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Dep. Leg. B-43072-81
ISSN 0212-3010
Diseño y maquetación: Gonzalo Rodríguez
Impresión: LOREDO, S. L. - Gijón
© Sociedad Española de Malacología
25,2014
New information on the marginellids of Sao Tomé and
Principe, with new records and the description of four new
species
Nueva información sobre los marginelidos de Sao Tomé y Principe,
con nuevas citas y descripción de cuatro especies nuevas
Emilio ROLÁN* & Sandro GORI**
Recibido el 5-XII-2012. Aceptado el 28-X-2013
ABSTRACT
Some marginellids collected in the islands of the Guiñean Gulf, Sao Tomé and Principe,
are studied, presenting new records, descriptions and figures of uncommon forms and
complementary information. Four new species are described.
RESUMEN
Se estudian algunos marginéiidos recolectados en las Islas del Golfo de Guinea, Sao
Tomé y Príncipe, y se presentan nuevas citas, descripciones y figuras de formas no habi¬
tuales, e información suplementaria. Se describen cuatro especies nuevas.
INTRODUCTION
The marginellids from the islands of
the Guiñean Gulf have been studied for
many years. Tomlin & Shackleford
(1912, 1913a & 1913b) mentioned from Sao
Tomé 13 species ñames from this group,
but many of them, such as Marginella mil¬
iaria (Linnaeus, 1758), M. bifasciata
Lamarck, 1822 and M. olivaeformis Kiener,
1834, belong to other geographical areas
and were not correctly identified. These
authors described four new species: Mar¬
ginella eveleighi, M. liparozona, M. melvilli
and M. chalmersi. Nobre (1886, 1887) also
recorded some species from these islands.
In more recent papers. Gofas & Fer-
nandes (1988) presented a revisión of the
family in the island of Sao Tomé, describ-
ing 11 species, of which 7 were new. Later,
Fern ANDES & Rolán (1992) provided new
records for both Sao Tomé and Principe
islands and described a new species. Since
these two papers and for many years, no
new publications were issued on the Mar-
ginellidae of these islands.
During recent years, the second
author (SG) visited several times these
islands, with many dives resulting in the
collection of a large amount of samples
(see collecting points in Figure 1 and
Table I); the Marginellids were set aside
in order to make a complete revisión of
the group but unfortunately some mater¬
ial was made available elsewhere so that
Cossignani (2012) described three new
species on the basis of only nine shells
and with scarce information, and further-
more made some mistakes that will be
discussed in the present work.
The study of several hundred speci-
mens and shells of Marginellidae is the
* Museo de Historia Natural, Universidad, 15782 Santiago de Compostela, Spain
** Via Sernesi, 7, 57123 Livorno, Italia
1
Iberus , 32 (1), 2014
6°30' 6°40' E
7°25' E
Figure 1 . Map of Sao Tomé and Principe islands, showing the location of collecting places.
Figura L Mapa de las islas de Sao Tomé y Principe, indicando la situación de los puntos de maestreo.
basis for the present work in which
some new records are made and 4 new
species are described.
MATERIAL AND METHODS
Most of the shells were collected by
scuba diving, in some cases at night.
Some places were visited and sampled
many times in nearly identical locations.
In some cases positions were recorded
with a portable GPS.
Abbreviations
MMM Malacologia Mostra Mondiale,
Cupra Marítima, Italy
MHNG Museum d'Histoire Naturelle,
Geneva, Switzerland
MHNS Museo de Historia Natural, San¬
tiago de Compostela, Spain
MNCN Museo Nacional de Ciencias
Naturales, Madrid, Spain
MNHN Muséum National d'Histoire
Naturelle, París, France
NMW National Museum of Wales,
Cardiff, Wales
CCS collection of Cedric Simbille, París,
France
CFB collection of Franck Boyer, Paris,
France
CFD collection of Francisco Déniz
Guerra, Canary Is., Spain
CJR collection of José Rosado, Maputo,
Mozambique
CPR collection of Peter Ryall, Maria
Rain, Austria
CSG collection of Sandro Gori, Livorno,
Italy
RESULTS
Genus Plesiocystiscus Coovert & Coovert, 1995
Plesiocystiscus violaceus spec. nov. (Figure 2A-H)
Type material: Holotype in MNHN (26621H, sp, Fig. 2A) and 2 paratypes in MNHN (26621P, sp
in alcohol). Other paratypes: MNCN 15.05/60085, 1 sp), CSG (2 sp); CPR (1 sp) and CJR (1 sp).
