachis from Angola (Prosobranchia, Col-

lumbellidae). Gloria Maris, 49 (3-4): 64-67.

Mifsud C. 2001. The genus Mitromorpha Car-

penter, 1865 (Neogastr opoda, Turridae) and its

subgenera with notes on the European species. Au-

thor, Malta. 32 pp.

Pelorce J. & Rolán E. 2007. The type specimen

of Anachis richardi (Gastropoda, Columbell-

idae) from the Cape Verde Archipelago. Noti¬

ciario SEM, 48: 40-41.

Remarks: Mitromorpha suarezi spec.

nov. has some similarity to M. hernandezi

spec. nov., in the pattern and colouration

but they can obviously be distinguished:

M. hernandezi has a slightly smaller shell,

its colour is lighter with yellowish on the

upper part of the whorls; the protoconch

is a little narro wer.

Mitromorpha saotomensis (Rolán &

Boyer, 2001) has a similar shape, a

darker shell, being more densely sculp-

tured. It could be considered a variation,

but in the same place where M. suarezi

was collected, there is also a population

of the typical M. saotomensis (Figs. 28-29)

living sympatrically without intergrada-

tions. This population is more similar to

that of Sao Tome only being a little

smaller. In Rolán & Boyer (2001: figs.

33-34) also were represented shells from

Principe 1. (Baía das Agulhas). In this

last species the presence of tubercles on

the protoconch was noticed several

times (Figs. 31-34). This could not be

shown in M. suarezi but perhaps the pro-

toconchs studied were a little eroded.

Xenética, of the University of Vigo; the

SEM micrographs were made by Jesús

Méndez and Inés Pazos of the CACTI of

the University of Vigo.

Rolán E. & Boyer F. 2001. The genus

Mitrolumna (Gastropoda, Turridae) in West

Africa. Iberus, 19 (19): 115-128.

Rolán E. & Boyer F. 2006. A new Anachis (Gas¬

tropoda: Columbellidae) from Gabon. No-

vapex, 7 (1): 25-27.

Rolan E. & Luque A. A. 2002. Two new species

of Columbellidae (Gastropoda: Buccinoidea)

from the Cape Verde Archipelago. Iberas, 20

(1): 73-83.

Rolán E. & Oliveira Á. de, 2009 «2008». A

new species of Anachis (Prosobranchia, Co¬

lumbellidae) from Cape Verde Archipelago.

Gloria Maris, 47 (4): 67-72.

Rolán E. 2001. A new species of Mitrella

(Neogastropoda, Columbellidae) from the

Cape Verde Archipelago. La Conchiglia, 33

(301); 11-12.

65

Iberus, 30 (1), 2012

Rolán E. 2004. A new species more oíMitrella

(Neogastropoda, Columbellidae) from the

Cape Verde Archipelago. La Conchiglia, 34

(311): 21-23.

Rolán E. 2005. Columbellidae (Gastropoda,

Neogastropoda) of the gulf of Guinea with

the description of eight new species. Iberus,

23 (2): 119-156.

Rolán E. 2005. Malacological Fauna from the Cape

Verde Archipelago. Conchbooks, Vigo. 482 pp

y 85 pls.

Rolán E. & Gori S. 2009. Two new species of

the genera Jujubinus and Mitrella (Mollusca,

Prosobranchia) from Sao Tome Island. Glo¬

ria Maris, 48 (1): 10-16.

66

© Sociedad Española de Malacología

Iberas, 30 (1): 67-83, 2012

The genus Gibberula (Gastropoda, Cystiscidae) in the Cape

Verde Islands with the description of a new species

El género Gibberula (Gastropoda, Cystiscidae) en el archipiélago de

Cabo Verde con la descripción de una nueva especie

Diego MORENO*

Recibido el23-V-201L Aceptado el 18-XI-2011

ABSTRACT

The species of the genus Gibberula from the Cape Verde islands are reviewed. There are

at least four species, three previously described: Gibberula lucia Jousseaume, 1 877, G.

rauli Fernandes, 1987 and G, rolani Cossignani and Cecalupo, 2005, and one new

species, G. elvirae n. sp. is here described. Gibberula rachmaninovi Keüner, 2003, is

considered here as a synonym of Volvarina sauliae (Sowerby, 1 846).

RESUMEN

Se revisan las especies del género Gibberula del archipiélago de Cabo Verde. Existen, al

menos, cuatro especies, tres descritas previamente: Gibberula lucia Jousseaume, 1 877,

G, rauli Fernandes, 1987 y G. rolani Cossignani y Cecalupo, 2005, y una especie nueva

que se describe en este trabajo: G. elvirae n. sp. Por último, Gibberula rachmaninovi Kell-

ner, 2003, es considerado un sinónimo de Volvarina sauliae (Sowerby, 1 846).

INTRODUCTION

The genus Gibberula belongs to the

Family Cystiscidae which, together with

the closely related Family Marginellidae,

are known as "MarginelliForm" gas-

tropods, and are one oF the most con-

spicuous elements oF the West AFrican

malacoFauna.

The moUuscan Fauna oF the Cape Verde

Islands has been studied by many authors:

Jousseaume (1877), Rochebrune (1881a,

b) , Dautzenberg (1910), Melvill and

Standen (1913), Burnay and Monteiro

(1977), Saunders (1977), García-Talavera

AND Bacallado (1979), CosEL (1982a, b,

c) , Fernandes (1987), Moreno and

Burnay (1999), Guerreiro and Reiner

(2000), Kellner (2003), Rolán (2005) and

Cossignani and Cecalupo (2005).

There are six species ñames For Gib¬

berula listed in the literature, either

described with a type locality in the

Cape Verde Islands or cited For the

Archipelago. The Mediterranean species

Gibberula miliaria (Linnaeus, 1758) and

G, philippii (Monterosato, 1878) were

erroneously cited in the Cape Verde

Islands and are not present in the archi¬

pelago. Another species, Gibberula rach¬

maninovi Kellner, 2003, described From

the Cape Verde islands as a Gibberula

will here be shown to belong to a diFFer-

ent genus, Volvarina, The genus Gib¬

berula is represented in the Cape Verde

Archipelago by Four species, either

endemic, as G. rauli Fernandes, 1987, G,

rolani Cossignani and Cecalupo, 2005,

* c/ Araña, apartamentos Las Dunas 2, 04150 Cabo de Gata, Almería (Spain).

67

IberuSy 30 (1), 2012

Table I. Localities studied in the Cape Verde Archipelago with species of the genus Gib bernia,

showing the coordinares and the origin of the samples (see ahhreviations used).

Tabla 1. Localidades de estudio en el archipiélago de Cabo Verde con especies del género Gihherula,

mostrando las coordenadas y origen de las muestras (ver abreviaturas utililizadas).

and the new species described in the

present paper, or with a very restricted

distribution area, as G. lucia

Jousseaume, 1877, cited also in Senegal

(Knudsen, 1956; Pin and Boyer, 1995).

The pigmentation patterns of the

head-foot and mantle in the living

animáis are usually bright and have been

shown to be stable and extremely infor-

mative for the separation of closely

related species (Fern andes, 1987; Gofas,

1989a, 1989b, 1990; Moreno and

Burnay, 1999) and will be given particu¬

lar attention in this paper. Also the radula

features are important for separating

species (Moreno and Burnay, 1999).

MATERIAL AND METHODS

Most of the material examined was

collected during the "Primera Expedi¬

ción Ibérica a Cabo Verde" (PEICV)

excursión to the Cape Verde Islands, in

which the author took part (August

1985). Specimens were hand-picked

while scuba-diving or snorkeling

between 0 and 5 m depth in the islands

of Sal, Boavista and Sao Vicente. Most

specimens were collected between -1

and -2 meters on rocky bottoms with

coral slabs. Other islands were visited

(Santa Luzia, Sao Nicolau and Santiago)

but only a limited sampling effort could

be carried out (snorkeling) and no speci¬

mens of this genus were obtained.

Other material studied was collected

by L.P Burnay in the years 1975-1976 (IIT),

by R. von Cosel in 1978 (Sao Vicente Island,

at MNHN), by E. Rolán in the years 1978,

1979, 1980, 1981, 1986, 1987, 1988 and 2001

(Sal, Boavista and Brava), and by A.D. De

Oliveira in 2005 (Santiago). Specimens of

Gibberula oryza (Lamarck, 1822), the type

species of the genus, were studied and

photographed alive at Punta Carnero,

Cádiz, in the Strait of Gibraltar (1997). All

localities studied are shown in Table I and

in the map of Figure 1.

The clarification of the complex taxo-

nomic problems in the genus is mainly

based on the observation of pigmentation

patterns, and therefore required collect-

ing and sorting the animáis alive. The

68

Moreno: The genus Gibberula in the Cape Verde Islands; description of a new species

Figure 1 . Map of Cape Verde Archipelago and localities where the studied material of the genus

Gibberula was collected (origin of the samples in brackets). Coordinates in Table I. Sal Island, 1:

Ponta Preta (ER); 2; Rigona (PEÍCV) and (ER); 3: Baia Palmeira (PEICV); 4: Rabo de Junco

(ER); 5: Baia Mordeira (ER); 6; Baia Algodoeiro (ER); 7: Santa Maria (PEICV); 8: Fiura

(PEICV); 9: Palhona (ER); 10: Monte Leste (ER). Sao Vicente Island, 11; Salamanza (PEICV);

12: Calhau (RVC-MNHN). Boavista Island, 13: Baia Sal Rei (PEICV); 14: Derrubado (ER). San¬

tiago Island, 15: Praia (AO). Brava Island, 16: Baia Pedrinha (ER); 17: Fuma (ER).

Figura 1. Mapa del archipiélago de Cabo Verde y localidades donde se recolectó el material estudiado del

género Gibberula (origen de las muestras entre paréntesis). Coordenadas en la Tabla /. Isla de Sal, 1:

Ponta Preta (ER); 2: Rigona (PEICV) y (ER); 3: Baia Palmeira (PEICV); 4: Rabo de Junco (ER); 5:

Baia Mordeira (ER); 6: Baia Algodoeiro (ER); 7: Santa Maria (PEICV); 8: Fiura (PEICV); 9:

Palhona (ER); 10: Monte Leste (ER). Isla de Sao Vicente, 11: Salamanza (PEICV); 12: Calhau (RVC-

MNHN). Isla de Boavista, 13: Baia Sal Rei (PEICV); 14: Derrubado (ER). Isla Santiago, 15: Praia

(AO). Isla Brava, 16: Baia Pedrinha (ER); 17: Fuma (ER).

living animáis were observed under a

stereomicroscope, and coloured draw-

ings prepared to take note of the patterns

and tones of pigmentation. In some cases,

colour slides were also taken.

The radulae were extracted from

adult specimens using potassium

hydroxyde on whole animáis soaked in

water. At least one radula of each

species was observed through a sean-

ning electrón microscope (MNCN).

Abbreviations used

AO: Alvaro David De Oliveira Ramos

and his private collection (Gulpil-

bares, Portugal)

ER: Emilio Rolán and his private collec¬

tion (Vigo, Spain).

IIT: Instituto de Investigagáo Tropical

(Centro de Zoología), Lisboa, Portu¬

gal.

MNCN: Museo Nacional de Ciencias

Naturales, Madrid.

MNHN: Muséum National d'Histoire

Naturelle, Laboratoire de Malacolo-

gie, Paris, France.

PEICV: Primera Expedición Ibérica al

Archipiélago de Cabo Verde (1985).

RVC: Rudo Von Cosel (MNHN, Paris,

France).

UAM: Universidad Autónoma de

Madrid, Spain (Departamento de

Biología, Zoología).

69

Ihems, 30 (1), 2012

SYSTEMATIC PART

Family Cystiscidae Stimpson, 1865

Genus Gibberula Swainson, 1840

Type species Gibberula zonata Swainson, 1840 by monotypy (= Volvaria oryza Lamarck, 1822), as

stated Gofas (1990: 115) (Fig. 17).

The Shell of the species of the genus

Gibberula is ovate-cylindrical, sometimes

somewhat pyriform with a very devel-

oped body whorl covering almost all of

the earlier whorls. The shell is translu-

cent in the smaller species, so that the

inner mantle may be visible by trans-

parency. The protoconch is paucispiral

as in all marginelliform gastropods. The

aperture is elongate and narrow and has

anteriorly a rounded siphonal canal.

The columella has plaits on its anterior

half, decreasing in size towards the

apical end. The outer lip, in adults, is

always thickened, with usually numer-

ous denticles on the inner margin. The

shell surface is glossy, without colour,

with spiral pigmented bands or more

rarely with wavy axial flames.

The animal (type 4 following

CoovERT AND CoovERT, 1995) is active,

fashmoving, and creeps on the foot

which is truncated anteriorly and

rounded posteriorly. The head is small,

deeply cleft in two parts and provided

with a pair of short tentados. The eyes

are small, without lateral bulges as in

the Marginellidae. The siphon lies over

the head and it is cleft on its lower side.

The mantle does not cover the shell as

other marginelliform gastropods. The

pigmented areas are on the upper part

of the foot, the head, the siphon and ten¬

tados, and the mantle. The most

common colours are black, orange and

opaque white. The upper part of the

foot, siphon and head are usually more

brightly pigmented. When the animal is

withdrawn, the pigmentation of the

inner mantle may be seen through the

shell.

The radula (type 3 following

COOVERT AND COOVERT, 1995), is

rachiglossan, reduced to a central

rachidian tooth per Une, which is V-

shaped, between 0.008 and 0.042 mm

wide, with several cusps (5-11). The

ratio shell length/ tooth width (L/Wr) in

the genus is between 118 and 309

(COOVERT AND COOVERT, 1995).

Gibberula lucia Jousseaume, 1877 (Figs. 2-4)

Gibberula lucia Jousseaume, 1877: 269-270, pl. V, figs. 11-13 (Santa Luzia Island, Cape Verde

Archipelago).

Other references:

Marginella (Glabella) lucia (Jousseaume, 1876) [sic]: Tryon, 1883.

Gibberula lucia Jousseaume, 1877: Tomlin, 1917; p. 277.

Marginella lucia (Jousseaume, 1876) [sic]: Knudsen, 1956; p. 88, Pl. 1, Fig. 4 (Gorée, Senegal).

Marginella lucia (Jousseaume, 1876) [sic]: Wagner and Abbott, 1967; p. 156.

Gibberula lucia Jousseaume, 1876 [sic]: Pin and Boyer, 1995; p. 56, Figs. 7-9 (Dakar, Senegal).

Other references (doubtful);

IGibberula miliaria (Linnaeus, 1758): Rochebrune, 1881b; p. 293 (Porto-Praya in Santiago Island,

Cape Verde Archipelago).

?Persicula miliaria (Monterosato) [sic]: García-Tala vera and Bacallado, 1979; p. 207 (Mindelo and

Gatas in Sao Vicente Island and Praia in Santiago Island, Cape Verde Archipelago).

Type material: 3 syntypes (MNHN) all of 3.5 mm length (Fig. 3 in the present paper).

Other material studied: Santiago Island: Praia, 1 sp., broken (AO) (Fig. 4).

70

Moreno: The genus Gibberula in the Cape Verde Islands; description of a new species

Figures 2-4. Gibberula lucia. 2: figures 11, 12 and 13 of the original description by Jousseaume

(1877); 3: three syntipes of G. lucia from Santa Luzia Island (MNFIN) (3.5 mm); 4: specimen

from Praia, Santiago Island collected by A. Oliveira (23-VII-2005) (4.0 mm).

Figuras 2-4. Gibberula lucia. 2: figuras 11, 12 y 13 de la descripción original de Jousseaume (1877);

3: tres sintipos de G. lucia de la isla de Santa Luzia (MNLIN) (3,5 mm); 4: ejemplar de Praia, isla de

Santiago, recogido por A. Oliveira (23-VII-2005) (4,0 mm).

Remarks: The shell descriptions of

Jousseaume (1877) and Knudsen (1956)

are good. The figure of this species by

Jousseaume (1877), reproduced here in

Fig. 2, is deficient, but that of Knudsen

(1956) from Senegal is better. Pin and

Boyer (1995) also made a description of

shells from Dakar (Senegal) showing

photographs of several specimens.

There are no data about the animal and

its colour pattern or radula.

Distribution: With the scarce data

available, the species lives in Santa

Luzia Island (Cape Verde Archipelago),

and in the area of Dakar (Senegal),

including Gorée Island, where it seems

to be more abundan!. So, this is the solé

species of the genus living in the Cape

Verde Islands that is not considered

endemic. It still has a very restricted

area of distribution, because Dakar is

the closest point of the African continent

to the Cape Verde Islands.

Discussion: The species, described

with material from the desertic island of

Santa Luzia by Jousseaume (1877), has a

shell with a very distinct colour pattern

of brown lines in four groups: two of

them comma-shaped that converge in

the central part of the body whorl,

another group near the suture and the

last one along the base of the shell. This

colour pattern matches that of the speci¬

mens from Gorée, Dakar (Senegal)

71

Iberus, 30 (1), 2012

studied by Knudsen (1956). There has

been no more material of this species

recorded from the Cape Verde Islands

since the description by Jousseaume

(1877), in spite of the numerous malaco-

logical expeditions to this area made in

the last decades of the XX Century The

species was not cited by several authors

who studied the malacofauna of the

Cape Verde Islands, including Cosel

(1982b) in a work about the molluscan

fauna of Santa Luzia, the type locality of

this species. Nevertheless a single speci-

men (Fig. 5) collected by A. Oliveira at

Praia (Santiago Island) (23-VII-2005),

confirms the presence of the species in

the Cape Verde Islands, and silences the

possible doubts that could exist about

the Jousseaume material.

Rolán (2005) reports the ñame Gíb-

berula lucia Jousseaume, 1876 [sic] erro-

neously as a synonym of Gibberula sp. 2.,

but not as a valid species. The species

cited as Gibberula sp. 2 by Rolán (2005)

was described in the same year as Gib¬

berula rolani by Cossignani and Ce-

CALUPO (2005), and it is hereafter treated

as a valid species, different from G. lucia.

Gibberula rauli Fernandes, 1987 (Figs. 5, 6, 14)

Gibberula rauli Fernandes, 1987: 265, figs. 3 and c (Sal, Boavista, Maio, Santiago and Sao Vicente

islands, all in the Cape Verde Archipelago).

Gibberula philippii (Monterosato, 1878) [misidentification]: García-Talavera and Bacallado, 1979;

p. 207 (Mindelo in Sao Vicente Island, Cape Verde Archipelago).

Gibberula rauli Fernandes, 1987: Rolán, 2005; p. 147, Figs. 642 and 778 (from throughout the Cape

Verde Archipelago).

Material studied (all from the Cape Verde Islands): Boavista Island: Derrubado, 2 sp. (ER). Sal

Island: Rabo de Junco, 2 sp. (ER). Rigona, 2 sp. (ER). Algodoeiro, 2 sp. (ER). Sao Vicente Island:

Salamanza, 75 sp. + 10 sp. (PEICV). Brava Island: Baia Pedrinha, 137 sp. (ER). Fuma, 5 sp. (ER).

Remarles: The shell description by Fer¬

nandes (1987) is correct. This author

included drawings of the shell and of the

living animal. The shell, which is very

transparent, is here shown photographed

(Fig. 5). The animal is active and fast. The

foot, rounded in its rear end, is longer

than the shell when the animal is moving

on the substratum. The head is small,

divided into two parts, and the tentacles

are very short. The pigmentation pattern

(Fig. 14) on the foot and on the inner

mande observed by transpareney, is

white, with extensive black areas, very

irregular in shape in the mande. The

siphon is white.

(Right page) Figures 5-13. Shells and radulae of species of Gibberula from Cape Verde Islands. 5,

6. Gibberula rauli, Salamanza, Sao Vicente (PEICV). 5: shells (2.2 and 2.1 mm); 6: radula. 7-11.

Gibberula rolani. 7: shell from Sal Rei, Boavista (PEICV) (5.1 mm); 8: shell from Rigona, Sal

(PEICV) (5.0 mm); 9: radula from Sal Rei, Boavista (PEICV); 10, 11: teratological double radula

from Sal Rei, Boavista (PEICV); 10: double radula showing the odontophoral cartilage hoods

typical of a cistiscid radula; 11: teeth details of double radula. 12, 13. Holotype of Gibberula

elvirae n. sp. from Salamanza, Sao Vicente (PEICV). 12: shell (4.8 x 3.0 mm); 13: radula.

(Página derecha) Figuras 5-13. Conchas y rádulas de especies de Gibberula de las islas de Cabo Verde.

5, 6. Gibberula rauli, Salamanza, Sao Vicente (PEICV). 5: conchas (2,2 y 2,1 mm); 6: rádula. 7-11.

Gibberula rolani. 7; concha de Sal Rei, Boavista (PEICV) (5,1 mm); 8: concha de Rigona, Sal

(PEICV) (5,0 mm); 9: rádula de Sal Rei, Boavista (PEICV); 10, 11: rádula teratológica doble de Sal

Rei, Boavista (PEICV); 10: rádula doble mostrando la forma de capucha del cartílago odontoforal,

típica de una rádula de cistíscido; 11: detalle de los dientes de la rádula doble. 12, 13. Holotipo de

Gibberula elvirae n. sp. de Salamanza, Sao Vicente (PEICV). 12: concha (4,8 x 3,0 mm); 13: rádula.

72

Moreno: The genus Gibberula in the Cape Verde Islands; description of a new species

73

Iberus, 30 (1), 2012

The radula is described and figured

here for the first time (Fig. 6). Radular

teeth have an arched form with a central

cusp prominent but narrower than some

lateral cusps. On each side of the central

cusp there is a small one, followed by

three lateral cusps, the first one large and

strong, and then two external ones,

smaller. There are 9 cusps in total. The

teeth are 0.017 mm width. As the shell

length of the specimen dissected for

radular study is 2.0 mm, the "shell

length/tooth width" ratio (L/Wr) is 118.

Habitat: This species lives in the

upper zone of the sublittoral, between 1

and 5 meters depth, on rocky bottoms

with sand and algae.

Distribution: An endemic species

from the Cape Verde Islands. Rolán

(2005) stated that G. rauU is known from

the entire archipelago. The islands

reported in the bibliography are: Sal

(type locality at Rabo de Junco Beach),

Boavista, Maio, Santiago and Sao

Vicente. In the present work it is cited

also from Brava Island. Gibberula philip-

pii is a Mediterranean species (Gofas,

1990) erroneously cited from the Cape

Verde Islands by García-Talavera and

Bacallado (1979).

Gibberula rolani Cossignani and Cecalupo, 2005 (Figs. 7, 8, 9, 10, 11 and 15)

Gibberula sp. 1: Rolán, 2005; p. 148, only part (Figs. 643-644) (endemic from the Cape Verde

Islands).

Gibberula sp. 2: Rolán, 2005; p. 148, (Fig. 645) (endemic from the Cape Verde Islands).

Gibberula rolani Cossignani and Cecalupo, 2005: p. 6-7, 6 Figs. (Mordeira in Sal Island, Cape

Verde Archipelago).

Other references (doubtful: could also be G. elvirae or G. lucia):

IGibberula miliaria (Linnaeus, 1758): Rochebrune, 1881b; p. 293 (Porto-Praya in Santiago Island,

Cape Verde Archipelago).

IGibberula orijza Lamarck, 1822: Dautzenberg, 1910; p. 43-44 (Cape Verde Archipelago).

IGibberula miliaria (Linnaeus, 1758): Melvill and Standen, 1913: p. 342 (Sao Vicente, Cape Verde

Archipelago).

Wersicula miliaria (Linnaeus, 1758): Saunders, 1977; p. 14 (Matiota in Sao Vicente and Palmeira in

Sal Island, Cape Verde Archipelago).

?Persicula miliaria (Monterosato) [sic]: García-Tala vera and Bacallado, 1979; p. 207 (Mindelo and

Gatas in Sao Vicente Island and Praia in Santiago Island, Cape Verde Archipelago).

IGibberula miliaria (Linnaeus, 1758): Cosel, 1982a; p. 20 (Sao Vicente, Sal and Santiago Islands,

Cape Verde Archipelago).

IGibberula miliaria (Linnaeus, 1758): Cosel, 1982c; p. 57 (Cape Verde Archipelago).

Material studied (all from Cape Verde Islands): Boavista Island: Sal Rei, 6 sp. + 1 juv. + 1 juv. + 1

juv. (UAM). Derrubado, 2 sp. + 1 juv (ER). Sal Island: without locality, 6 sp. + 1 sp. + 1 sp. (IIT),

Palmeira, 1 sp. + 1 sp. + 2 sp. (UAM). Ponta Preta, 10 sp. + 9 juv. (ER). Palhona, 6 sp. + 4 sp. + 4 sp.

+ 4 juv. (ER). Rabo de Junco, 27 sp. + 10 juv. (ER). Mordeira, 5 sp. + 1 sp. (ER). Monte Leste, 8 sp.

+ 3 sp. (ER). Santa María, 1 sp. (UAM). Rigona, 1 sp. + 1 sp. (UAM), 6 sp. + 1 juv. (ER). Algodoeiro,

4 sp. (ER). Fiura, 1 sp. (UAM). ?Brava Island: Baia Pedrinha, 1 sp. (ER).

Description: The species description

by Cossignani and Cecalupo (2005) is

basically correct, but based only on the

shell. This is pyriform, narro w in the an¬

terior end with a broad and rounded

body whorL There are 5-6 plaits in the

columella, decreasing in size posteriorly,

and 14-18 small denticles on the inner

side of the outer lip. In general the shell

has a dark brown sutural band, not al-

ways present. Several specimens are

plain white (Fig. 7) or a palé yellow, and

other shells have two or three spirals

bands of a very palé brown (Fig. 8).

Some specimens have one broad spiral

and brown band in the central part of the

body whorl, a result of the fusión of both

anterior pigmented lines.

74

Moreno: The genus Gibberula in the Cape Verde Islands; description of a new species

The animal, described here for the

first time (Fig. 15, from Boavista Island

material) is active and slithers fast on

the substratum with the foot, which is

larger than the shell when totally

extended. The foot is ovate in the

metapodium and truncated in the

propodium, translucenhwhite with

peripheral orange lines and patches,

and a conspicuous central line on the

posterior end (Fig. 15). Some specimens

do not have the metapodium pig-

mented, but have orange lines in the

propodium. The head is small with a

central split. There is one orange line on

each side of the head from the inner part

of the eyes to the anterior end of the

snout. The tentados are short, with a

small and central band, also orange. The

siphon is small, of a palé yellow colour.

The inner mantle, that can be observed

by transparency, is also very variable,

dark with white lobes, orange with

white patches, or clear with dark or

orange dots.

The radula is described and figured

here for the first time (Fig. 9, from Boa-

vista Island material). Radular teeth

were studied from two specimens. The

teeth have an arched form with a promi-

nent central cusp. There are on each side

of the central cusp a small one (not

always present) and 4 (3 or 5 in some

cases) strong lateral cusps (there are 9-

10 cusps in total). The same pattern of

cusps are observed every three teeth, so

that two adj acent teeth never have the

same cusps, and the lateral cusps of

similar length do not match in the same

position. A teratological specimen was

observed with two complete radulae in

the same bucal bulb (Figs. 10-11), one of

them placed over the other. The teeth of

both radulae are similar to that previ-

ously described for a normal specimen.

The average tooth width is 0.026 mm.

As the average shell length of the speci¬

mens is 5.0 mm, the "shell length/tooth

width" ratio (L/Wr) for this species is

192.

Habitat: This species lives in the

upper sublittoral area, between 1 and 5

m depth, where it was observed under

stones on rocky bottoms with sand.

Distribution: G. rolani is an endemic

species of the Cape Verde Islands. We

have found this species only on Sal and

Boavista islands, where it is frequent.

We studied also a single specimen from

Brava Island that probably belongs to

this species, with a similar form and 2

spiral bands of palé brown, one central

and another on the base. Some of the

references of Gibberula miliaria from the

Cape Verde Islands (considered here as

corresponding to G. rolani) cited other

islands in its distribution, such as Santi¬

ago (Rochebrune, 1881b; Cosel, 1982a)

and Sao Vicente (Saunders, 1977;

García-Talavera and Bacallado,

1979; Cosel, 1982a). If G. rolani really

lives on Santiago and Sao Vicente, a fact

that should be confirmed with new

material, the species would achieve a

distribution throughout all the island

groups of the archipelago.

Discussion: The description of this

species by Cossignani and Cecalupo

(2005) was made immediately (Novem-

ber 2005) after the publication in April

of the same year of the book by Rolán

(2005) on the malacological fauna of the

Cape Verde Archipelago. Rolán (2005)

included two Gibberula species without

specific ñames (sp. 1 and sp. 2, on page

148), stating that they were in course of

description. The hasty description of

Gibberula rolani by Cossignani and

Cecalupo (2005) disregards the ethical

recommendation of the International

code of Zoological Nomenclature

because they knew the situation regard-

ing the case and stated that their new

species was present in the book by

Rolán (2005) as '"Gibberula sp. 1 (Figs.

643-644, 776) and Gibberula sp. 2 (Fig.

645) pag. 148".

It is most probably this species

which has been misidentified as Gib¬

berula miliaria (Linnaeus, 1758) by

several authors, but G. miliaria lives only

in the Mediterranean Sea (Copas, 1990).

There are obvious differences in the pig-

mentation pattern between both species,

particularly on the foot, where G. rolani

has a clearly defined pattern with

peripheral orange lines and patches.

Also, the head of G. rolani has single

75

Iberus, 30 (1), 2012

Figures 14-16. Live animáis of Gibberula species from Cape Verde Islands. 14: Gibberula rauli,

Salamanza, Sao Vicente (PEICV) (2.2 mm); 15: Gibberula rolani, Sal Rei, Boavista (PEICV) (5.0

mm); 16: holotype of Gibberula elvirae n. sp., Salamanza, Sao Vicente (PEICV) (4.8 mm).

Figuras 14-16. Animales vivos de especies de Gibberula de las islas de Gabo Verde. 14: Gibberula rauli,

Salamanza, Sao Vicente (PEIGV) (2,2 mm); 15: Gibberula rolani. Sal Rei, Boavista (PEIGV) (5,0

mm); 16: holotipo de Gibberula elvirae n. sp. Salamanza, Sao Vicente (PEIGV) (4,8 mm).

orange lines and a small band in the ten-

tacles, whereas in G. miliaria the foot

and head patterns are more elabórate,

with different colours and small dots

(CoovERT, 1987; Gofas, 1990). The

radula of G. miliaria was studied by

Bandel (1984, Fig. 297: as Persicula mi¬

liaria, from Banyuls-sur-Mer, France,

Mediterranean Sea) and reproduced by

CoovERT (1989, Fig. 19). The teeth of G.

rolani are similar to those of G. miliaria,

but in the former the lateral cusps are

stronger than in the latter species. Also,

the teeth of G. rolani have 8-9 cusps

against 9-11 in G. miliaria.

About the teratological radulae

studied (Figs. 10-11) it seems that in this

strange specimen there are two radular

sacs, instead of one, but only a pair of

odontophoral cartilage hoods (Fig. 10),

characteristic of the "cysticid radula", so

called by Coovert and Coovert (1995).

There is no Information of a similar case

known in the marginelliform gas-

tropods (Coovert and Coovert, 1995).

Dautzenberg (1910) included the

Cape Verde Islands in the distribution of

the species Gibberula oryza, but this

species lives only on the continental

African coasts from Senegal to the Strait

of Gibraltar (Gofas, 1990). Some shell

patterns of G. rolani, like those with a

broad band remind of the neotype of G.

oryza figured by Gofas (1990, Fig. 2), or

shells without bands, which look like

specimens of G. oryza from Ceuta (Fig. 3

in Gofas, 1990). Flowever, the shell of G.

rolani is smaller (up to 5 mm length) than

that of G. oryza (up to 8 mm length). The

colour pattern of the animal is also diffe¬

rent between these two species; G. rolani t

has only orange lines and patches, |

76

Moreno: The genus Gibberula in the Cape Verde Islands; description of a new species

Figures 17-18. Live animáis of Gibberula species. 17: Gibberula oryza. Punta Carnero, Cádiz,

Spain (7 mm); 18: holotype of Gibberula elvirae n. sp., Salamanza, Sao Vicente (PEICV) (4.8

mm).

Figuras 17-18. Animales vivos de especies de Gibberula. 17: Gibberula oryza. Punta Carnero, Cádiz,

España (7 mm); 18: holotipo de Gibberula elvirae n. sp. Salamanza, Sao Vicente (PEICV) (4,8 mm).

restricted in general to the pedal margin

of the foot, while in G. oryza the pigmen-

tation pattern is outspread, with orange,

yellow and violet small dots over all the

foot (Gofas, 1990: Pl. 1, Figs. a-a'; see also

Fig. 17 in the present paper). The radula

of G. oryza of a specimen from Ceuta was

studied by Gofas (1990, Fig. 1). The

arched form of the teeth and the number

of cusps are very similar between G.

rolani and G. oryza (9-10 and 11 cusps.

respectively), but in the first species the

tooth width is significantly smaller (0.026

mm) than that of the second (0.038 mm).

The "shell length/tooth width" ratio

(L/Wr) is similar in both species, 192 and

187 respectively, but the length shell of

the first species is 5 mm, while in G. oryza

it is more or less 7 mm (Gofas, 1990 did

not State the shell length of the specimen

dissected for obtention of the radula

figured).

77

Iherus, 30 (1), 2012

Gibberula elvirae n. sp. (Figs. 12, 13, 16 and 18)

Gibberula sp. 1: Rolán, 2005; p. 148, only part (Fig. 776) (endemic from Cape Verde Island).

Other references (doubtful);

IGibberula oryza Lamarck, 1822: Dautzenberg, 1910; p. 43-44 (Cape Verde Archipelago).

IGibberula miliaria (Lirmaeus, 1758): Melvill and Standen, 1913: p. 342 (Sao Vicente, Cape Verde

Archipelago).

IPersicula miliaria (Linnaeus, 1758): Saunders, 1977; p. 14 (Matiota in Sao Vicente and Palmeira in

Sal Island, Cape Verde Archipelago).

IPersicula miliaria (Monterosato) [sic]: García-Talavera and Bacallado, 1979; p. 207 (Mindelo and

Gatas in Sao Vicente Island and Praia in Santiago Island, Cape Verde Archipelago).

IGibberula miliaria (Linnaeus, 1758): Cosel, 1982a; p. 20 (Sao Vicente, Sal and Santiago Islands,

Cape Verde Archipelago).

Type material: Holotype, specimen 4.8 x 3.0 mm, from Salamanza, Sao Vicente (Cape Verde Islands),

from a lot of one specimen (UAM no. 229) collected on 20-VIII-1985 by PEICV (deposited in MNCN,

no. 15.05/60007); Paratypes 1 and 2 (the latest with the siphonal canal broken) from Baia do Calhau,

Sao Vicente, from a lot of five specimens (two broken) (MNHN, Rudo von Cosel ColL, 21-XII-1978)

(both deposited in MNHN).

Other material studied: ?Brava Island: Baia Pedrinha, 3 sp. (ER).

Type locality: Salamanza, Sao Vicente, 1 m depth.

Etymology: The species is dedicated to my daughter Elvira Moreno Martín 11 years oíd, for her

constan! effort, sensitivity and enthusiasm without limits.

Description: The shell is oval-cylin-

drical, quite elongate, with the body

whorl covering almost all the previous

spirals. There are 6-7 plaits in the col-

umella, decreasing in size adapically.

The outher lip goes from the sutural

area to the siphonal canal, is swollen

and has 15-16 well defined denticles on

the inner side. The surface is smooth

and brilliant and somewhat translucent,

so that the internal mantle can be seen

by transparency. The protoconch is pau-

cispiral like in all marginelliform gas-

tropods, and the adult shell has approxi-

mately 2.5 whorls. The aperture is

narrow and elongated, and it has in its

anterior part a wide siphonal canal. The

shell colour seems very constan!, with

four brown bands, plus one in a subsu¬

tural position. These four bands, more

or less equidistan!, are narrow but well

contrasted, except the second one (num-

bered from de suture) which is broader

than the others, and constitutes the

central band very characteristic of the

species.

The animal is active, with the foot

longer than the shell when it is com-

pletely extended. The foot is truncated

in the anterior part and rounded in its

rear end. The head is small, divided into

two parts with a pair of very short ten-

tacles. The eyes are near the base of the

tentacles. The siphon, placed on the

head, is short and wide. The pigmenta-

tion pattern is very typical, with intense

colours: white, black and orange (Figs.

16 and 18). The head is black, with two

yellow specks in the center and two or¬

ange ones on the snout. Behind each eye

there is an orange speck, as on the end

of every tentacle. The siphon has black

spots and the end is bordered with or¬

ange. The foot presents lateral and ra¬

dial black bands, which reach the pedal

border, whereas in the anterior and pos¬

terior parts they end before reaching the

rim. The margin of the foot has a thin

orange line, which is continued on the

propodium or anterior part, and inter-

mittent on the central part and end of

the metapodium. On the propodium

and the posterior metapodium numer-

ous orange dots exist. The internal man¬

tle, seen by transparency, is black in

colour with large white, irregular, but

very definite patches. The only animal

observed was the holotype.

The radula has approximately 150

tooth lines (data from the only animal

78

Moreno: The genus Gibberula in the Cape Verde Islands; description of a new species

studied, the holotype). The teeth are

0.036 mm width (Fig. 13). As the shell

length of the specimen dissected for

radular study is 4.8 mm, the "shell

length/tooth width" ratio (L/Wr) is 133.

The teeth are arched and have a more

prominent central cusp with 4-5 minor

denticles on each side (10-11 cusps in

total). The central cusp has a broad base

and is blunt. The adjacent cusps to the

central one (1 or 2 in number) are

smaller that the other lateral denticles.

Every 3 teeth, approximately, the same

pattern of denticulation is observed.

There are never two identical contigu-

ous teeth and the lateral cusps of equal

size do not coincide with the same line.

Habitat: This species lives in the

upper sublittoral area, between 1 and 2

m depth, on a rocky bottom with algae.

Distribution: Gibberula elvirae n. sp. is

an endemic species of the Cape Verde

Islands. We have found this species with

certainty only on Sao Vicente Island,

where seems to be limited to the North-

East coast. A lot of 3 specimens from the

island of Brava (Baia Pedrinha) proba-

bly belongs to this species, considering

the presence of 4 bands on the shell as

described previously. The species has

not been found on the eastern islands,

such as Sal and Boavista, in spite of the

numerous samples obtained from both.

Discussion: Gibberula elvirae n. sp. is

different from any other species known

in this genus, both for the coloration

and shape of the shell, and for the pig-

mentation pattern of the animal, inclu-

ding the other species with similar

length shell that live in the Cape Verde

Islands: G. lucia and G. rolani. The

central expanded band of the shell

recalls certain patterns of G. oryza, but

the latter does not possess the thin basal

bands, next to the siphonal canal and to

the suture, respectively. As stated by

Gofas (1990), when the shells of G. oryza

have a pattern with spiral pigmentation,

they can show either a broad and central

band like the neotype, or two broad and

parallel bands in a central position (his

figures 2 and 4, respectively). Flowever,

the shell of G. elvirae n. sp. has up to 4

bands, and only the second (from the

suture) is broader than the others. The

length and general shape of the shell is

also different; G. elvirae is smaller (less

than 5 mm) and of cylindrical shape,

while G. oryza is larger (up to 8 mm) and

piriform.