2
RoláN & GORI: New Information on the marginellids of Sao Tomé and Principe
Table L Coordinates of sampling places. Only the place ñame is given where a GPS was not used.
Tabla I. Coordenadas de los lugares de maestreo. Solamente se da el topónimo cuando no se usó un GPS.
Type locality: Sete Pedras, SE of Sao Tomé Island.
Etymology: The specific ñame alindes to the violet colour of the soft parts.
Description : Shell smalf shiny and
translucent, oblong, swollen posteriorly,
with a low and very globose spire.
Outer lip thickened and curved, inter-
nally smooth. Columelia with four weak
plaits, aperture narrow and elongate,
wider on the anterior part.
Soft parts: Described from the exam-
ined material after being fixed, but it was
recorded that this colour pattern was
3
IberuSy 32 (1), 2014
Figure 2. Plesiocystiscus violaceus spec. nov. A: holotype, 2.6 mm (MNHN); B-F: paratypes, with
soft parts; G: group of paratypes in alcohol; H: colour pattern.
Figura 2. Plesiocystiscus violaceus spec. nov. A: holotipo, 2,6 mm (MNHN); B-F: paratipos, con
partes blandas; G: grupo de paratipos en alcohol; H: patrón de color.
similar when alive: Animal with a red-
violet colour, on all the body parts visible
by transparency. It is a little darker in the
anterior and posterior parts comprised
within the shell; there are some small
oval lighter spots, not very numerous;
the visible parts of head, siphon and foot
are whitish with purple dots.
Dimensions: Holotype 2.6 mm.
Paratypes of similar size.
Habitat : Collected brushing spheroid
concretions of a coralline alga in 27-30 m
depth. These concretions are frequent on
the bottom surrounding the islands of
the Guiñean gulf .
Distribution: Only known from type
locality.
Remarks: The differences with other
described species from that area are
seen in the soft parts, the shell being
very similar:
- Plesiocystiscus gutta (Gofas & Fer-
nandes, 1988) is very dark in colour,
visible by transparency, with numerous
yellowish microspots and a few red
ones; the foot is whitish with some
brown spots on the anterior part,
including the head, and yellow on the
caudal part.
- Plesiocystiscus josephinae (Femandes
& Rolán, 1992) was collected in Principe
Island and it has, visible by trans¬
parency within the shell, very dark parts
anteriorly and posteriorly, with a wider
space between them which is pink with
small orange spots. The foot is whitish-
translucent and only the propodium is
orange.
Genus Volvarina Hinds, 1844
Volvarina insulana Gofas & Fernandos, 1988 (Figure 3A-E)
Volvarina insulana Gofas & Femandes, 1988: 13, figs. 14-15, pl. ID. [Type locality: Esprainha].
4
ROLÁN & GORI: New Information on the marginellids of Sao Tomé and Principe
Figure 3. Vaharina insulana Gofas & Fernandes, 1988; A-D: shells, 8.9, 8.1, 8.4, 8.2 mm,
Tinhosa Grande, 15 m, Principe Island (CSG); E: living animal on a dark background.
Figura 3. Volvarina insulana Gofas & Fernandes, 1988; A-D: conchas, 8,9; 8,1; 8,4; 8,2 mm,
Tinhosa Grande, 15 m, isla del Principe ( CSG); E: animal vivo sobre fondo oscuro.
Type material: Holotype in MNHN (represented in Gofas & Fernandes, 1988, fig. 14a, 14b).
Other material examined: Principe Island: 7 s, Pedra Galé, 10-35 m (CSG); 1 s, Tinhosa Pequeña,
41 m (CSG); 5 s, Sete Brazas, 21 m (CSG); 5 s, Sete Brazas, 20 m (CSG); 5 s, Tinhosa Grande, 15
m(CSG); 2 sp, Ponta Cabra, Baia de Santo Antonio, 10 m (CSG); 2 s, Tinhosa Pequeña West, 42 m
(CSG); 1 s, Ponta da Mina, 10 m (CSG); 1 s, Ilheu Santana, Baia de Santo Antonio, 10 m (CSG); 1 sp.