The animal of G. elvirae n. sp. was

illustrated by Rolán (2005, Eig. 776) as

Gibberula sp. 2 (actually that specimen is

the holotype designated here and that

photograph the same image reproduced

here in Eigure 18). The colour pattern of

the animal of G. elvirae is different from

that of G. oryza shown by Gofas (1990:

Pl. 1, Eigs. a-a') and in the present paper

(Eig. 17), in having: 1) several black

patches on the foot, on the head and on

the siphon, 2) the orange dots of the

propodium arranged forming a well

defined line, and 3) the tentacles with a

broad orange band.

Like for the previous species, the

"V"-shape of the teeth and the number

of cusps are very similar between G.

elvirae n. sp. and G. oryza (10-11 and 11

cusps, respectively). Also, the tooth

width is similar between both species

(0.036 and 0.038 mm, respectively), but

in the first species the "shell

length/tooth width" ratio (L/Wr) is 133

and in G. oryza is 187, because the shell

in the last species is larger. Gofas (1990)

did not State the shell length of the dis¬

sected specimen used to obtain the

figured radula, and here it is assumed to

be of 7 mm, an average of the specimens

mentioned by the same author from the

Strait of Gibraltar.

Other species described from the Cape Verde Islands

Gibberula jousseaumi Rochebrune,

1881, described with type locality in

Porto Praia, Santiago Island, was cited

later as Marginella jousseaumi by Burnay

AND Monteiro (1977), as Gibberula

jousseaumi by Cosel (1982a), and as Per-

sicula jousseaumi by Guerreiro and

Reiner (2000), all of them with material

79

Iberus, 30 (1), 2012

from Sal Island. However, this ñame

was considered as a subjective synonym

of Volvarina sauliae (Sowerby, 1846) by

Moreno and Burnay (1999) in their

revisión of the genus Volvarina in the

Cape Verde Islands. Volvarina sauliae has

a shell that resembles a Gibberula, but

has a typical radula of the genus Volva¬

rina, with rectangular and "comblike"

shaped teeth, flat and multicusped (type

6 following CoovER, 1989), while the

radula of genus Gibberula is of type 3

(according to the same author), with

teeth strongly concave in the basal edge,

"V"-shaped and relatively few cusps.

The taxon Gibberula rachmaninovi

Kellner, 2003, was introduced recently in

a very poor description based only on a

shell from Sal Island, without animal

DISCUSSION

The genus Gibberula is represented in

the Cape Verde Archipelago by four

species, three previously described {Gib¬

berula lucia Jousseaume, 1877; Gibberula

rauli Fernandes, 1987; and Gibberula

rolani Cossignani and Cecalupo, 2005),

and the new species described here Gib¬

berula elvirae n. sp.

Gibberula jousseaumi Rochebrune,

1881, and Gibberula rachmaninovi Kellner,

2003, are considered here synonyms of

Volvarina sauliae (Sowerby, 1846). The

ñames of the Mediterranean species Gib¬

berula miliaria (Linnaeus, 1758) and G.

philippii (Monterosato, 1878) are erro-

neously included in the Cape Verde

Islands because these species are not

presen! in the archipelago.

Among the Gibberula species that

live in Cape Verde Islands, there are two

clearly differentiated groups, which are

equivalen! to those already known from

the Mediterranean, where Gofas (1990)

speaks about the "group of G. oryza”

and the "group of G. philippii'' , based on

shell length, shell transparency, pigmen-

tation patterns of the animal and most

frequent colours. In Cape Verde waters,

one group is constituted by "large"

species (for the dimensions of the

genus), of 3.5-5 mm long (as G. lucia, G.

and radula data. Rolán (2005) in his

book on the mollusks from the Cape

Verde Islands considers as a valid

species Volvarina sauliae (p. 146, Figs 635-

636, from Fuma, Brava Island, which is

the same specimen shown by Moreno

AND Burnay, 1999, Fig. 37, actually Corn¬

ing from Baia Pedrinha in the same is¬

land), and also Gibberula rachmaninovi (p.

148, Figs. 1193-1194, holotype from Santa

Maria, Sal Island). Moreno and Burnay

(1999: p. 113 and Fig 35) designated and

figured a neotype of Volvarina sauliae to

stabilize the nomenclature of this

species. The holotype of G. rachmaninovi

matches the neotype of V. sauliae in all

respects, and therefore the first ñame is

considered here a synonym of Volvarina

sauliae (Sowerby, 1846).

rolani and G. elvirae n. sp.), whereas

another group is represented by only

one "small" species, of approximately 2

mm long (G. rauli). The number of Gib¬

berula species (four, in the presen!

paper) presen! in the Cape Verde Archi¬

pelago is smaller than that of Volvarina

species (nine, in Moreno and Burnay,

1999), a genus of the family Marginelli-

dae. It is curious that the opposite

happens in the Mediterranean, where

there are more species of Gibberula (also

nine, according to Gofas, 1990), than of

Volvarina (only one, according to Gofas,

1989a). So, in the North Eastern Atlantic

Ocean the diversity of Volvarina species

is highest in warm waters, and specially

in the Cape Verde Islands, while the

diversity of Gibberula species is highest

along températe coasts, principally in

the Mediterranean Sea. Also along the

Angolan coast, in the southeast Atlantic

Ocean, there is a higher diversity of

species for the genus Gibberula, with at

least 11 described species (Gofas,

1989b).

The radular teeth, their size, shape

and number of cusps, are also importan!

to discriminate species in the genus Gib¬

berula. Among the studied species the

smallest radula is that of G. rauli, with

80

Moreno: The genus Gibberula in the Cape Verde Islands; description of a new species

only 0.017 mm tooth width. However,

the "shell length/tooth width" ratio

(L/Wr) is 118, exactly the lowest valúes

cited for the genus by Coovert and

CoovERT (1995). This means that the

radula is the largest in comparison to

the Shell among the Gibberula species

known. On the contrary the radula of G.

rolani, although larger (average of 0.026

mm tooth width), is proportionally

smaller in relation to the animal, with a

"shell length/tooth width" ratio (L/Wr)

of 192. The largest radula of the studied

species is that of G. elvirae n. sp. (0.036

mm tooth width), almost the same size

as that of G. oryza. Nevertheless, the

height of the shell of G. elvirae n. sp. (up

to 5 mm) is clearly smaller than that of

G. oryza (up to 8 mm), and therefore, the

"shell length/tooth width" ratio (L/Wr)

in this species is smaller (133) that that

of the other species mentioned: G. rolani

and G. oryza (with L/Wr ratio of 192 and

187, respectively).

All species studied here have a

restricted distribution area and are

endemic to the Cape Verde Islands,

except G. lucia Jousseaume, 1877, cited

also from Dakar, Senegal (Knudsen,

1956; Pin and Boyer, 1995), the closest

point of the African continent to the

Cape Verde Islands. There are relatively

few data about the distribution of the

species within the Archipelago, in part

due to the difficulty in determining the

species in the absence of information on

the animal colour pattern and radular

data. Also, the majority of previous pub-

lications that included Gibberula species

from Cape Verde Islands cited ñames of

Mediterranean species (G. miliaria and

G. philippii), or from the continental

coasts (as G. oryza) and therefore these

records are difficult to assign to the

species now considered valid in the

Archipelago.

The species that has a wider distribu¬

tion is G. rauli, that lives on Sal (type

locality), Boavista, Maio, Sao Vicente,

Santiago and Brava. Accordingo to

Rolán (1991, on the genus Conus), and

Moreno and Burnay (1999, on the

genus Volvarina), there are several natural

barriers between islands in the Cape

Verde Archipelago. The main barrier

divides the Archipelago into two groups,

that of the South East (including Sal,

Boavista, Maio, Santiago, Fogo and

Brava) and that of the North West

(including Sao Nicolau, Santa Luzia, Sao

Vicente and Santo Antáo). In the first

group there are also important barriers

between Sal and Boavista-Maio, and

between Santiago-Fogo-Brava and Maio-

Boavista. So, G. rauli is present in all the

groups and subgroups of islands in the

Archipelago, and probably lives on all

the islands. Gibberula rolani, described

from Sal is here confirmed from Boavista,

where it is common. A single specimen

from Brava studied here, considered with

some doubts as belonging to this species,

might indicate its presence in the South¬

ern group of islands. The oíd material

(from publications before 1987) that cited

the Mediterranean species G. miliaria

could correspond to G. rolani, with

doubts because they might correspond

also to other species. These distribution

data inelude also Sao Vicente of the

North group of islands. If the presence of

G. rolani was confirmed in Brava and Sao

Vicente, the distribution of the species

would inelude all groups and important

sub-groups of islands. Nevertheless, for

the moment, with confirmed informa¬

tion, its distribution is restricted to Sal

and Boavista, the easternmost islands

which are closest to the African coast.

Gibberula elvirae n. sp. is described here

only from Sao Vicente specimens which

is the type locality. There is a small lot of

3 specimens from Brava (FR) that have a

very palé coloured shell, but with a

pattern similar to that of the type mater¬

ial of G. elvirae n. sp. For the moment, this

new species could be considered as

endemic to Sao Vicente, with reserva-

tions, pending verification of the small

lot of 3 specimens from Brava or its pres¬

ence on other islands. Finally, G. lucia, the

first species described from Cape Verde

Islands, is the only one that we can con-

sider to be not endemic, being found also

in Senegal. There were no data of this

species in the Cape Verde Islands since its

description and all along the XIX and XX

centuries, whereas on the contrary, there

81

Iberus, 30 (1), 2012

was good material from Senegal. In the

present work a single specimen collected

at Santiago (AO), confirms the presence

of the species in the Cape Verde Islands,

if there was any doubt about the correct

origin of the material studied by

Jousseaume.

ACKNOWLEDGEMENTS

The author is grateful to the following

persons: Emilio Rolán, for his continuous

support with the loan of his collection,

personal observations and especially for

his friendship; Luis Pisani Burnay for his

collaboration and help in the study of the

marginelliform gastropods from the

Cape Verde Islands; Angel A. Luque

(UAM), for the loan of the PEICV collec¬

tion; José Templado (MNCN), for his con¬

tinuous help; all members of the PEICV

for their collaboration during the expedi-

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and other Mesogastropoda and Neogas-

tropoda. Zoologische Verhandelingen, 214: 1-

188, 22 PL

Burnay L.P. and Monteiro A. A. 1977.

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pp.

CoovERT G.A. 1987. A literature review and

summary of marginellid external anatomy.

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Revisión of the Supraspecific Classification

of Marginelliform Gastropods. The Nau-

tilus, 109 (2-3): 43-110.

CosEL R. VON 1982a. Ergebnisse deutsch-

-portugiesischer Sammelreisen auf den

Kapverdische Inseln (República de Cape

Verde Is.). Vorláufige Liste der marinen Mol-

lusken. Courier Forschungsinstitut Sencken-

berg, 52: 15-25.

CoSEL R. VON 1982b. Marine Mollusken von

Santa Luzia, Branco und Razo (Kapverdis¬

che Inseln). Courier Forschungsinstitut

Senckenberg, 52: 27-33.

CoSEL R. VON 1982c. Marine Mollusken der

Kapverdischen Inseln. Übersicht mit zoo-

geographischen Anmerkungen. Courier

Forschungsinstitut Senckenberg, 52: 35-76.

tion (Emilio Rolán and Luis Pisani Bur¬

nay -both organizers-, Angel A. Luque

from UAM, José Templado from MNCN,

Jesús Ortea from Oviedo University,

Manuel Ballesteros from Barcelona Uni¬

versity, Emilio Rolán-Álvarez from Santi¬

ago de Compostela University, Eva María

Llera from Oviedo University, Mirucha

Garrido, Ana Ballesteros, Ana Burnay,

Eduardo Espín, Andrés Lorenzo and

Diego Sarmiento); José Bedoya (MNCN),

who made the scanning photographs;

Serge Gofas (MNHN), for his invaluable

comments and information given on sev-

eral occasions; Philippe Bouchet, Serge

Gofas and Virginio Héros (MNHN) for

sending types and making available the

MNHN collection (including that of

Rudo von Cosel); Alvaro David De

Oliveira Ramos (Gulpilhares, Portugal)

who obtained and sent recent and impor-

tant information (including a photo-

graph) of Gibberula lucia.

CossiGNANi T. AND Cecalupo A. 2005. Des-

crizione di una nuova marginella (Gastro-

poda: Prosobranchia, Cystiscidae) delLAr-

cipelago di Capo Verde. Malacologia Mostra

Mondiale, 49 (nov.): 6-7.

Dautzenberg P. 1910. Contribution a la faune

malacologique de l'Afrique occidentale. Ac-

tes de la Société Linnénne de Bourdeaux, 44: 1-

174, 4 pL

Fernandes F. 1987. Descrizione di tre nuove

specie di Marginellidae (Mollusca: Gastro-

poda) delle Isole Capo Verde. Argonauta, 3

(3-4): 259-267.

García-Talavera F. and Bacallado J.J. 1979.

Nuevas aportaciones a la fauna de Gasteró¬

podos marinos (Mollusca, Gastropoda) de

las islas de Cabo Verde. Boletín del Insituto Es¬

pañol de Oceanografía, 6 (328): 202-208.

Gofas S. 1989a. Le genre Volvarina (Margine¬

llidae) dans la Méditerranée et FAtlantique

du Nord Est. Bollettino Malacologico, 25 (5-8):

159-182.

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marinhos da ilha de S. Vicente (Arquipélago de

Cabo Verde). Camara Municipal de Oeiras,

Europress, Póvoa de Santo Adriáo, 279 pp.

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Mollusques nouveaux. Bulletin de la Société

Zoologique de France, 1: 265-273, pl. 5.

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pical West Africa (Stenoglossa). Atlantide Re-

port, 4: 7-110, 4 pl.

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molluscs obtained by the Scotish Antarctic

Exp.-Species from St. Vicent, Cabo Verde Isl.

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48 (2): 18, 340-342.

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Volvarina (Gastropoda: Marginellidae) in the

Cape Verde Islands. Journal ofConchology, 36

(5): 83-124.

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faune de FArchipel du Cap Vert. Nouvelles Ar¬

chives du Muséum d'Histoire Naturelle, Paris,

2(4): 215-340.

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tropoda) en el Archipiélago de Cabo Verde (África

Occidental). Universidad de Santiago de Com-

postela. Tesis Doctoral, 653 pp.

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Verde archipelago. Part 1. Polyplacophora and

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83

k.

© Sociedad Española de Malacologia

Iberas, 30 (1); 85-89. 2012

Alvania garrafensis Peñas & Rolán 2008 (Gastropoda:

Rossoidae) from Croatian waters

Alvania garrafensis Peñas & Rolán 2008 (Gastropoda: Rossoidae) en

aguas croatas

Enzo CAMPANI*, Stefano BARTOLINI** and María Teresa SPANU***

Recibido el 3-VII1-201 1. Aceptado el 2-XII-201 1

ABSTRACT

We report on the presence of numerous specimens of Alvania garrafensis Peñas & Rolán

2008 from shell grit dived in shallow waters near severa! islands of the Quarnero Gulf,

North Adriatic Sea, Croatia. The species was up to now known oniy from the Garraf iit-

toral, Barcelona, East Spain and the present records greatly extend its geographical distri-

bution. Hypotheses that could explain such a large range extensión are considerad.

RESUMEN

Se documenta el registro de numerosos ejemplares de Alvania garrafensis Peñas y Rolán

2008 en cascajos conchíferos recolectados por buceo en aguas someras de varias islas

del golfo Quarnero, norte del mar Adriático, Croacia. La especie se conocía hasta la

fecha sólo en el litoral del Garraf, Barcelona, este de España, así que la nueva cita

amplía considerablemente su distribución geográfica. Se adelantan hipótesis que podrían

explicar una ampliación tan considerable.

INTRODUCTION

Alvania garrafensis Peñas & Rolán

2008 was described on a few specimens

trawled at 90 m depth off Cubelles, west

end of Garraf, after which the species

was named. Its known distribution was

hitherto limited to the type locality only

A localized distribution has to be

expected since the species has a shell pro-

toconch of 1,5 whorls, suggesting its non

- planktonic development and henee not

a wide geographical range (Scheltema

1978; 1986). In this respect our records

from Croatia seem quite strange and

surely need to be at least pointed out and,

if possible, understood.

MATERIALS AND METHODS

Most of the A. garrafensis specimens

we examined were found dead in shell

grit obtained through diving during late

Summer 2008 at various stations near

the islands of Krk and Prvic, Quarnero

Gulf, Croatia, at 35/50 m depth. The

material was picked from sandy or

muddy bottoms below the coralliferous

slopes which in a few steps connect the

rocky coast to the bottom. A quantity of

~ 15 litres of shell grit was collected

altogether. From this material ~ 340

shells of A. garrafensis were extracted;

* Corso G. Mazzini 299, 57126 Livorno, Italy, E-mail enzo.campani@fastwebnet.it, corresponding author.

** Via Ermete Zacconi 16. 50137 Firenze, Italy, E-mail: stefmaria.bartolini@alice.it

***Via Vivaldi traversa, 8 - 07041 Alghero, Italia. E-mail: mariate.spanu@alice.it

85

Iberus, 30 (1), 201’2

this species was the commonest Alvania

in the sorted material.

This large amount of shells were dis^

tributed as follows: Krk, Sokol Point

45m 120 shells; Krk, loe. Baska 38m 18

sh.; Prvic, Kita Bay 45m ~ 100 sh.; Prvic,

Gladni Potok Bay 50m 105 sh.

We could also examine two other

samples, courtesy of Peter Sossi, Trieste:

Prvic, 21m 6 sh; Plavnik, 40m 4 sh.

TAXONOMY

COENOGASTROPODA Cox, 1960

Rissoidae Cray, 1847

Alvania Risso 1826

Alvania garrafensis Peñas & Rolán 2008

Figures 1-6 show all the features

needed to compare our specimens with

A. garrafensis. Beyond the general shape,

we point out the 1.5 whorls, ~ 400 pm

wide shell protoconch with its very

characteristic spiral sculpture of 12 more

or less fragmentad threads; a body

whorl a bit less than 70% of the total

height; an axial sculpture of orthocline

cords interrupted at the base margin; a

weak spiral sculpture of 7 cords on the

body whorl, 4 between the two sutures

and 3 on the base. All these features fit

nicely with the holotype original

description and images. This however

has only three whorls at 2,8 mm height,

compared with four whorls at 3,5-4 mm

height of our Croaban specimens. We

thus conclude that the Spanish original

material contained only shells that were

slightly immature. In our Opinión this

fact can accommodate a difference we

observed between the two: while our

full grown specimens show lyrae all

over the lip internal margin, only 2-3 of

these are observed in the holotype shell

near the lowest part of the lip. We con-

sider that a minor difference and we are

confident that our Croatian specimens

have to be assigned to A. garrafensis. We

moreover sent a few of our specimens to

Dr. Emilio Rolan, who produced the

SEM images of figs. 1-3 and endorsad

our assignment. A hint to this possible

assignment was also present in a discus-

sion topic on the "Natura Mediterrá¬

neo" Italian online discussion forum

<www.naturamediterraneo.com>,

accessed November 2011.

We nevertheless compared our spec¬

imens with the two species Alvania

consociella Monterosato 1884 and Alvania

disparilis Monterosato 1890. As for the

first we based our observations on per¬

sonal material from our collections and

on images from the literatura (Gian-

NUZZI Savelli, Pusateri, Palmeri &

Ebreo, 1997; Scaperrotta, Bartolini &

Bogi, 2011). For the second we obtained

a photo of a specimen 3,6 mm high,

from the Monterosato collection in

Rome Zoological Museum (ZMR), cour¬

tesy of A. Pierullo. This photo enabled a

comparison of this species with our

Croatian specimens.

Alvania consociella has deep and canalic-

ulated sutures, almost fíat whorls, more

numerous spiral chords which form

nodules Crossing the axial ones, giving a

nodulose look to the shell shape; it often

has few spirals, which are white on a darker

background; its protoconch shows a papil-

lae sculpture that is quite different from

that of A. garrafensis. The photo of A. dis-

parilis we saw shows a shell shape more

squat than A. garrafensis (height to width

ratio ~ 1,6 versus 1,9 in Croatian speci¬

mens), with rounded whorls and a

rounded aperture; the sutures are deep

and canaliculated (unlike in garrafensis);

the thin orthocline axial chords are much

narrower than their interspaces (wider in

garrafensis) and more numerous; the spiral

chords are more numerous than in gar¬

rafensis, as thin as the axial ones, and the

axial to spiral Crossing forms a regular

squared reticule (totally absent in A. gar¬

rafensis); finally a few (3-4) thin, but well

86

Campani ET AL: Álvania garraf ensis (Gastropoda: Rossoidae) from Croatian waters

Figures 1-3. Alvania garrafensis SEM images. 1: adult shell 4 mm high, Krk Island, 45 m; 2: proto-

conch of the same specimen, side view; 3: protoconch of the same specimen, apical view.

Figuras 1-3. Alvania garrafensis imágenes de MEB. 1: concha adulta de 4 mm de alto, isla de Krk, 45

m; 2: protoconcha del mismo ejemplar, vista lateral; 3: protoconcha del mismo ejemplar, vista apical.

raised and marked smooth spiral chords

are evident on the base. All these diffe-

DISCUSSION

Alvania garrafensis is apparently

much commoner and widespread in

Croatian waters then in the type locality.

rences are clearly seen comparing Figure

7 to Figures 1-6.

It was found in large numbers and in

many places in Croatia, always in the

depth range 25-50 m, while in the case

87

Iberus, 30 (1), 2012

Figures 4-6. Alvania garraf ensis. 4: specimen from Prvic, 50 m, 4 mm high; 5, 6: different views of

the protoconch of the same specimen, greatly enlarged.

Figuras 4-6. Alvania garrafensis. 4: ejemplar de Prvic, 50 m, 4 mm de alto; 5, 6: diferentes vistas de la

protoconcha del mismo ejemplar, a mayor aumento.

of Garraf only a few specimens coming

from a single and much deeper locality

(90 m.) were found.

Moreover to our knowledge the

species has never been recorded in any

locality intermediate between Garraf and

Croatia, e.g. French or Italian coasts,

although it may well have been over-

looked due to its quite recent description.

A possible explanation would be to

regard (Rolan, prívate communication)

A. garrafensis as a native species in Croa-

tian waters, arrived to Garraf via some

"man assisted" transportaron mecha-

nism, for instance with the trade of

Ostrea edulis from the North Adriatic Sea

to the Iberian Península. A different

Opinión (from A. Peñas, prívate commu¬

nication) is that maybe A. garrafensis is

widely present in the West Mediter-

ranean, but yet unsampled in most areas

in deep waters; moreover to his know¬

ledge there is no Ostrea edulis import in

the type location area. We think that the

hypothesis of a Croatian origin for the

species arrived in Garraf via some "man

assisted" transportation mechanism

seems very unlikely and that the

hypothesis of a hitherto overlooked

species is to be preferred.

We conclude that the problem of the

geographical distribution of A. garrafen-

88

Campani ET AL: Alvania garraf ensis (Gastropoda: Rossoidae) from Croatian waters

Figure 7. Alvania disparilis, Palermo, 3.6 mm, specimen in Monterosato collecdon, Zoological

Museum of Rome.

Figura 7. Alvania disparilis, Palermo, 3,6 mm, ejemplar en la colección Monterosato, Museo Zoológico

de Roma.

sis is unclear as yet and its full under-

standing needs some further research in

regions intermedíate between the Garraf

area and the Adriatic Sea, where it

seems well established.

ACKNOWLEDGEMENTS

This paper wouldn't have been pos-

sible without the generous help of Dr.

Emilio Rolan, Vigo, Spain, who pre¬

pared the SEM images of our shells,

confirmed our species assignment,

BIBLIOGRAPHY

Giannuzzi Savelli R., Pusateri F., Palmeri A.

& Ebreo C.”^ 1997. Atlante delle Conchiglie ma¬

rine del Mediterráneo Vol II. Edizioni "La Con-

chiglia" Roma. 258 pp.

Monterosato T.A. di 1890. Conchiglie della

profonditá del mare di Palermo. Naturalista

Siciliano, 9 (6): 140-151; 9 (7): 157-166; 9 (8):

181-191.

Peñas A., Rolán E. & Ballesteros M. 2008. Se¬

gunda adición a la fauna malacológica del li¬

toral de Garraf (NE de la Península Ibérica.

Iberas, 26 {2):15-42.

shared with us his opinions on the

species geographical distribution and.

revised our paper We thank him a lot

for that. We thank also Peter Sossi,

Trieste, for letting us examine some of

his specimens, Mrs. Angela Pierullo,

Rome, for providing us with a photo of

an A. disparilis type specimen and Dr.

Massimo Appolloni, responsible for the

ZMR malacological collection for per-

mission to publish this image. We

finally thank an anonymous referee who

suggested a few relevant changos to the

paper.

SCAPERROTTA M., BaRTOLINI S. & BOGI C. 2011.

Accrescimenti Vol III. L'Informatore Piceno,

Ancona. 184 pp.

ScHELTEMA R.S. 1978. On the relationship bet¬

ween dispersal of pelagic veliger larvae and

the evolution of marine proosobranch gas-

tropods. In Battaglia B., Beardmore J. A. (Eds.);

Marine organisms: genetics, ecology and evolu¬

tion. Plenum, New York, p. 303-322.

ScHELTEMA R.S. 1986. On dispersal and plank-

tonic larvae of benthic invertebrates: an eclec-

tic OverView and summary of problems. Bu-

lletin of Marine Science, 39: 290-322.

89

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© Sociedad Española de Malacología

Iberas, 30 (1): 91-102, 2012

A review of species-limits in some Cryptazeca (Gastropoda:

Azecidae)

Reconsideración de los límites entre especies en Cryptazeca

(Gastropoda: Azecidae)

David T. HOLYOAK and Geraldine A. HOLYOAK*

Recibido el ll-Xl-2011. Aceptado el 3-XII-20 11

ABSTRACT

Species-limits are reviewed for severa! taxa in the land-snail genus Cryptazeca, based

mainly on measurements of shell size and shape. Compared to previous studies, larger

samples of shells from more numerous populations were available for analysis. Widely

overlapping ranges of variation in shell size and form in C. monodonta (from SW. France)

and C. kobeíti (NW. Spain) imply that these allopatric taxa are likely to be conspecific, so

the latter ñame is synonymised with the formen The species lives amongst leaf-litter in moist

deciduous woodland. C. subcylindrica (SW. France) has shells that are similar to those of

some populations of the allopatric and widely variable C. vasconica (NW. Spain), henee

the latter ñame is synonymised with the formen The species lives amongst debris in crevi-

ces of limestone rocks, sometimes inside caves. An identificaron key to the species is pre-

sented, based on characters of adult shells. The proposed taxonomic changes have impli-

cations for the conservation status of the taxa involved, since their current threat status on

the lUCN Red List should probably be reduced now that each of the two species recogni-

sed as valid has a wider range and more populations known.

RESUMEN

Se realiza una reconsideración de las características que permiten separar especies dentro

del género de moluscos terrestres Cryptazeca, basados en la forma y medidas de la con¬

cha. En comparación con otros estudios previos, este análisis se ha efectuado sobre un mayor

número de ejemplares y procedentes de un mayor número de poblaciones. El amplio sola-

pamiento observado en lo concerniente al tamaño y forma de la concha en C. monodonta

(del SW, de Francia) y C. kobelti (NW. España) implica que estos dos taxones alopátricos

son muy probablemente conespecíficos, por lo que este último nombre es incluido en la sino¬

nimia del primero. Esta especie vive entre la hojarasca en suelos húmedos de bosques cadu-

cifolios. Por otro lado, las conchas de C. subcylindrica (SW. Francia) son similares a las de

varias poblaciones de la muy variable y alopátrica C. vasconica (NW. España), por lo que

este último nombre se incluye en la sinonimia de C. subcylindrica. Esta especie vive entre

los restos vegetales acumulados en fisuras de rocas calizas, y a veces también dentro de

cuevas. Se presenta una clave para la identificación de las especies del género, basada en

los caracteres conquiológicos de individuos adultos. Los cambios taxonómicos propuestos

tienen implicaciones sobre el estatus de conservación de los taxones implicados, de forma

que la categoría de amenaza con que figuran en la lista roja de la UlCN deberá proba¬

blemente ser menor, ya que ambas especies reconocidas como válidas tienen ahora un mayor

rango de distribución y se conocen un mayor número de poblaciones.

* Quinta da Cachopa, Barcoila, 6100-014 Cabezudo, Portugal; holyoak9187@hotmail.com

91

Iberus, 30 (1), 2012

INTRODUCTION

The genus Cryptazeca De Folin & Bé-

rillon, 1877 comprises small land-snails

occurring in woodland, among limes-

tone rocks and in cave habitáis in SW.

France and parts of NW. Spain. Difie¬

ren! authors have regarded them as

members of the Ferussaciidae in the su-

perfamily Achatinoidea of the infraor-

der Sigmurethra (Schileyko, 1976, 1999;

Bank, Bouchet, Falkner, Gittenber-

GER, FÍAUSDORE, VON PROSCHWITZ AND

Ripken, 2001; Falkner, Bank and von

Proschwitz, 2001; Falkner, Ripken

AND Falkner, 2002) or Cochlicopidae

within the infraorder Orthurethra

(ZiLCH, 1959; Gittenberger, 1983; Gó¬

mez AND Angulo, 1987, 1990; Gómez,

1990b, 1991). The uncertainty about

their wider relationships has been resol¬

ved recently by Madeira, Elejalde,

Chueca and Gómez (2010) from a phy-

logenetic study of rRNA sequence data.

They demónstrate a sister-group rela-

tionship of Cryptazeca and Hypnophila,

both of which are closely allied to Azeca

(Azecidae), but placed more distan!

from CochUcopa and far from Ferussacia.

The review by Gittenberger (1983)

recognised four species of Cryptazeca,

with two restricted to SW. France in

Dept. Pyrénées-Atlantiques (C. rnono-

donta (De Folin & Bérillon, 1877), C.

suhcylindrica De Folin & Bérillon, 1877)

and two restricted to parts of NW. Spain

from Prov. Asturias eastwards to Prov.

Navarra (C. vasconica (Kobelt, 1894), C.

kobelti Gittenberger, 1983). Since then

two more species have been described

from caves in NW. Spain (C. spelaea Gó¬

mez, 1990a; C. elongata Gómez, 1990b). A

total of six species has therefore been re¬

cognised in recent literature (e.g. Gó¬

mez, 1990b; Fauna Europaea DataBase

Project, 2011). Detailed studies of the

genital anatomy of C. monodonta and C.

vasconica (Gómez and Angulo, 1987;

Gómez, 1991) revealed that they differ

significantly in penis structure. Both of

the cavernicolous species C. spelaea and

C. elongata also show peculiar fea tures in

their genital anatomy (Gómez, 1990a,

1990b), as well as much larger shells

than those of all congeners from open

habitats. The existence of distinct spe¬

cies is also supported by finds of two

pairs of taxa living together at the same

localities, C. monodonta with C. suhcylin¬

drica in SW. Trance (Gittenberger, 1983;

at Grotte de Sare: FIolyoak and Sed-

don, 1985) and C. kobelti with C. vasco-

nica in Spain (S. of Orduña: Gittenber¬

ger, 1983; Dance, Holyoak, Seddon

AND Tattersfield, 1986).

When Gittenberger (1983) wrote his

seminal review, most of the specimens

available to him were obtained in the

nineteenth century Efforts to refind three

of the species were unsuccessful at that

time, so his more recent material consis-

ted only of eight shells of C. vasconica.

However, all of the species were refound

in the following two decades, some of

them at several localities (Holyoak &

Seddon, 1985; Dance et al., 1986;

Gómez and Angulo, 1987; Altonaga,

Gómez, Martín, Prieto, Puente and

Rallo, 1994). The present authors' field-

work from 2001-2011 has added more

specimens and additional localities, but

also disclosed populations of Cryptazeca

that could not be identified to species

using the key to shells presented by

Gómez (1990b). Our initial suspicions

that undescribed species are involved

were set aside when fuller studies

showed greater variability in shell size

and form in the known species than is

recognised in existing literature. In parti¬

cular, the allopatric C. monodonta and C.

kobelti appear to be linked by many inter¬

medíate shells and the very variable C.

vasconica has populations linking it to the

allopatric C. subcylindrica. This paper the¬

refore reconsiders species-limits in these

two species pairs, based on many more

specimens than were available for Git-

tenberger's (1983) review (cf. Table II).

MATERIAL AND METHODS

Field collections by the authors were

made by direct searching and by sieving

leaf-litter and other debris using nested

2.0 and 0.5 mm mesh sieves to obtain

"fines" that were subsequently searched

92

Holyoak .'VND HolyoaK: a review of species-limits in some Cryptazeca

a

b

Figure 1. Locations of shell measurements given inTables I and II. a-b, shell height (SH), c-d, shell

breadth (SB), b-e, height of body whorl (BH), b-f, height of aperture (AH).

Figura 1. Representación de las medidas de la concha recogidas en las Tablas I y 11. a-b, altura de la concha

(SH), c-d, anchura de la concha (SB), b-e, altura de la última vuelta (BH), b-f, altura de la abertura (AH).

at low magnifications using a stereo-

microscope. AIl shells found were retai-

ned, to avoid possible bias in favour of

large or well preserved specimens.

Localities and altitudes were recorded

from 2007 onwards using Garmin hand-

held GPS, accurate to within 10 metres.

From 2007 onwards sites were also

given consecutive serial numbers (e.g.

E153). Habitat notes (including bedrock

type and vegetation) and associated

Mollusca were recorded at all sites.

Adult snails could readily be distin-

guished from immatures by the thicke-

ned edge to the peristome. Drawings of

shells were prepared using a drawing

tube on a Meiji RZ series stereomicros-

cope. Figure 1 shows the locations of

measurements of shell height, shell

RESULTS

Measurements of shells are listed

in Table I and summarised and

compared with those from the litera-

breadth, body-whorl height and aperture

height. These measurements were made

on adult shells using Infinity Analyze©

software on images taken with an Infinity

1 camera on a Meiji RZ series stereomi-

croscope. The measurements on the

images were reproducidle to ± < 0.01

mm, but unavoidable slight tilting of the

shells almost certainly caused minor

additional loss of precisión.

Details of the material studied are

listed in the Appendix. Specimens

reported by Holyoak & Seddon (1985)

and Dance et al. (1986) were studied

on loan from the Dept. of BioSYB of

National Museum and Gallery of Wales,

Cardiff, U.K. (NMW.Z). Almost all other

specimens studied are retained in the

Collection of G.A. and D.T. Holyoak.

ture in Table IL Figure 2 gives dra¬

wings of representativo shells of each

taxon.

AZECIDAE Watson, 1920

Cryptazeca De Folin & Bérillon, 1877

Cryptazeca monodonta (De Folin & Bérillon, 1877)

Azeca monodonta De Folin & Bérillon, 1877, Bul!. Soc. Borda Dax, 2 (1); 199, pl. [1], figs 1, 2 [Type

locality: "Bramepan", St. Pierre-dTrube SE. of Bayonne, Pyrénées-Atlantiques, France]. See

Gittenberger & Kosten (1983) for bibliographical details.

93

Iberus, 30 (1), 2012

Table I. Measurements (mm) of shells of Cryptazeca. N = number of intact adult shells measured,

SH = shell height, SB = shell breadth, BH = body-whorl height, AH = aperture height; see Fig. 1

for locations of measurements.

Tabla I. Dimensiones (mm) de las conchas de Cryptazeca. N ^ número de conchas adultas intactas

medidas, SH = altura de la concha, SB = anchura de la concha, BH = altura de la última vuelta, AH

= altura de la abertura; ver Fig. 1 para la forma en que se han tomado las medidas.

94

Holyoak and HolyoaK: a review of species-limits in some Cryptazeca

Table IL Summary of measurements (mm) of shells of Cryptazeca species and comparisons with

data from literature. N = number of intact adult shells measured; SH = shell height; SB = shell

breadth; measurements (mm) from literature are cited in the form in which they were published;

measurements of the lectotype of C. subcylindrica are based on Gittenberger (1983: fig. 12).

Tabla II. Resumen de las medidas (mm) de las conchas de las especies de Cryptazeca j/ comparación con

los datos de la literatura. N - número de conchas adultas intactas medidas; SH = altura de la concha;

SB = anchura de la concha; las medidas (mm) de procedentes de la bibliografía son citadas en la forma

en que fueron publicadas; las medidas del lectotipo de C. subcylindrica se basan en Gittenberger

(1983: fig 12).

Cryptazeca kobelti Gittenberger, 1983, Zool. Meded., 57 (23): 313 iType locality: Conca d'Orduña,

Spain].

C. kobelti was described from just four

oíd shells from Conca de Orduña in Prov.

Navarra (Gittenberger, 1983) [the actual

localities were probably in an area that is

now within Prov. Alava] and compared

with ca 24 oíd shells of C. monodonta. The

present study used 72 shells of C. mono¬

donta (from 2 localities) and 28 of C. kobelti

(from 4 localities in 2 areas). Based on the

larger samples now available it is clear

that these taxa overlap widely in shell size

and form (Pables I, II; Fig. 2). The key by

Gómez (1990b) gave shell height/shell

breadth as 2.2-2.35 in C. monodonta and

2. 1-2.2 in C. kobelti, whereas our measu¬

rements give ratios respectively of 2.05-

2.33 and 2.07-2.28 so that most individual

shells are inseparable on this basis. In the

original description of C. kobelti, Gitten¬

berger (1983: 313) described C. monodonta

as "more spindle-shaped and less palé in

colouration (judging after only one fresh

specimen of C. kobelti and several of C.

monodonta)”. However, in our material the

shapes of individual shells of the two taxa

can be closely matched (cf. Fig. 2). There

is also no difference whatever in the

brown coloration of shells of the two taxa

when modern specimens collected alive

are compared. Gittenberger 's "fresh"

shells must anyway have been about 90

years oíd when he studied them.

Locality data reported by Altonaga

ET AL. (1994) for C. kobelti included sites in

the Spanish Provinces of Guipúzcoa and

Vizcaya situated between the type-loca-

lity near Orduña and the localities for C.

monodonta in Dept. Pyrénées-Atlantiques

95

Iberus, 30 (1), 2012

(Fig. 3A), Our few specimens from near

Madariaga (Prov. Guipúzcoa) could only

be identified arbitrarily as either C. kobelti

or C. monodonta. Henee, it seems clear

that C. kobelti should be regarded as a

synonym of C. monodonta. The species

lives amongst leaf-litter in moist deci-

duous woodland, including hillside

Fagiis sylvatica woods near Orduña (ele-

vation ca 540-570 m) and near Madariaga

(at elevations of 656 and 675 m), and

more varied woodland with Qiiercus,

Acer, Fraxinus excelsior, Corylus avellana

and other trees and shrubs at Grotte de

Sare (ele vation ca 220 m). Most if not all

the localities overlie iimestone bedrock

and there is much exposed rock at some

of them, whereas several others lack rock

exposures in the places where the species

was collected alive.