Baia das Agulhas, 10 m (CSG). Sao Tomé Island:
10 m (MHNS).
Description : See Gofas & Fernandes
(1988). The soft parts from specimens of
Principe have a transparent foot with
milk whíte blotches (Fig. 3E).
Distribution : Sao Tomé and Principe Is.
Remarks: The radula of this species
was presented in Fernandes &
s, Ilheu Santana, 21 m (CSG); 35 s, Esprainha, 2~
Rolán (1992: fig. 1A). This species
was described as having a shell with
three fading darker bands. In the
material from Principe, these bands
are more marked (Figs. 3 A-D). Some
shells were totally cream without
bands.
5
Iberus , 32 (1), 2014
Genus Marginella Lamarck, 1799
Marginella helmatina Rang, 1832 (Figure 4A-I)
Marginella helmatina Rang, 1832: [Type locality: see Remarks: Principe Island?].
Type material: In MNHN (0530) a lot of 4 syntypes, one of which is represented on the institutional
website chttp:/ / coldb.mnhn.fr>
Other material examined: Principe Island: 1 sp (Fig. 4), 1 s, 1 j, Ponta da Mina, (CSG); 3 sp, Ponta
da Mina, 6 m, Santo Antonio (CCS, CJR); 1 s, Ponta Cabra (CSG); 2 s, Ponta Marmita, 13 m (CCS,
CJR); 12 sp, Baia das Agulhas, 3-7 m (CSG). Gabon: 1 sp, Port Gentil, 20-25 m (CSG).
Description : See Rang (1832). The soft
parís (dorsum of the foot, siphon and
tentacles) are translucent with numerous
milk-white and some orange spots.
Dimensions: Up to 26 mm.
Habitat : This species prefers calm
water and sandy bottoms, between 3
and 13 m.
Distribution : This species is known
from Principe Island and there are
records from Gabon (Bernard, 1984) and
from Senegal.
Remarks : Rang (1832) wrote after the
description of this species: "Nous avons
trouvé la Marginelle helmatine sur la cote
d' Afrique, depuis Vembouchure de la
Gambie jusqu'aux Bisagots". However the
label of the MNHN specimen reads "He
de Prince". In this situation we consider
that the syntype represented would be
more similar to the population here
studied, whereas the material seen from
Senegal: (http: / / www.femorale.com.
br / shellphotos / detail.asp?species=Mar-
ginella+helmatina+Rang%2C+1832) is
evidently rather different morphologi-
cally and it could be a different species,
or a population with different characters
(wider and thicker shell than the popu¬
lation from Principe). The shells of the
population mentioned from Gabon
(Bernard, 1984 and Fig. 41) are also quite
different and their relationship with the
insular population is still to be studied.
Marginella gemina Adams, 1850 (Figures 5A-0, 6A-H)
Marginella gemma Adams, 1850. [Type locality: "East Africa"].
Marginella gorii Cossignani, 2012. Malacologia Mostra Mondiale, 75: 29. [Type locality: Principe
Island].
Type material: Holotype of Marginella gemma Adams, 1850 in (NHMUK 1961135)(Gofas & Fer-
n andes, 1988: fig. 31a, 31b); holotype oí Marginella gorii in MMM (Cossignani, 2012, figs. without
number).
Other material examined: Principe I.: 5 sp, Tinhosa Grande, 15 m (CSG); 1 s, Ponta da Graga, 22
m, under rocks (CSG); 3 sp, 1 s, Ponta da Graga, 22 m (CSG); 1 sp, 1 s, Pedra Galé, 10 m (CSG); 13
sp, Pedra Galé, 35 m (CSG); 2 sp, Pedra Galé, 10 m (CSG); 3 sp, Ilheu dos Mosteiros,20 m (CSG); 1
sp, Tinhosa Grande, 15 m (CSG); 1 sp, 1 j, Tinhosa Grande, 12 m (CSG); 4 sp, 1 s, Tinhosa Grande,
15 m (CSG); 3 sp, Tinhosa Pequeña, 42 m (CSG); 1 sp, Sete Brazas, 10 m (CSG); 12 sp, Sete Brazas,
26 m (CSG); 4 sp, 17 m (CSG); 2 sp, Pedra do Kaki, 16 m (CSG); 2 sp, Tinhosa Pequeña, 25 m (CSG);
11 sp, 1 s, 2 j, Bonné de Joquei, 15 m (CSG); 2 sp, Ilheu Santana, Santo Antonio, 10 m (CSG).