Cry-ptazeca suhcylindrica De Folin & Bérillon, 1877

Cryptazeca monodonta var. subcylindrica De Folin & Bérillon, 1877, Bidl. Soc. Borda Dax, 2 (4): 443,

pl. 3, fig. 6 [Type locality: "Bramepan", St. Pierre-dTrube SE. of Bayonne, Pyrénées-Atlanti-

ques, France]. See Gittenberger & Kosten (1983) for bibliographical details.

Ferussacia (Hypnophila?) vasconica Kobelt, 1894, Icón. Land- & Süsswass.-Moll, (N.F.) 7 (1-4): 37, pl.

188, fig. 1200 [Type locality: Orduña, Prov. Alava, Spain].

Recent collections of C. vasconica

show a wide range of variability in shell

size and shape (Figs. 2A-F, Table I). In dif-

ferent populations the shell height ranges

from 2.9-3.0 mm (N. of Araotz, Prov. Gui¬

púzcoa) to a mean of 4.4 mm (Desfilade¬

ro de los Beyos, Prov. Asturias) and shell

shape ranges from subcylindrical to na-

rrowly conical or narrowly fusiform (Fig.

2). Shell colour is typically light brown

(lighter than in C. monodonta), but colour-

less shells oceur inside caves (Gómez,

1990b: 373; Altonaga et al., 1994: 76).

Measurements of our material of C.

subcylindrica from near the Grotte de Sare

(Dept. Pyrénées-Atlantiques) are within

the overall range of shell size and shape

found in C. vasconica (Table I, Fig. 2). The

freshest empty shells (at least one collec¬

ted alive) from Grotte de Sare inelude

two that are palé brown and one that is

almost colourless (part of NMW.Z.

1993.052.10530). As pointed out by Git¬

tenberger (1983) and Gómez (1990b:

373), the transverse sculpture in C. subcy¬

lindrica is strong; on the freshest shells it

is closely similar to that of C. vasconica. A

single adult specimen from our eastern-

most Spanish locality for C. vasconica ne¬

ar the Puerto de Usateguieta (Prov. Nava¬

rra) is so cióse to the C. subcylindrica from

Grotte de Sare in shell size and form (cf.

Figs. 2F and 2G) that identifying them as

sepárate species seems arbitrary.

The lectotype of C. subcylindrica

figured by Gittenberger (1983: 314) is a

very narrow shell (shell height/ shell

breadth 3.08), whereas the measurements

he gives for other specimens widely

overlap those of our material from Grotte

de Sare. Although he reported the shell of

that species as colourless, this is not true

of some from Grotte de Sare, whereas

some C. vasconica are colourless, as noted

above. Henee most C. subcylindrica (inclu¬

ding our material from Grotte de Sare)

show shells that are within the range of

variation found in different populations

of C. vasconica, only the lectotype being

notably narrow (although the C. vasconica

shell from Puerto de Usateguieta appro-

aches this with shell height/ shell breadth

of 2.84). It therefore seems better to regard

these taxa as conspecific, with C. vasconica

treated as a synonym of C. subcylindrica

which has the older ñame. The alternative

of dividing C. vasconica into several closely

split species might eventually prove to be

justifiable, but there are too many Ínter-

medíate populations for this to be warran-

ted on the basis of the shell data alone. Alt¬

hough anatomical and molecular compa-

risons are desirable, we have failed to

obtain living C. subcylindrica on recent

visits to sites near its French type locality.

C. subcylindrica sensu lato is an elusive

snail that is usually found in small

numbers in more or less shaded places

96

Holyoak and HolyoaK: a review of species-limits in some Cryptazeca

Figure 2. Drawings of representative shells of Cryptazeca species. A: C. vasconicay by San Ignacio

turning, Prov. Asturias; B-D: C. vasconicay W. of Ranero, Prov. Vizcaya; E: C vasconicay NE. of

Araotz, Prov. Guipúzcoa; F: C vasconica. Puerto de Usateguieta, Prov. Navarra; G: C. subcylindricay

Grotte de Sare, Pyrénées-Atlantiques; H: C. monodontay Grotte de Sare, Pyrénées-Atlantiques; I: C

kobeltiy 3 km aiong road S. of Orduña, Prov. Alava; J: C. kobeltiy 4 km along road S. of Orduña, Prov.

Alava; K: C kobeltiy site El 49, E. of Madariaga, Prov. Guipúzcoa; L: C monodontay Grotte de Sare,

Pyrénées-Atlantiques; all specimens are in the Collection of GA. and D.T. Holyoak.

Figura 2. Dibujos de varias conchas representativas de las especies de Cryptazeca. A: C. vasconica, de San

Ignacio turning, Prov Asturias; B-D: C. vasconica, W. de Ranero, Prov. Vizcaya; E: C. vasconica NE.

de Araotz, Prov, Guipúzcoa; E: C. vasconica. Puerto de Usateguieta, Prov. Navarra; G: C. subcylindrica,

Grotte de Sare, Pyrénées-Atlantiques; H: C. monodonta, Grotte de Sare, Pyrénées-Atlantiques; I: C. kobelti,

3 km por la carretera al S. de Orduña, Prov. Alava; ]: C. kobelti, 4 km por la carretera al S. de Orduña,

Prov. Alava; K: C. kobelti, lugar E149, E. de Madariaga, Prov. Guipúzcoa; L: C. monodonta, Grotte

de Sare, Pyrénées-Atlantiques; todos los ejemplares están en la Colección de G.A. y D.T. Holyoak.

97

Iberus, 30 (1), 2012

Figure 3. Distribution of some Cryptazeca species mapped in 10 km squares of the U.T.M. grid. A:

C monodonta sensu lato, closed symbols= records originally determined as C monodonta, open

symbols= records originally determined as C kobelti, large symbols= authors’ records, small

symbols= records from literature; B: C. subcylindrica sensu lato, closed symbols= records originally

determined as C subcylindrica, open symbols= records originally determined as C. vasconica, large

symbols= authors’ records, small symbols= records from literature. The report of C. monodonta

from Prov. Asturias by Bech (1986) is not mapped because the species identification seems unli-

kely (the shell height was given as 3.15 mm, shell breadth [erroneously?] as 3.8 mm).

Figura 3. Distribución de varias especies de Cryptazeca en cuadrículas UTMs. de 10 km de lado. A: C.

monodonta sensu lato, círculos negros^ citas determinadas originalmente como C. monodonta, círcu¬

los blancos^ citas determinadas originalmente como C. kobelti, símbolos grandes^ citas de los autores,

símbolos pequeños- citas procedentes de la literatura; B: C. subcylindrica sensu lato, círculos negros-

citas determinadas originalmente como C. subcylindrica, círculos blancos^ citas determinadas original¬

mente como C. vasconica, símbolos grandes^ citas de los autores, símbolos pequeños^- citas procedentes

de la literatura. La cita de C. monodonta de la Prov. Asturias de Bech (1986) no ha sido representada

en el mapa ya que la identificación de la especie es dudosa (la altura de la concha indicada es de 3, 15

mm, su anchura [¿errónea?] es de 3,8 mm).

that have exposed limestone rocks, some-

times under closed cover of deciduous

woodland, sometimes on more open

crags or limestone pavement with only

scattered trees and shrubs. Litres of sie-

vings from ground litter commonly held

only a few shells and repeated visits to

the same locality were often unsuccessful

in finding more. Our only locality that

produced a large sample (32 snails or

shells, 12 of them living) was at the damp

bottom of a solution hollow in limestone

pavement, 5 m below the ground surface,

where permanently shaded by the

98

Holyoak and Holyoak: a review of species-limits in some Cryptazeca

surrounding rocks and by trees (W. of

Ranero, Prov. Vizcaya). The species is also

reported from inside caves (see above).

Overall, this species appears much more

closely restricted to sheltered and shaded

rock crevices than C. monodonta, with no

confirmed occurrences in woodland leaf-

litter at sites lacking exposed rocks.

DISCUSSION

Revised key to Cryptazeca species

Reduction of the number of species

recognised within Cryptazeca greatly sim-

2 " Shell height/shell breadth > 2.8

- Shell height/shell breadth < 2.8

plifies Identification of its species. The

key to adult shells presented by Gómez

(1990b: 373) can be revised as follows:

. 2

. 3

C. elongata

. C. spelaea

1 - Shell height >5.5 mm (cave species) .

- Shell height <5.5 mm (in open habitats or caves)

3 - Shell height/ shell breadth > 2.35, shell with strong transverse sculpture .

. . . C. subcylindrica

- Shell height/shell breadth < 2.35, shell with transverse sculpture weak or nil . . .

. C. monodonta

Conservation status of the enlarged

C. monodonta and C. subcylindrica

Three of the four taxa of Cryptazeca dis-

cussed above are included in the lUCN

Red List (Mollusc Specialist Group, 1996):

C. kobelti as Endangered (under criteria B1

+ 2c), C. monodonta and C. subcylindrica

both as Vulnerable (under criterion D2),

whereas C. vasconica is regarded as Lower

Risk/ near threatened. Two of the taxa are

now protected by law in France, as Mollus-

que proteges, C. monodonta under Article

4, C. subcylindrica as a Rare species under

Article 1. Our proposal to regard C. kobelti

as conspecific with C. monodonta adds three

French localities with modern records to

those in Spain, thus increasing the species'

range and presumably reducing its global

threat status to Vulnerable. Nevertheless,

it remains a localised endemic occurring

only in sites that are species-rich for land-

snails and which need to be protected (see

Appendix). The proposal to regard C. vas¬

conica as conspecific with C. subcylindrica

will greatly increase the species' range and

number of localities with modern records,

presumably implying that the overall

threat status should be Lower Risk/ near

threatened (i.e. no longer on the Red List).

ACKNOWLEDGEMENTS

Thanks are due to the Dept. of

BioSYB at National Museum and

Gallery of Wales, Cardiff, U.K. (NMW.Z)

for loans of specimens and Jennifer

Gallichan and Dr Graham Oliver for

helping with arrangements for the loan.

Thanks are also due to Jonathan Ablett

for assistance in studies at the Natural

History Museum, London, U.K. Figure 3

was prepared using the DMAP software

written by Dr A.J. Morton.

BIBLIOGRAPHY

Altonaga K., Gómez B., Martín R., Prieto

C.E., Puente A.I. and Rallo A. 1994. Estu¬

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100

Holyoak and Holyoak: a review of species-limits in some Cryptazeca

APPENDIX. MATERIAL STUDIED.

Data are Usted in sequence as province, locality ñame, habitat description, associa-

ted species, altitude, U.T.M. grid reference, date, collectors' initials, collector's field

number, number of shells or specimens (immature or adult, whether alive or dead

when collected), collection, registration number if any. Abbreviations: CGAH =

Collection of G.A. & DT. Holyoak, DTH = D.T Holyoak, GAH = G.A. Holyoak, MH =

M„ Holyoak, MBS = M.B. Seddon, NHMUK = Natural History Museum, London,

U.K.; NMW.Z. = Department of BioSYB, National Museum and Gallery of Wales,

Cardiff, U.K.; sh = shells, sp = spirit specimens.

C. kobelti SPAIN, Prov. Alava: by N625 ca 3 km S. of Orduña, steep limestone hill-

side with Fagiis woodland, damp beneath, with few saplings & bushes, "ca 610 m

alt.", VN3, 26 Aug. 1984, DTH, MH, MBS, 43 sh (adults & immatures, alive & dead),

NMW.Z. 1993.052. 10453; by A625 ca 3 km S. along road from Orduña, from leaf-litter

on N.-facing slope in beechwood, ca 540 m alt., 30T 049797/47574, 9 June 2007, GAH,

DTH, site 33, 6 sh (adults, alive), CGAH; by A625 ca 4 km S. along road from Orduña,

from leaf-litter on N.-facing slopes in beechwood, ca 570 m alt., 30T 04977/47557, 9

June 2007, GAH, DTH, site 34, 5 sh (2 immatures & 3 adults, alive & dead), CGAH;

Prov. Guipúzcoa: E. of Madariaga (NW. of Azkoitia), N.-facing slope with few limes¬

tone crags, under mature beech wood, 656 m alt., 30T 05560/47840, 8 May 2011, GAH,

DTH, site E148, 1 sh (adult, alive), CGAH; E. of Madariaga (NW. of Azkoitia), N.-

facing slope with few limestone crags & beech wood, 675 m alt., 30T 05551/47842, 9

May 2011, GAH, DTH, site E149, 6 sh (1 adult & 5 immatures, alive & dead), CGAH.

C. monodonta TRANCE, Dept. Pyrénées-Atlantiques: "Basses-Pyrénées", Ex. A.M.

Norman Coll. (Ex Morelet), 3 sh, NHMUK.1958.9.3.1-3; 1 km SW. of Ustaritz {ca 8 km

NW. of Cambo), mixed deciduous wood with Corylus understorey in valley of small

stream, 24 Aug. 1980, DTH, 23 sh (alive & dead), NMW.Z. 1993.052.04432; near Grotte

de Sare, 5.5 km S. of Sare, mixed deciduous wood on rocky limestone slope, moss-rich

ground vegetation, 23 Aug. 1980, DTH, 29 sh (alive & dead), NMW.Z. 1993.052.04408;

Grotte de Sare, rocky limestone slope, shaded by mixed deciduous trees, very mossy

with dead wood lying, ca 300 m alt., XNl, 6 Aug. 1983, DTH, MBS, 6 sh,

NMW.Z. 1993.052.07652; Grotte de Sare, rocky limestone slopes shaded by mixed deci¬

duous wood, moist & mossy, XNl, 28 Aug. 1984, DTH, MH, MBS, 56 sh (dead),

NMW.Z. 1993.052.10528; near Grotte de Sare, limestone slopes near stream in deci¬

duous woods, ca 200 m alt., 43°16' N., 1°34'W., 19 May 2001, GAH, 4 sh (adults, alive

& dead), CGAH; Grotte de Sare, rocky limestone slope with deciduous woodland

cover, 220 m alt., 30T 061587/479160, 7 May 2011, GAH, DTH, site E146, 1 sp & 11 sh

(6 adults [1 damaged] & 6 immatures, 1 alive & 10 dead), CGAH.

C. subcylindrica TRANCE: Dept. Pyrénées-Atlantiques: near Grotte de Sare, 5.5

km S. of Sare, mixed deciduous wood on rocky limestone slope, moss-rich ground

vegetation, with Cryptazeca monodonta, 23 Aug. 1980, DTH, 3 sh (2 immatures & 1

adult, dead), NMW.Z. 1993.052.04420; Grotte de Sare, rocky limestone slope, shaded

by mixed deciduous trees, very mossy with dead wood lying, ca 300 m alt., XNl, 6

Aug. 1983, DTH, MBS, 1 sh (immature, dead), NMW.Z. 1993.052.07653; Grotte de Sare,

rocky limestone slopes shaded by mixed deciduous wood, moist & mossy, XNl, 28

Aug. 1984, DTH, MH, MBS, 5 sh (1 immature & 4 adults, alive & dead),

NMW.Z.1993.052.10530; Grotte de Sare, rocky limestone slope with deciduous wood¬

land cover, 220 m alt., 30T 061587/479160, 7 May 2011, GAH, DTH, site F146, 2 sh (1

adult & 1 immature, dead), CGAH.

101

Iberus, 30 (1), 2012

C. vasconica SPAIN, Prov. Alava: by N625 ca 3 km S. of Orduña, steep limestone

hillside with Fagus woodland, damp beneath, with few saplings & bushes, ca 610 m

alt., VN3, 26 Aug. 1984, DTH, MH, MBS, 3 sh (dead), NMW.Z.1993.052.10455; Prov.

Asturias: near AS261, by San Ignacio, limestone scree on wooded slope by stream

bank, ca 270 m alt., [approximately] 30T 03302/47871, 24 June 2007, GAH, site 46, (5

adults & 5 immatures, dead), CGAH; near N625 in N. part of Desfiladero de los Beyos

(by turn to San Ignacio), base of limestone crag with ledges, scree & ephemeral water-

fall, shaded by low deciduous woodland, 184 m alt., 30T 032997/478728, 13 May 2011,

GAH, DTH, site E159b, 3 sh (2 adults [1 broken] & 1 immature, dead), CGAH; Prov.

Cantabria: just SE. of Llaguno (SW. of Castro-Urdiales), limestone road-cutting, crag &

deciduous woodland on slope, 252 m alt., 30T 04786/47940, 10 May 2011, GAH, DTH,

site E153, 8 sh (2 adults & 6 immatures, 3 alive & 5 dead), CGAH; S. of CA250 just

NW. of Otañes, base of limestone crag with few oak trees, block-scree locally, 136 m

alt., 30T 04830/47985, 10 May 2011, GAH, DTH, site E152, 5 sh (3 adults & 2 immatu¬

res, dead but 1 shell fresh), CGAH; Prov. Guipúzcoa: by GI3592 NE. of Araotz, rocky

limestone slopes & crags, above river, with patchy scrub & low woodland, 398 m alt.,

30T 05459/47610, 9 May 2011, GAH, DTH, site E150, 3 sh (adults, alive), CGAH; Prov.

Navarra: by NA4150 ca 2 km E. of Puerto de Usateguieta, rocky limestone slopes with

deciduous woodland (mainly beech), 644 m alt., 30T 059231/477150, 8 May 2011,

GAH, DTH, site E147, 2 sh (1 adult & 1 immature, alive & dead), CGAH; Prov.

Vizcaya: W. of Ranero (E. of Ramales de la Victoria), from litter at base of deep shaded

hollows on limestone slope, ca 500 m alt., 30T 04692/47902, 14 & 15 June 2007, GAH,

DTH, site 40, 3 sp & 29 sh (18 adults [1 broken] & 14 immatures, 12 alive & 20 dead),

CGAH; by BI2543 near Dima, rocky limestone slope with crags and patches of Hazel

coppice, 247 m alt., 30T 05209/47746, 9 May 2011, GAH, DTH, site E151, 4 sh (2 adults

& 2 immatures, 1 alive & 3 dead), CGAH.

102

Iberus, 30 (1): 103-110, 2012

© Sociedad Española de Malaeología

Reproduction in two sympatric Iberian nudibranch species

(Gastropoda: Opisthobranchia) with contrasting develop-

ment types

Reproducción en dos especies simpátricas de nudibranquios

(Gastropoda: Opisthobranchia) de la Península Ibérica con tipos de

desarrollo alternativos

Gonzalo CALADO*, Cláudia SOARES** and Ricardo C. NEVES***

Recibido el 16-XI-201 1. Aceptado el 7-X1I-201 1

ABSTRACT

Doriopsillo areolato and D. pelseneeri are two sympatric species from the Iberian Penin-

sula that contrast in their reproductiva modes. In this study we investigated the develop-

mental mode of both species, comparing egg number and egg volume. In addition, we

investígate whether the type of development influences the amount of energy allocated for

reproduction.

Animal weight was found to be positively correlated with egg number per egg mass but not

with egg volume, which seems to be a specific character in both species. Moreover, our results

suggest that both species allocate the same amount of energy for reproduction, in terms of

body weight, though a clear difference in the way these resources are used was actually

observed. Doriopsillo areolato invested 2.22 ± 1 .87% of body weight (n=3ó) essentially for

egg production, whereas D. pelseneeri invested 5.72 ± 3.31% of body weight (n-l 1) on

rich gel matrix spawn masses. The different type of development could provide an explana¬

ron for the difference in the geographical range of both species, but further studies are required.

RESUMEN

Doriopsillo areolato y D. pelseneeri son dos especies simpátricas en la Península Ibérica,

que contrastan en sus modos de reproducción. En este trabajo se investigó el modo de

desarrollo de ambas especies, comparando el número y el volumen de huevos. Además,

se investiga si el tipo de desarrollo influye en la cantidad de energía dedicada a la repro¬

ducción.

El peso del animal muestra una correlación positiva con el número de huevos por masa

de huevos, pero no con el volumen del huevo, que parece ser un carácter específico en

ambas especies. Nuestros resultados sugieren que ambas especies asignan, en términos

de peso corporal, la misma cantidad de energía para la reproducción, pero la forma en

que estos recursos se utilizan es diferente. Doriopsillo areolato, invirtió 2,22 ± 1,87% de

su peso corporal ¡n = 36), esencialmente para la producción de huevos, mientras que D.

pelseneeri ha invertido 5,72 ± 3,31% de su peso corporal (n = 1 1) en la matriz de gel

enriquecido de las masas. El tipo de desarrollo podría explicar la diferencia en la distribu¬

ción geográfica de las dos especies, pero se necesitan más estudios.

Universidade Lusófona de Humanidades e Tecnologías, Av. do Campo Grande, 376, 1749 - 024 Lisboa,

Portugal (e-mail: goncalo.calado@ulusofona.pt)

** Centro de Ciencias do Mar Universidade do Algarve, Campus de Gambelas 8005-139 Faro, Portugal

*** Biozentrum, University of Basel, Klingelbergstrasse 50, CH-4056 Basel, Switzerland

103

Iberus, 30 (1), 2012

INTRODUCTION

As far as their reproduction is con¬

cerned, opisthobranchs display a diver-

sity of strategies (Rudman and Willan,

1998). Several works report on species

displaying difieren! larval development

types: planktotrophy, lecithotrophy, and

direct development (Hadfield and

Switzer-Dunlap, 1984; Hadfield and

Miller, 1987; Bouchet, 1989; Todd,

1983). However, the selective factors in

determining these larval strategies

remain to be ascertained. According to

Hadfield and Miller (1987), for any of

those three "different means to the same

end" the time required to reach the juve-

nile stage is variable; usually, the larger

the egg, the longer the pre-hatching

period.

Doriopsilla areolata (Bergh, 1880) and

Doriopsilla pelseneeri (d'Oliveira, 1895)

are two radula-less, porostomate nudi-

branchs belonging to the family

Dendrodorididae. They are amongst

the most common nudibranchs inhab-

iting the Iberian coasts. Although

sympatric and occasionally feeding

upon the same sponges (GC, personal

observations), both species have rather

different types of development. Doriop¬

silla areolata has a planktotrophic

development type, producing egg

masses that are formed by numerous

small eggs, while D. pelseneeri displays

a metamorphic, direct development

type and produces a lower number of

larger eggs (Soares and Calado,

2006).

The known geographical range is

also different. Doriopsilla areolata is

known to occur between the North coast

of Spain and the Cape Verde Islands

(including the Mediterranean Sea),

Ghana and Angola, whereas D. pelse¬

neeri is restricted to the Atlantic and

Mediterranean coasts of the Iberian

Peninsula. The aim of this study is to

compare the developmental mode of

both species, comparing egg number

and egg volume. Further, we intend to

test whether the type of development

influences the amount of energy allo-

cated for reproduction.

MATERIALS AND METHODS

Animáis of both species were col-

lected from two localities along the

Arrabida coast (West Portuguese Coast)

and Ria de Ferrol (Galicia, NW Spain).

The exact localities can be found in

So ARES AND CALADO (2006). All speci-

mens were collected by hand while

scuba diving at 5-15 m depth from

March to July of 2003, in the period

reported as the peak of their reproduc¬

tion period.

Animáis were maintained under lab-

oratory conditions as described else-

where (Soares and Calado, 2006).

Specimens were paired periodically for

copulation and after approximately one

week, egg masses were laid and

removed immediately. In order to estí¬

mate the number of eggs, spawn masses

were measured (top, base length, and

width). We collected a total number of

61 Doriopsilla areolata and 39 D. pelse¬

neeri, from which we studied 43 and 21

laid egg masses, respectively. Each set

animal-spawn mass was weighed in

fresh for later use in calculations.

As for Doriopsilla areolata, three Imm

wide transverse sections of the egg

masses were cut for observation under a

stereomicroscope. Spawns from D. pelse¬

neeri were observed intact under a stere¬

omicroscope. Immediately after oviposi-

tion, and before any cleavage was

observed, eggs were counted in a

known egg mass area and its larger

diameter measured to estímate egg

number per egg mass and egg volume,

respectively.

In order to establish a calibration

curve fresh weight/dry weight, three

Doriopsilla pelseneeri specimens and four

D. areolata specimens were weighed

without water excess and dried at 60°C

to constan! weight (Smith and Sebens,

1983). Afterwards, a fraction of all dried

samples were used to calcúlate ash

valúes by ignition (3 hours at 550°C in a

muffle furnace), by the estimation of

ash-free dry weight equivalents

(Havenhand and Todd, 1988; Todd,

1979). The same procedure was used on

five egg masses of each species, as well

104

Calado ET al.: Reproduction in two sympatric Iberian nudibranch species

D. pekeneeri adult weigrh (g)

Figure 1 . Relationship between egg number per egg mass and adult animal weight for (A) D. areo-

lata (R2= 0.563) and (B) D. pelseneeri (R2= 0.768).

Figura 1. Relación entre el número de huevos por masa de huevo y peso del animal adulto en (A) D.

areolatá (R2- 0,563) y (B) D. pelseneeri (R2= 0,768).

as in two spawn masses of D. pelseneeri,

from which eggs were previously

removed manually. For D. areolata it was

assumed that egg masses were entirely

composed of eggs, since it was impossi-

ble to detach eggs from the feeble gel

matrix. A measure of the reproductive

effort per egg mass was calculated as

the ratio between the dry weight of a

single egg mass and the dry weight of

the progenitor adult inferred from the

fresh weight by the calibration curve.

Finally, how much energy (inferred by

the amount of biomass) is allocated per

egg mass and per egg unit, was calcu¬

lated.

Statistical analyses were conducted

using STATISTICA (Release 6.0). Regres-

sion analysis was performed to investí¬

gate the relationship between both egg

number and egg volume (mean valúes)

and adult body weight. Differences

between egg masses of the distinct pop-

ulations (Portugal and Spain) were also

investiga ted.

RESULTS

Egg number per egg mass and egg

volume

As presented in Soares and

Calado (2006), the egg number per egg

mass in D. areolata was estimated to be

between 5,500 - 240,000 eggs (n-43),

whereas in D. pelseneeri egg number

varied from 300 to 5,000 (n=15) eggs per

egg mass.

In both species it is clear that egg

number in a single spawn mass is posi-

tively correlated with animal weight

(R2= 0.563 for Doriopsilla areolata and

R2= 0.768 for D. pelseneeri) as can be

seen in Figure 1. The geographical area

105

Iberus, 30 (1), 2012

D. areolata adult weigrh (g)

Figure 2. Relationship between egg volume and adult animal weight for (A) D. areolata (R2=

0.006) and (B) D. pelseneeri (R2= 0,240).

Figura 2. Relación entre el volumen del huevo y peso del animal adulto en (A) D, areolata (R2= 0,006)

y (B) D. pelseneeri (R2= 0,240).

from where animáis were collected

seems to have no influence on this

analysis. However, specimens from

Galicia were observed to be slightly

smaller and therefore to deposit smaller

egg masses than those collected in Por¬

tugal.

Moreover, it was observed that egg

volume does not appear to co-vary with

animal weight (R2= 0.006 for Doriopsilla

areolata and R2= 0.240 for D. pelseneeri)

as illustrated in Figure 2. Indeed, egg

volume seems to be a specific character

in both species.

Reproductive investment in the

two species

Dry weight analysis revealed that

Doriopsilla pelseneeri's body contains on

average 87.94 ± 2.22% (n=3) of water.

whereas D. areolata contains 84.20 ±

2.41% (n=4), which reveáis a similar

hydrate condition in both species. From

the ash content analysis it was estimated

that D. pelseneeri contains 4.30 ± 1.42%

(n=3) of inorganic matter in the body,

whereas D. areolata contains 7.95 ±

1.34% (n:=4) (mean valúes).

Doriopsilla pelseneeri egg masses con-

tained 90.57 ± 3.03% (n=5) of water and

4.13 ± 0.89% (n=5) of inorganic matter,

whereas D. areolata egg masses con-

tained 92.73 ± 2.08% (n“5) of water and

4.55 ± 0.85% (n=2) of inorganic matter

(mean valúes). In the former species egg

masses are made of 39% of eggs and

61% of gel matrix, which contains 88.53

± 1.66% (n=2) of water and 6.90 ± 1.08%

(n~2) of inorganic matter. Since for D.

areolata it has been impossible to detach

106

Calado ET al.: Reproduction in two sympatric Iberian nudibranch species

Weight % invested

per egg (xlO'^)

0 1

3 4 5 6 7

9 10 11 12 13 14 20 21 22 23 24 25

B

D. aerolata

D. pebeneeri

Weight % invested

per egg masse

- j ^ ^ - 1

0 1 2 3 4 5 6 7 8 9 10

Figure 3. Relationship between weight invested (A) per egg mass and (B) per egg (%) for D. areo-

íata and D. pelseneeri.

Figura 3. Relación entre el peso invertido (A) por unidad de masa de huevos y (B) por huevo (%) para

D. areolataj D. pelseneeri.

eggs from the jelly matrix, egg masses

were assumed to be made up nearly

100% of eggs.

With these valúes we obtain that D.

pelseneeri invested 0.0012 ± 0.0009% of

their weight per single egg (n^ll, mean

valué of the range 0.00009% - 0.00243%),

while D. areolata invested 0.00026 ±

0.00009% of their body weight per

single egg (n-36, mean valué of the

range 0.0000032% ^ 0.00039%) (Figure

3A). The investment per egg mass was

also calculated; Doríopsilla pelseneeri

specimens studied invested 5.72 ± 3.31%

of their body weight (n=ll, mean valué

of the range 0.305% - 9.479%), whereas

D. areolata, invested 2.22 ± 1.87% (n=36,

mean valué of the range 0.012% -

4.767%) (Fig. 3B).

DISCUSSION

Developmental patterns in opistho-

branchs have been a subject of much

Ínteres! in the last decades (e.g. Had-

FIELD AND SWITZER-DUNLAP, 1984; FIaD-

EIELD AND MlLLER, 1987; TODD, 2001,

Krug, 2009). However, obtaining a final

consensus has not been possible so far

and some authors claim that there is no

single answer to what determines the

107

Iberus, 30 (1), 2012

developmental mode of any individual

opisthobranch species (Hadfield and

Miller, 1987). Although sympatric and

feeding on the same food Ítems, Doriop-

silla pelseneeri and D. areolata exhibit dif-

ferent developmental patterns. Soares

AND Calado (2006) demonstrated that

D. areolata with planktotrophic develop“

ment takes approximately half the time

till hatching than D. pelseneeri with

direct deveiopment.

Even with rather opposite develop¬

mental modes both species allocate

about the same amount of energy for

reproduction, as inferred from differ-

enees in body weight. However, there is

a clear difference in the way these

resources are used. Doriopsilla areolata

expends most resources producing

spawns with a high number of small

eggs, and a very scarce gel matrix.

Larvae hatch just a few days after egg

mass deposition, as is typical for obliga-

tory planktotrophs, and thus not many

nutritivo reserves are needed. In opposi-

tion, D. pelseneeri produces a small

number of large, yolky eggs and a dense

gel matrix in which they are embedded.

The matrix is used for mechanical and,

most certainly, for Chemical protection

of the embryos developing inside the

egg-capsule. Since we did not have

access to the complete reproductivo

period, we were unable to calcúlate the

total reproductivo effort for these two

species and therefore to compare it with

others (e.g. Todd, 1979; DeFreese and

Clark, 1983; Gibson and Chía, 1991).

Another conclusión arises from our

results, larger adult specimens produce

higher numbers of eggs per egg mass.

This has been observed for both species

studied herein. Allen, Krug and Mar-

SHALL (2009) report for the sacoglossan

Elysia stylifera that larval size is deter-

mined by the extra-capsular yolk, but not

by egg size. Todd, Lambert and Davies

(2001), based on previous studies with

Adalaria próxima (Alder and Hancock,

1854) (Lambert, Todd and Thorpe,

2000) generalise that between-population

variation in egg size is genetically deter-

mined. Such a pattern was not observed

herein. Egg size appears to be species

specific regardless of the geographic area,

and henee of the population.

As Strathmann (1986) refers, "dif-

ferences in size and body plan account

for much of the variation in types of

larval deveiopment". Recently and

using opisthobranchs as a model,

Goddard (2004) agrees with this point

of view. This author argües that phylo-

genetic constraints on developmental

mode can be severe. For instance, if a

feeding structure is lost the possibility

of it being regained is very remóte, con-

straining the whole lineage to a non-

feeding developmental mode. In gas-

tropods, however, this process could

have happened more easily since

lecitotrophic larvae often retain struc-

tures used in the process of feeding

(Strathmann, 1978).

Since planktotrophy is clearly the

most frequently observed deveiopment

mode, Hadeield and Miller (1987)

assume that this is also the most primi¬

tivo developmental strategy amongst

opisthobranchs. They argüe that the

evolutionary direction will be towards

lecithotrophic-planktonic deveiopment

and from there to direct deveiopment, in

which the opisthobranchs with ameta-

morphic deveiopment are the most

evolved forms. According to Todd

(2001), a larger egg should not be the

trigger for this chango and for Hadfield

AND Miller (1987) it is not clear if the

selectivo pressures responsible for this

have been the same in all opisthobranch

species. Intense predation on early juve¬

niles could have selected for increased

size, leading to a direct deveiopment

mode, but in another situation the opis¬

thobranch can be the predator and may

adapt its own life cycle to grow and

reproduce before a short-living prey is

exhausted (Todd and Doyle, 1981). The

costs for this adaptation will be, in part,

a reduction in opisthobranch size and

fecundity, and production of larger,

lecithotrophic eggs or directly develo-

ped offspring (pelagic larva could be

dispensable if prey is not too patchy)

will be favoured (Hadfield and

Miller, 1987). According to Todd (1991)

a consensus is lacking and probably

108

Calado ET al.: Reproduction in two sympatric Iberian nudibranch species

there are special cases for particular

species.

Planktotrophic species have a high

fecundity and great dispersal potential,

whereas species with direct develop-

ment have a reduced fecundity with a

higher larval stage protection and

usually hatch and settle in the same

places where their progenitors live

(ScHELTEMA, 1986). The data available

on the two species herein studied

support this idea. Doriopsilla areolata,

with planktotrophic development, has a

wide geographic range whereas D. pelse-

neeri, with direct development, has a

much more restricted distribution,

mostly limited to the Iberian Peninsula

(see also Valdés and Ortea, 1997).

Potential dispersión capability of a

species is dependen! on many physical

and biological habitat conditions

(ScHELTEMA, 1986), however, and since

both species feed on a variety of

sponges widespread in the Lusitanian

province (GC, personal observations),

no host constraint should be involved in

their actual distribution.

Studying two sympatric dorid nudi-

branchs with different larval strategies,

Todd (1979) claimed that planktotrophic

species were 'Torced" into that develop-

mental mode since an insufficient

number of lecithotrophic eggs would be

produced to maintain its population sta-

bility. This author also suggested that

selection always favours the strategy

BIBLIOGRAPHY

Allen R.M., Krug P.J. and Marshall D.J.

2009. Larval size in Elysia stylifera is deter-

mined by extra-embryonic provisioning but

not egg size. Marine Ecology Progress Series,

389: 127-137.

Bouchet P. 1989. A review of poecilogony in

gastropods. Journal of Molluscan Studies, 55:

67-78.

DeFreese D.E. and Clark K.B. 1983. Analy-

sis of reproductive energetics of Florida

Opisthobranchia (Mollusca: Gastropoda).

International Journal of Invertebrate Repro¬

duction, 6: 1-10.

which produces the larger offspring sur-

vivorship even if it is energetically inef-

ficient. However, as seen above, other

authors claim that energetic factors

seems to be a lesser deterministic factor

for the selection of developmental

pattern.

As Hadfield and Miller (1987)

already pointed out, there is "no valid

reason to assume that species currently

found together evolved in sympatry or

under the conditions in which they are

now found". Furthermore, it is not

certain that planktotrophy and lecitho-

trophy are different processes aiming at

the same final results (Todd, 1991). We

are convinced that further phylogeo-

graphic studies on the populations of

Doriopsilla areolata and Doriopsilla pelse-

neeri will help to elucidate the origins of

the speciation process and therefore the

evolutionary relationships between

these two species.

ACKNOWLEDGMENTS

We would like to thank Dr. Henrique

Cabral for his help on statistical analy-

sis. Fátima Martins and Peter Wirtz

kindly provided the photos of the

spawn masses of D. areolata and D. pelse-

neeri that illustrate Figure 3A. The

second author holds a grant from the

Fundaíjáo para a Ciencia e Tecnologia,

Portugal (BD/82339/2011).

Gibson G.D. and Chía F.-S. 1991. Contrast-

ing reproductive modes in two sympatric

species of Haminaea (Opisthobranchia:

Cephalaspidea). Journal of Molluscan Stud¬

ies, 57: 49-60.

Goddard J.H.R. 2004. Developmental mode

in benthic opisthobranch mollusks from the

northeast Pacific Ocean: feeding in a sea of

plenty. Canadian Journal ofZoology, 82:1954-

1968.

Hadfield M.G. and Miller S.E. 1987. On de¬

velopmental patterns of Opisthobranchs.

American Malacological Biilletin, 5: 197-214.

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Hadfield M.G. and Switzer-Dunlap M. 1984.

Opisthobranchs. In: The Mollusca (Reproduc-

tion), A. S. Tompa, N. H. Verdonk and J. A.

M. van den Biggelaar, eds. pp. 209-350. Aca-

demic Press, New York.

Havenhand J.N. and Todd C.D. 1988.

Physiological ecology of Adalaria próxima

(Alder et Hancok) and Onchidoris muricata

(Müller) (Gastropoda: Nudibranchia). I.

Feeding, growth, and respiration. Journal of

Experimental Marine Biology and Ecology,

118:151-172.

Krug P.J. 2009. Not my "Type": Larval dis-

persal dimorphisms and bet-hedging in

opisthobranch life histories. Biological Bu-

lletin, 216: 355-372.

Lambert W.J., Todd C.D. and Thorpe J.P.

2000. Variation in growth rate and repro-

ductive output in British populations of the

dorid nudibranch Adalaria próxima: conse-

quences of restricted larval dispersal. Ma¬

rine Biology, 137: 149-159.

Rudman W.B. and Willan R. 1998. Opistho-

branchia. Introduction. In: Mollusca: The South¬

ern Synthesis. Fauna of Australia, Vol. 5, Part

B, P.L. Beeseley, G.J.B. Ross, A. Wells, eds. pp.

915-942. CSIRO Publishing, Melbourne.

SCHELTEMA R.S. 1986. On dispersal and plank-

tonic larvae of benthic invertebrates: an ecletic

OverView and summary of problems. Bid-

letin of Marine Science, 39: 290-322.

Smith D.A. and Sebens K.P. 1983. The physi¬

ological ecology of growth and reproduc-

tion in Onchidoris aspera (Alder and Han¬

cock) (Gastropoda: Nudibranchia). Journal

of Experimental Biology and Ecology, 72:287-

304.

Soares C. and Calado G. 2006. Spawn and

early development of two sympatric species

of the genus Doriopsilla (Opisthobranchia:

Nudibranchia) with contrasting develop¬

ment strategies. Records of the Western Aus-

tralian Miiseum. Supplement 69: 103-109.

Strathmann R.R. 1978. The evolution and loss

of feeding larval stages of marine inverte¬

brates. Evolution, 32:894-906.

Strathmann R.R. 1986. What Controls the type

of larval development? Summary statement

for the evolution session. Bulletin of Marine

Science, 39: 616-622.

Todd C.D. 1979. Reproductivo energetics of

two species of dorid nudibranchs with plank-

totrophic and lecitotrophic larval strategies.

Marine Biology, 53: 57-68.

Todd C.D. 1983. Reprodiictive and trophic ecology

of nudibranch mollusks. In: The Mollusca (Ecol-

ogy), W. D. Russel-Hunter, ed. pp. 225-259.

Academic Press, New York.

Todd C.D. 1991. Larval strategies of nudibranch

mollusks: similar means to the same end?

Malacologia, 32: 273-289.

Todd C.D. and Doyle R.W. 1981. Reproduc¬

tivo strategies of marine benthic inverte¬

brates: a settlement-timing hypothesis. Ma¬

rine Ecology Progress Series, 4: 75-83.

Todd C.D., Lambert W.J. and Davies J. 2001.

Some perspectives on the biology and ecol¬

ogy of nudibranch mollusks: generalisations

and variations on the theme that prove the

rule. Bollettino Malacologico, 37: 105-120.

Valdés a. and Ortea J. 1997. Review of the

genus Doriopsilla Bergh, 1880 (Gastropoda:

Nudibranchia) in the Atlantic Ocean. The

Veliger, 40: 240-254.

lio

© Sociedad Española de Malacología

Iberas, 30 (1): 111-117, 2012

Asían openbill stoú^ Anastomus oscitans as a predator of the

invasive alien gastropod Pomacea canaliculata in Thailand

La cigüeña asiática de pico abierto Anastomus oscitans como un

depredador del caracol manzana Pomacea canaliculata^ especie exó¬

tica invasora en Tailandia

j

Weerachon SAWANGPROH*’***\ Philip D. ROUND** and Pilai POONSWAD***

Recibido el 19-IX-2011. Aceptado el 15-X1I-2011

ABSTRACT

The freshwater goiden apple snail Pomacea canaliculata, native to South and Central

America, and the West Indies, ¡s now a major agricultural pest in Thailand paddy fieids.

There is anecdotal evidence that the Asian openbill Anastomus oscitans consumes these

snails, but no reports on its predation efficiency. Our study showed that the number of

storks visiting the fieids correlated with a decrease in snail abundance (rs= 0.902, p=

0.0001). Storks are tactile predators and preferred adult snails. This is probably more pro-

fitable for the birds, but keeps the P. canaliculata population 'sustainable'.

RESUMEN

El caracol manzana dorado Pomacea canaliculata, una especie de agua dulce originaria

de América del Sur y Central y del Caribe, es actualmente una plaga agrícola importante

en los arrozales de Tailandia. Se sabe que las cigüeñas asiáticas de pico abierto Anasto¬

mus oscitans consumen estos caracoles, pero no hay datos sobre la eficacia de esta pre¬

dación. En este trabajo se muestra que hay una correlación entre el número de cigüeñas

que visitan los arrozales y una disminución en la abundancia de caracol (rs= 0,902, p=

0,0001). Las cigüeñas son depredadores táctiles y prefieren caracoles adultos. Esto es

probablemente más rentable para las aves, pero mantiene la población de P. canaliculata

en niveles "sostenibles".

INTRODUCTION

Goiden apple snails Pomacea canali¬

culata (Lamarck, 1822) are freshwater

gastropod molluscs originally native to

South America, Central America, and

the West Indies (Pain, 1972). They were

first introduced into Taiwan in Asia in

1979-80, to be cultivated as a high-

protein food source for local people and

export to industrialized countries

(Naylor, 1996). Subsequently, they were

* Program in Conservation Biology, School of Multidisciplinary Studies, Mahidol University, Kanchanaburi

Campus, 199 Moo 9 Lumsum Subdistrict, Saiyok District, Kanchanaburi Province 71150 Thailand

** Department of Biology, Faculty of Science, Mahidol University, Rama VI Road, Rachadhavi District,

Bangkok 10400 Thailand

*** Department of Microbiology, Faculty of Science, Mahidol University, Rama VI Road, Rachadhavi District,

Bangkok 10400 Thailand

^ Corresponding author: E-mail: kawsw@mahidol.ac.th

Iberus, 30 (1), 2012

introduced to other Asían countries,

including Japan, the Philippines, China,

South Korea, Malaysia, Thailand, Indo¬

nesia and Vietnam. In Thailand, P. cana-

liculata were first imported in 1982-83,

probably from Japan, Taiwan and the

Philippines via Chatuchak weekend

market for culturing as aquarium pets.

These snail farms were subsequently

abandoned, allowing the snails to

invade all natural freshwater environ-

ments and become a serious pest in

paddy fields, particularly in central

Thailand (Janyapeth and Archawa-

KOM, 1998).

P canaliciilata is a dioecious, sexually

dimorphic gastropod, with mature

males smaller than mature females

(SiTTi, 1988; Estebenet and Cazzaniga,

1998). Reproduction is continuous in

tropical zones (Martín, Estebenet and

Cazzaninga, 2001) and begins at ages

of 2-3 months after hatching (CowiE,

2002) or when they have reached a size

of > 25 mm (Sawangproh unpubl data).

Mating occurs at any time of the day

and in all seasons of the year, as long as

there is a sufficient supply of water

(Naylor, 1996). Females can lay eggs

weekly, on substrates from the bare

ground of canal banks to vegetation

above the water line. Clutch size varíes

from 400-3000 eggs, depending on the

availability of food and the size of the

reproductive female (Sitti, 1988; Janya¬

peth AND Archawakom, 1998), indica-

ting that control of snails of reproduc¬

tive age may be an effective control stra-

tegy. The control of apple snails as agri¬

cultura! pests using various biological

control agents has been extensively

reviewed (egrets - Agosta and Pullin,

1989; firefly larvae - Kondo and

Tanaka, 1989; water bugs - Aroonsri-

MORAKOT, 1993; fish - Halwart, 1998;

fire ants - Yusa, 2001; carp - Ichinose,

Tochihara, Wada, Suguiura and

Yusa, 2002; leeches - Aditya and Raut;

2005). However, studies of other orga-

nisms that prey on apple snails are still

needed.

Common large waders foraging in

paddy fields and irrigation canals in

central Thailand inelude herons and

egrets (Ardeidae), and a stork species

the Asían openbill Anastomus oscitans

(Boddaert, 1783) that feeds exclusively

on mollucs, especially the large native

freshwater apple snails Pila spp. (Kahl,

1971; McClure, 1974; Lauhachinda

1969; Lekagul and Round, 1991;

PooNSWAD, 1979). However, in response

to the spread of the alien P. canaliculata

in the aquatic environment, the native

Pila snail population has decreased

(Lauhachinda, Senawong, Phongs-

MAS, Udomchoke, Homchan and

Makatan, 1999) and the Asían openbill

has shifted its diet from Pila (PooNS-

WAD, 1979) to P canaliculata snails

(Khobkhet, 2000). The Asían openbill is

believed to be the most effective natural

predator of P. canaliculata (Khobkhet,

2000 and various farmers pers. comm.).

The objectives of this study were to

determine the feeding behavior of the

Asían openbill on P. canaliculata and

assess its efficieney as a predator on

these snail populations in paddy fields

of central Thailand.

MATERIALS AND METHODS

Study area

The study was conducted in Nakhon

Pathom Province (14° 00' N; 100° 14' E)

in central Thailand, where P. canaliculata

infestation is severe. We conducted our

studies on various paddy field blocks,

each about 2 acres in area, that oceurred

alongside 16-20 km of the local road No.

3004 just north of Salaya Campus,

Mahidol University (13° 50' N; 100° 18'

E). We also studied snail abundance and

size in Donthong Village (DT; 13° 47' N;

100° 16' E) and Sanchao Village (SC; 13°

50' N; 100° 16' E), which are about 6 km

apart and where no storks had ever

been sighted.

Flock size and feeding behavior of

storks

We counted the number of storks at

25 blocks of paddy fields along road No.

3004 and recorded their feeding beha¬

vior at first sighting through lOx binocu-

lars.

112

Sawangproh ET al.: Asían openbili stork predation on Pomacea canaliculata in Thailand

Figure 1 . Varying sizes of empty shell of P. canaliculata snails left in the paddy fields after stork

foraging (H= height; W= width).

Figura 1. Distintos tamaños de conchas vacias de caracoles P. canaliculata abandonados en los campos

de arroz después de excursiones alimenticias de la cigüeña (H= altura, W= anchura ).

Snail abundance and measurement

We distinguished snails that were

recently consumed by storks from those

that had died earlier, either from stork

predation or other causes by looking at

the fresh muscle remaining attached to

their shells and opercula, or at visceral

masses and reproductive organs, such as

albumen, found near the shells. We mea-

sured the size i.e. height and width of live

snails to the nearest 0.01 mm using

vernier calipers and then returned them

to the same fields, whereas dead shells

were brought to the laboratory for mea¬

surement (Figure 1). We also studied

snail abundance and size in DT and SC.

Effects of storks on snail abun¬

dance and size

To determine the effect of the storks

on snail abundance, we studied the

flock sizes of the storks and snail densi-

ties at another seven blocks of paddy

fields along road No. 3004, over ten con¬

secutivo weeks from late February to

April 2006. To determine the effect of

Asian openbili predation on snail sizes,

we randomly sampled live snails and

preyed-on shells using quadrat sam-

pling (quadrat size= 1x1 m, n- 20) after

the storks had been feeding in the seven

paddy fields from July-April 2006. All

paddy fields visited were at the phase of

field preparation for rice planting, when

rice stalks are burnt and the fields

flooded.

Statistical analysis

We analyzed flock sizes of Asian

openbili quantitatively and their feeding

behavior descriptively. To determine the

efficiency of stork in controlling snail

populations, we plotted the percentage of

snails preyed on by storks against the

number of storks visiting each field.

Based on flock size, we classified our 25

paddy fields during this study into three

classes: 1) 12 low openbill-pressure sites,

where visits by storks were known to be

frequent but with low abundance (mean

flock size 16.4 ± 15.3 storks/flock, n= 12),

2) 13 high openbill-pressure sites, where

stork visits were known to be frequent

and abundance high (53.0 ± 31.8

storks/flock, n= 13), with 3) one stork-

free site (DT-SC) as control. We classified

the data of snail sizes from all these

paddy fields and correlated them with

the three distinct classes based on stork

abundance and frequency of stork visit.

We categorized the snails into 2 size-

classes on their reproductive potential: a)

juvenile or non-reproductive (< 25 mm

high) and adult or mature (> 25 mm high;

based on Carlson, Brónmark, and

Hansson (2004). Finally, using the Good-

ness of fit test, we determined snail size

preferences as consumed by storks.

113

Iberus, 30 (1), 2012

Figure 2, Seven paddy fields showed decreasing snail abundance corresponding to the numbers of

foraging storks except the last four weeks when snails were brought into the fields accompanied by

water flooding.

Figura 2. Siete arrozales mostraron una disminución de la abundancia de caracoles correspondiente al

número de cigüeñas alimentándose, con la excepción de las cuatro últimas semanas, cuando más caraco¬

les fueron llevados a los campos acompañados por inundaciones.

RESULTS

Abundance of the Asían openbill

and its feeding behavior

Flock sizes of storks visiting the

paddy fields ranged from 6-133 storks

per flock (39.5 ± 30.5 storks/flock, n=

24). Stork feeding sites were characteri-

zed by flooded fields with water depth

ranging from 2.3-16.0 cm (6.4 ± 3.4 cm,

24). The flock sizes of storks were

positively correlated with snail abun¬

dance (live and preyed-on snails) in the

fields (rs- 0.6112, p= 0.0082).

We observed that storks searching

for snails probed with their bilis under

and adjacent to floating vegetation and

rice stalks. The stork slightly opened its

bilí and stabbed through the water.

When a snail was located, the stork

quickly closed its bilí, pinned the snail

to the ground with its mandibles and

then pushed the tip of the lower mandi¬

ble in to open snail operculum. Once the

operculum was opened, the stork then

inserted its mandibles, lifted up the

snail, shook its bilí to free the snaiTs

body from its shell, swallowed the flesh

and discarded the shell and operculum.

The effect of the Asían openbill on

snail abundance

Snail abundance at seven paddy

fields decreased in tándem with decrea¬

sing numbers of visiting storks, except

during the last four weeks of the study

when the fields were flooded with water

by farmers, bringing new snails from

nearby irrigation canals to settle in the

fields (Figure 2). In addition, we found

that the percentage of preyed-on snails

was positively related to the number of

visiting storks (rs= 0.902, p= 0.0001).

The effect of Asían openbill preda-

tion on snail sizes

Storks took a longer time (61.0 ± 50.7

seconds, n- 9) to successfully extract large-

sized snails (63.2 ± 10.99 mm in length, n=

80) comparing to the few seconds requi-

red to swallow each small-sized snail.

Storks apparently preferred large-sized (=

mature adult) snails to juvenile (= imma-

ture small-sized) snails (Table I).

DISCUSSION

Abundance of Asían openbills and

their feeding behavior

The flock size of storks varied depen-

ding on Pomacea snail abundance. All

paddy fields visited by storks were burnt

and then flooded shortly afterwards. At

this time, the habitat allows snails to

come out of their hibernating burrows,

grow and reproduce, making them avai-

lable to storks in the fields. We observed

1 14

Sawangproh ET al.: Asían openbill stork predation on Pomacea canaliculata in Thailand

Table 1. Summary of snail abundance, preference of snail sizes chosen by storks, and total number

of storks at three different sites, (juveniles immature, adult= matute)

Tabla /. Resumen de abundancias de los caracoles, tamaños de los caracoles preferidos por las cigüeñas, y

número total de cigüeñas en tres sitios diferentes. (juveniP inmaduro; adulto- maduro).

storks repeatedly visiting the same

paddy fields when snails were available,

but not fields with high, dense rice stalks

where it was possibly difficult to find

snails. When searching for snails, the

stork probes with its bilí in shallow water

with the tips of the mandibles slightly

open, griping a snail once fotmd and then

withdrawing its soft body from the shelL

This foraging method is known as tacto-

location (CouLTER, Balzano, Johnson,

King and Shannon, 1989) and is similar

to snaihlocating behavior in limpkins

Aramus guarauna (Linnaeus, 1766) (Sny-

DER AND Snyder, 1969). Extraction of Pila

snails by Asian openbill is the same as

described by Kahl (1971) where the tip

of the lower mandible is usually inserted

into the shell and the body dislodged

without help from the feet. V'^en feeding

on native Pila or exotic P. canaliculata

snails, Asian openbills did not damage

the shells from these observations. We

also observed Asian openbills discard the

albumen glands of mature female snails

on the ground where they fed. Possibly,

the yolk and eggs of this snail are distas-

teful, as suggested by Snyder and Kale

(1983), and the yellowish albumen cons-

picuous and presumably aposematic to

avian predators.

The effect of Asian openbills on

apple snail abundance and snail size

Large flocks of openbills could gre-

atly reduce snail populations in paddy

fields. Even though the cost of handling,

extracting, and consuming large-sized

snails was high compared to small-sized

snails, storks preferred to eat large-sized

snails, apparently because large-sized

snails gave more profitability (more

energy returns) as predicted by optimal

foraging theory (Pyke, 1984). Small-sized

snails, which openbills swallowed whole,

included much more indigestible mate¬

rial (hard shells and corneous opérenla),

than the large-sized snails where open¬

bills left the hard shells (Figure 1) and

tough opérenla in the fields.

115

IberuSy 30 (1), 2012

The preference for large-sized or

adiilt snails by openbills may negatively

affect the reproductive success of the

snails, and consequently their popula-

tion dynamics. On the other hand, the

remaining small-sized snails are

allowed to grow and reproduce, inci-

dentally making the population a 'sus-

tainable' food source for the storks.

Overall, openbills were considered

effective agents in controlling the snail

population while crops were present;

particularly those of reproductive size

that would cause most crop damage

(Table I).

The limpkin, a tactile predator like

the Asian openbill, acts as an agent of

disruptive selection on P. paludosa snail

size by selecting average-sized snails

disproportionately more often than

small or large snails (Reed and Janzen,

1999). Snail kites Rostrhamus sociabiíis

(Vieillot, 1817), which feed exclusively

on P. paludosa snails in their native range

in Florida, Venezuela, and Colombia,

search for prey visually (Beissinger,

1983, 1990; Snyder and Kale, 1983) and

seemed to be agents of directional selec¬

tion against lighter colored snails (Reed

and Janzen, 1999). From our observa-

BIBLIOGRAPHY

Acosta B.O. and Pullin R.S.V. 1989. Envi-

ronmental impact of the golden apple snail

(P omacea sp.) on rice farming system in the

Philippines. Siimmary report of the workshop

held at the Freshwater Aquacultiire Center Lu-

zon State University Nueva Ecija, Philippines.

9-10 November 1989. 34 pp.

Aditya G. and Raut S.K. 2005. Feeding of the

leech Glossiphonia iveberi on the introduced

snail Pomacea bridgesii in India. Aquatic Eco-

logy, 39 (4): 465-471.

Aroonsrimorakot S. 1993. Biology of small

water bug {Sphaeroderma molestum (DUF.) in

the control of golden apple snail {Pomacea

canaliculata). M.Sc. thesis. Mahidol University,

Bangkok, Thailand, 75 pp.

Beissinger S.R. 1983. Hunting behavior, prey

selection, and energetics of snail kites in Gu¬

yana: consumer choice by a specialist. The

AuK 100: 84-92.

Beissinger S.R. 1990. Alternative foods of a diet

specialist, the snail kite. TheAuk, 107: 327-333,

tion, storks evidently showed size prefe¬

rence for adult snails over juvenile

snails at every study site (Table I).

CONCLUSIONS

The Asian openbill, a residen! bird,

is an importan! predator of the mature

stage of the invasive alien golden apple

snail in Thailand, where rice planting is

practiced all year round. Based on snail

size and abundance, openbills could

potentially feed on and then lower snail

population of all sizes, but preferred the

large-sized snails that were probably of

a more profitable size.

ACKNOWLEDGEMENTS

We are very grateful to the UDC

scholarship for support of our expenses

until we finished our field work. We

would like to extend special thanks to

the Department of Biology, Faculty of

Science, Mahidol University for allo-

wing US to use some facilities. Finally,

we have to thank Dr. Alan Kemp for

checking the English manuscript.

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SON L. 2004. Invading Herbivory: The gol-

den apple snail alters ecosystem functio-

ning in Asian wetlands. Ecology, 85 (6):

1575-1580.

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C.E. AND Shannon P.W. 1989. Conservation

and captive management of storks. Proceedings

of an International Workshop at the New

York's Zoological Society's Wildlife Survival

Center, Saint Catherine's island, Georgia,

U.S.A., 80 pp.

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as agricultural pests: their biology, impacts,

and management. In Barker G.M. (Ed): Mo-

lluscs as Crop Pests. CABI, New York, pp 145-

192.

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Sex-related differential growth in Pomacea

canaliculata (Gastropoda: Ampullariidae).

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116

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Harwart M. 1998. Cyprinus carpió and Ore-

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of the golden apple snail Pomacea canaliculata

- Effect of predator size, prey size, and prey

density. Asian Fisheries Science, 11: 31=42.

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GUIURA R. AND Yusa Y. 2002. Influence of

common carp on apple snail in a rice field eva-

luated by a predator - prey logistic model. In¬

ternational Journal ofPest Management, 48 (2):

133-138.

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cial seminar on the topic of "Cherry snail" at

Louis Tavern hotel in Bangkok, Thailand.

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openbili storks. Journal ofOrnithology, 112: 21-

35.

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rakadee Press, Thailand, 260 pp.

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mental study of predation by the larvae of the

firefly. Lucióla lateralis Motschulsky (Cole¬

óptera: Lampyridae) on the apple snail, Po¬

macea canaliculata Lamarck (Mesogastropoda:

Pilidae). Applied Entomology and Zoology, 33

(4): 211-216.

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the life history of open-billed stork, Anasto-

miis oscitans (Boddaert). M.Sc. thesis. Kaset-

sart University, Bangkok, Thailand, 51 pp.

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D., Udomchoke V., Homchan U. and Ma-

katan N. 1999. Crop protection and control

from the golden apple snail Pomacea canali¬

culata (Orbigny) by physiological means and

biocontrol. Final Report, Kasetsart University,

44 pp.

Lekagul B. and Round P.D. 1991. A guide to

the birds of Thailand. Saha Karn Bhaet, Bang¬

kok, Thailand, 457 pp.

McClure H.E. 1974. Migration and survival ofthe

birds of Asia. Sahamitr Karn Pim, Bangkok,

Thailand, 476 pp.

Martín P.R., Estebenet A.L. and Cazzaniga

N.J. 2001. Factors affecting the distribution of

Pomacea canaliculata (Gastropoda: Ampulla-

riidae) along its southernmost natural limit.

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sessing the cost of the golden apple snail in

Asia. Ambio, 25 (7): 443-448.

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survey. Journal of Conchology, 27: 453-462.

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- billed storks {Anastomus oscitans) and their

relationships to Pila snails and domestic

ducks. M.Sc. thesis. Mahidol University,

Bangkok, Thailand, 100 pp.

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tical review. Anniial Review of Ecology and

Systematics, 15: 523-575.

Reed W.L. and Janzen F.J. 1999. Natural se-

lection by avian predators on size and colour

of a freshwater snail {Pomacea flagellata). Bio¬

logical Journal of the Linnean Society, 63 (3):

331-342.

SiTTi S. 1988. Selective feeding on some aqua-

tic plants by the golden apple snail {Pomacea

canaliculata). M.Sc. thesis. Kasetsart Univer¬

sity, Bangkok, Thailand, 89 pp.

Snyder N.F.R. and Kale H.W. 1983. Mollusk

predation by snail kites in Colombia. The

Auk, 100: 93-97.

Snyder N.F.R. and Snyder H. A. 1969. A com-

parative study of mollusc predation by Limp-

kins, Everglade Kites, and Boat-tailed Grac-

kles. Living Bird, 8: 177-223.

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snail Pomacea canaliculata (Gastropoda : Am¬

pullariidae) by the fire ant Solenopsis geminata.

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117

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Iberus 30 (1) 2012

Houart R. and RoláN E. The genus Pradoxa Fernandes & Rolán, 1993 (Gastropoda: Murici-

dae) in SaoTomé, Príncipe and Annobón

El género Pradoxa Fernandes & Rolán, 1993 (Gastropoda: Muricidae) en SaoTomé, Principe y

, Annobón . 1-14

FíOLYOAK D.T. and Holyoak G.A. A taxonomic revisión of Oestophora barhula (Rossmássler,

' 1838) and O. barbolla (Servain, 1880), two Iberian endemic land-snail species (Gastro¬

poda: Trissexodontidae)

Revisión taxonómica de Oestophora barbula (Rossmássler, 1838) y O. barbella (Servain,

1880), dos especies de caracoles terrestres endémicas de la península ibérica (Gastropoda: Trisse-

xodontidae) . A . ' . . 15-40

PUSATERI R, Giannuzzi-Savelli R. and Oliverio M. A revisión of the Mediterranean Raphi-

tomidae 1: on the sibling species Raphitoma contigua Monterosato, 1884 and Raphitoma

spadiana n. sp. (Gastropoda, Conoidea)

Revisión de los Raphitomidae del Mediteráneo 1: las especies hermanas Raphitoma contigua

Monterosato, 7 5d4j)/ Raphitoma spadiana sp. (Gastropoda, Conoidea) . 41-52 •

Rolán E. and Go'RI S. New species of neogastropods from the islands of the Gulf of Guinea,

West Africa

Nuevas 'especies de neogasterópodos de las islas del golfo de Guinea, Africa occidental . . . 53-66

Moreno D. The genus Gibberula (Gastropoda, Cystiscidae) in the Cape Verde Islands with the

description of a new species

El género Gibberula ( Gastropoda, Cystiscidae) en el archipiélago de Cabo Verde con la descrip¬

ción de una nueva especie . 67-83

Campani E., Bartolini S. and Spanu M.T Alvania garrafensis Peñas & Rolán 2008 (Gas¬

tropoda: Rossoidae) from Croatian waters

Alvania garrafensis Peñas & Rolán 2008 ( Gastropoda: Rossoidae) en aguas croatas _ 85-89

Holyoak D,.T and Holyoak G.A. A review of species-limits in some Cryptazeca (Gastropoda:

Azecjdae)

Reconsideración de los limites entre especies en Cryptazeca ( Gastropoda: Azecidae) ... 91-102

Calado G.* Soares C. and Neves R.C. Reproduction in two sympatric Iberian nudibranch

species (Gastropoda: Opisthdbranchia) with contrasting development types

Reproducción en dos especies simpátricas de nudibranquios ( Gastropoda: Opisthobtanchia) de

la Península Ibérica con tipos de desarrollo alternativos . , . 103-110

Sawangproh W, Round PD. and PoONSWAD E Asian openbill sx.oúiAnastomus oscitans ás a predatpr

of the invasive alien gastropod Pomacea canaliculata in Thailand

La cigüeña asiática de pico abierto Anastomus oscitans como un depredador del caracol

manzana Pomacea canaliculata, especie exótica invasora en Tailandia . 111-117

ISSN 0212-3010

(QL

^01

.11^3

12.

WevMS

REVISTA DE LA

SOCIEDAD ESPAÑOLA

DE MALACOLOGÍA

Oviedo, ¡ulio 2012

Iberus

Revista de la

Sociedad Española de Malacología

Comité de Redacción (Board of Editors)

Editor de Publicaciones (Editor-in-Chief)

Serge Gofas Universidad de Málaga, España

Director de Redacción (Executive Editor)

Gonzalo Rodríguez Casero Mieres del Camino, Asturias, España

Editora Ejecutiva (Managing Editor)

Eugenia M" Martínez Cueto-Eelgueroso Mieres del Camino, Asturias, España

Editores Adjuntos (Associate editors)

Iberus gualtieranus (Linnaeus, 1758), una especie emblemática de la península Ibérica, que da

nombre a la revista. Dibujo realizado por José Luis González Rebollar “Toza”.

Iberus

-gS^HSOl^

AUG 28 2012

REVISTA DE LA

SOCIEDAD ESPAÑOLA

DE MALACOLOGÍA

Vol. 30 (2)

Oviedo, julio 2012

Iberus

Revista de la

Sociedad Española de Malacología

Iberus publica trabajos que traten sobre cualquier aspecto relacionado con la Malacología. Se

admiten también notas breves. Iberus edita un volumen anual que se compone de dos o más números.

Instrucciones para los autores

Los manuscritos deben remitirse a; Serge Gofas, Editor de Publicaciones, Departamento de Bio¬

logía Animal, Universidad de Málaga, Campus de Teatinos, s/n, 29071, Málaga, España.

Los trabajos se entregarán por triplicado (original y dos copias). Se recomienda a los autores leer

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Contents list published in Aquatic Science and Fisheries Abstracts and Zoological Records,

BIOSIS.

Dep. Leg. B-43072-81

ISSN 0212-3010

Diseño y maquetación: Gonzalo Rodríguez

Impresión: LOREDO, S. L. - Gijón

© Sociedad Española de Malacología - - — _ — _ — _ Iberus, 30 í(£): 1-6, 2012

Oscilla g^lilae, a new species of Pyramidellidae (Mollusca,

Gastropoda, Heterobranchia) from the Eastern

Mediterranean

Oscilla galilae, una nueva especie de Pyramidellidae (Mollusca,

Gastropoda, Heterobranchia) del Mediterráneo oriental

Cesare BOGI* ** ***, Selahattin Ünsal KARHAN” and Mehmet Baki YOKE§”*

Recibido el 26-1-2012. Aceptado el 12-III-2012

ABSTRACT

The finding of some specimens of a small pyramidellid along the Mediterranean coasts of

Turkey, Israel and Cyprus, previously reported off the south-eastern coast of Turkey (Buzzu-

rro & Greppi, 1996; Buzzurro ef al., 2001) as Hinemoa cylindrica (de Folin, 1879), indu-

ced US to revise this identification. Hinemoa cylindrica is of lndo=Pacific origin and has

been originally attributed to the genus Jo minea Brown, 1827 (not Risso, 1826). Later, it

has been transferred to the genus Hinemoa Oiiver, 1915 (Buzzurro et al., 2001). Howe-

ver, some details of its morphology do not agree with the description and figure of Jami-

nea cylindrica given by de Folin (1 879). For a more suitable generic placement, we com¬

pared the species to the members of some closely resembling taxa in Pyramidellidae Cray,

1840 (e.g., Cingulina, A.Adams, 1860, H/nemoo Oiiver, 1915, Miralda A. Adams,

1864, Odetta de Folin, 1870 and Oscilla Adams A., 1861). Its morphoiogical characte-

ristics have led us to assign it to the genus Oscilla as a new species.

RESUMEN —

El hallazgo de algunos ejemplares de un pequeño piramidélido en las costas mediterrá¬

neas de Turquía, Israel y Chipre, previamente citado en la costa sur-oriental de Turquía

(Buzzurro y Greppi, 1996;. Buzzurro et al, 2001) como Hinemoa cylindrica (de Folin,

1 879), llevó a revisar esta identificación. Hinemoa cylindrica es de origen Indo-Pacífico y

ha sido atribuida originalmente al género Ja minea Brown, 1827 (non Risso, 1826). Poste¬

riormente, se ha transferido a! género Hinemoa Oiiver, 1915 (Buzzurro et al., 2001). Sin

embargo, algunos detalles de su morfología no cuadran con la descripción y la figura de

Jaminea cylindrica dada por de Folin (1879). Con vistas a una identificación genérica

más adecuada, se compararon la especie con miembros de algunos grupos taxonómicos

más similares en Pyramidellidae Gray, 1 840 (por ejemplo, Cingulina, A.Adams de 1 860,

Hinemoa Oiiver, 1915, Miralda A. Adams, 1 864, Odetta de Folin de 1 870 y Oscilla A.

Adams, 1861). Sus características morfológicas nos han llevado a describir una nueva

especie en el género Oscilla.

* Cío Lippi Ello, Via Icilio Wan Bergher, 24 1-57122 Livorno, Italy. E-mail: bogicesare@tiscali.it

** División of Hydrobiology, Department of Biology, Faculty of Science, Istanbul University, 34134

Vezneciler, Istanbul, Turkey. E-mail: unsalkarhan@yahoo.com

*** Department of Molecular Biology & Genetics, Faculty of Arts & Sciences, Hali^ University, 3438 1 Sisli,

Istanbul, Turkey. E-mail: bakiyokes@halic.edu.tr

1

Iberus, 30 (2), 2012

INTRODUCTION

Ongoing malacological research and

the examination of numerous sediment

samples collected along the coasts of

Israel, in particular by dredging opera-

tions carried out off the port of Haifa,

have produced many interesting discove-

ries including numerous species of Indo-

Pacific origin (e.g., Bogi & Galil, 1997,

2000, 2007, 2009). This time we recorded

the discovery of some specimens of a

pyramidellid gastropod already reported

from the south-eastern coast of Turkey

(Buzzurro, Hoarau, Greppi & Pelorge,

2001) and identified by the authors as

Jaminea cylindrica de Folin, 1879, assigned

to the genus Hinemoa Oliver, 1915, albeit

tentatively. This species, whose type loca-

lity indicated by de Folin (1879) is

Borneo (South China Sea) has a wide

Indo-Pacific distribution, but has not yet

been reported from the Red Sea (Debcker

& Orlin, 2000). Flowever, morphological

characteristics of newly found specimens

make them easily distinguishable from

de Folin's species. Upon examination,

morphological characteristics of the

shells have led us to assign them to an

undescribed species of the genus Oscilla

A. Adams, 1861, but not Hinemoa. A des-

cription of the new species along with a

discussion of its taxonomic affinities is

presented herein.

TAXONOMY

Pyramidelloidea Cray, 1840

Pyramidellidae Cray, 1840

Oscilla A. Adams, 1861

Oscilla galilae n. sp. (Fig. lA-F)

Type material: The holotype (1.40 x 0.65 mm) (Fig. lA-E) is depositad in the National Collections

of Natural History at Tel Aviv University with the catalog number TAU MO 73668. The paratypes

are depositad in the following collections: paratypes A (Fig. IF) and E (1.55 x 0.68 mm and 1.27 x

0.60 mm, respectively) in the collection of S.Ü. Karhan and M.B. Yoke§ (Istanbul, Turkey); paraty¬

pes B, C and D (1.36 x 0.61 mm, 1.20 x 0.55 mm and 1.35 x 0.62 mm, respectively) in the collection

of C. Bogi (Livorno, Italy); paratypes F and G (1.71 x 0.70 mm, 1.52 x 0.65 mm, respectively) in the

collection S. Raveggi (Firenze, Italy).

The holotype comes from dredging carried out in Haifa Bay, Israel (32° 54.544' N, 35° 04.093' E) at

10.5 m depth on 31.V.2009; paratype A from sediments collected manually using SCUBA in Carmel

Reef, off Haifa (32° 50.529' N, 34° 56.637' E) at 21 m depth on 19.XL2009; paratypes B and C from

material collected by dredging carried out in the Port of Haifa (32° 54.433' N, 35° 01.661' E) at 25

m depth on 14.IV.2010; paratype D collected from the Port of Haifa (32° 54.357' N,35° 02.793' E) at

20 m depth on 12.X.2010; paratype E from sand sample collected using SCUBA off Ta§ucu, Mersin,

Turkey (36° 14.530' N, 33° 48.359' E) at 5 m depth on 03.IX.2007, and paratypes F and G off Capo

Greco, Cyprus (34° 58.3618' N, 34° 04.3682' E) at 30 m depth on August 2011. The holotype and

paratypes B, C and D were collected complete with soft parts.

Type locality: Sandy bottoms in and around the Port of Haifa (Mediterranean coast of Israel) at

depths between 10 and 25 m.

(Right page) Figure 1. A-E: Oscilla galilae n. sp., holotype (TAU MO 73668), actual height 1.40

mm. A: frontal view; B: lateral view; C: protoconch, lateral view; D: protoconch, apical view; E:

detall of the scuipture; F: paratype A, actual height 1.55 mm, frontal view. G: Jaminea cylindrica^

original illustration given by DE FOLIN (1879).

(Página derecha) Figura 1. A-E: Oscilla galilae n. sp., holotipo (TAU MO 73668), altura real 1,40

mm. A: vista frontal; B: vista lateral; C: protoconcha, vista lateral; D: protoconcha, vista apical, E:

detalle de la escultura, F: paratipo A, altura real 1,55 mm, vista frontal. G: Jaminea cylindrica, ilus¬

tración original por DE FOLIN ( 1 879).

2

Bogi ET al.: Oscilk galike, a new Pyramidellidae from the Eastern Mediterranean

3

Iberus, 30 (2), 2012

Etymology: The new species is named after Dr. Bella S. Galil, in recognition of her extensive con-

tributions to taxonomy and to knowledge of the fauna from the Eastern Mediterranean Sea. The

epithet is a noun in the genitive case.

Description: Shell of médium size for

genus, attaining 1.71 mm height, elongate,

cylindrical or slightly oval; color uni-

formly whitish. Teleoconch consisting of

about three whorls, somewhat convex in

profile, separated by welhmarked and

slightly oblique suture. Protoconch

smooth, fíat and heterostrophic, type C

(van Aartsen, 1977), with about 0.5

whorl; diameter about 300 jum in ho-

lotype. The first whorl of teleoconch with

two spiral ribs, visible only by transpa-

rency inside the spire. Spiral sculpture

consisting of very pronounced spiral ribs,

rounded, almost uniform that tends to

increase with growth. Interspaces between

the ribs narrower than the ribs themseh

ves and crossed by numerous axial lame-

lias. Aperture about one-third of total

height, oval, with a thin outer lip; colu-

mellar tooth distinct; umbilicus absent.

Distribution: Geographical distribu-

tion of the species seems to be restricted

to shallow sublittoral areas in a few

localities on Israeli, Turkish and Cypriot

coasts in the eastern Mediterranean Sea.

Discussion: After examination of the

present specimens (fig. 1 A-F), our atten-

tion was drawn (J. Pelorce, pers. comm.)

to the fact that they correspond perfectly

to those previously collected on the

Turkish coast and identified as Hinemoa

cylindrica (de Folin, 1879), a new combi-

nation proposed for Jaminea cylindrica by

Buzzurro et al. (2001).

However, we have been able to note

some morphological details that do not

agree with the description and illustra-

tion of Jaminea cylindrica given by de

Folin. The original description of /. cylin¬

drica is as follows (de Folin, 1879: 266):

"Testa minutissima, sucylindrica, satis

lata, pauló elongata, alba; anfractus

embryonales II, obliquis ápice occulto;

normales III, in longitudine rapide

augentes, liris validis spirali bus ornati;

ultimo máximo, 1/3 testae aequante;

apertura ovata pauló obliqua, margine

columellare undulato, intus dentato.

Long.il mm 4; lat.: 0 mm 5.

Tres petite espéce, fort jolie et fort bien

caractérisée par la position oblique de ses

tours embryonnaires dont le sommet parait

caché; puis par safarme presque cylindrique,

peu allongée et relativament large. Les tours

normaux, au nombre de trios, s'allongent

rapidement, en demeurant presque de la

meme largeur. lis sont ornes de cordons

spiraux proportionnellement forts et bien

exprimes. L'ouverture est ovale, légérement

oblique; son bord columellaire est nodulé par

les courbes qu'il décrit; il est armé, au dedans

et vers son milieu, d'une dent assez sensible. "

Oscilla galilae n. sp. and Jaminea cylin¬

drica differ in several features. In O.

galilae there is a dense axial sculpture

consisting of small lamellae in the

interspaces between the spiral ribs (Fig.

lE), whereas in /. cylindrica spiral ribs

are in contad with each other without

any interspaces bearing axial sculpture

(Fig. IG). The new species differs from J.

cylindrica also in general shape of the

shell as the latter is more cylindrical, its

top whorl is more obtuse, and its sutu-

ring is more oblique (de Folin, 1879;

pl.9, fig.5). Spiral sculpture also differs

as the new species has more prominent

spiral cords than those of /. cylindrica.

Buzzurro et al. (2001) have also

noticed these discrepancies, but all the

same, probably taking into account the

cylindrical profile, well-marked colume-

llar tooth, flat protoconch and smooth first

whorl of teleoconch, they have assigned

their specimens to the species described

by de Folin (1879). Unfortunately it was

not possible to compare our specimens

with the holotype of /. cylindrica as it is not

present in the collection of de Folin stored

in MNHN (París, France) and is presumed

to be lost (V. Heros, pers. comm.). Since

the type material is no longer available, /.

cylindrica should therefore be considered

a nomen dubium.

In the family Pyramidellidae Gray,

1840 there are many morphologically

similar genera, some of which show only

minor differences from others. These difi

ferences are often insufficient to justify

4