Description : This is a very variable
species in size and mainly in pattern
colouration: In Figures SA-O, we repre-
sent 15 specimens that are very differ¬
ent, and between these specimens there
are intergradations so that they consti-
tute forms of one single species.
Soft parts: the animal is almost
transparent with very fine milk-white
small spots everywhere (foot, siphon
and tentacles) (Figs. 6G-H).
Dimensions: The holotype of M.
gemma is 11.6 mm. The holotype of M.
gorii is 11.25 mm. The larger shells col-
ó
RoláN & GORI: New Information on the marginellids of Sao Tomé and Principe
Figure 4. Marginella helmatina Rang, 1832. A-G: Living specimens on dark background, Ponta da
Mina, Santo Antonio, Principe Is; H: shell, 19.2 mm, Ponta Cabra, Baia de San Antonio, 10 m
(CSG); I: shell, 24.0 mm, Port Gentil, Gabon, 20-25 m (CSG).
Figura 4. Marginella helmatina Rang, 1832. A-G: animales vivos sobre fondo oscuro, Ponta da Mina,
Santo Antonio, Isla del Principe; H: concha, 19,2 mm, Ponta Cabra, Baia de San Antonio , 10 m
(CSG); I: concha, 24,0 mm, Port Gentil, Gabon, 20-25 m (CSG)
7
Iberus , 32 (1), 2014
Figure 5 A-O. Marginella gemma Adams, 1850. A, B: shell, 11.3 mm, Pedra Galé, 10; C-E: shells,
14.3, 14.3, 13.4 mm, Tinhosa Grande, 15 m; F: shell, 12.5 mm, Pedra Galé, 35 m; G, H: shells,
12.2 and 13.3 mm, Mosteiros Fora, 20 m; I, J: shell, 11.6 mm, Tinhosa Grande, 15 m: K: 12.6
mm, Sete Brazas, 20 m; L-O: shells, 12.6, 12.6, 13.7, 13.7 mm, Pedra Galé; all from Principe
island (CSG).
Figura 5 A-O. Marginella gemma Adams, 1850. A, B: concha , 11,3 mm, Pedra Galé, 10; C
conchas, 14,3; 14,3; 13,4 mm, Tinhosa Grande, 15 m; F: concha, 12,5 mm, Pedra Galé, 35 m; G,
conchas, 12,2 y 13,3 mm, Mosteiros Fora, 20 m; I, J: concha, 11,6 mm, Tinhosa Grande, 15 m:
12,6 mm, Sete Brazas, 20 m; L-O: conchas, 12,6; 12,6; 13,7; 13,7 mm, Pedra Galé; todas de isla
Principe ( CSG).
8
I: ^
ROLÁN & GORI: New informatíon on the margínellids of Sao Tomé and Principe
lected reach up to 14.5 mm, but some
populations can ha ve smaller shells.
Habitat and distribution : The species
was collected only in Principe Island in
many localities, under rocks, on rocky
bottoms. We noted that the shells inside
Baia de Santo Antonio (with calm water)
are smaller and less colourful. The
largest and more colourful specimens
were found in the small islet outside the
bay, where the water is rougher.
Remarks: Gofas & Fernandes (1988:
fig. 31a, 31b) represented the holotype of
Marginella gemma Adams, 1850 (in
BMNH 1961135). These figures are rather
similar to our Figure 5I-J, which represent
a form with light colour but only a varia-
tion. The shell shown by Fernandes &
Rolán (1992, fig. 2) has a different
pattern more similar to our Figure 5E.
Cossignani (2012) described the
species here studied as Marginella gorii,
the holotype and paratypes being rather
similar to our Figures 5L O. This shows
that an overview of the variability of a
species is needed before describing a
new one based on only a few specimens.
This species is larger than M. spinacia
Gofas & Fernandes, 1988, and has a
wider shell.
Marginella cf. melvilli Tomlin & Shackleford, 1913 (Figure 6I-M)
Marginella melvilli Tomlin & Shackleford, 1913: 11, pl. 1, figs 1-2. [Type locality: Sao Tomé island].
IMarginella liparozona Tomlin & Shackleford, 1913: 43. [Type locality: Sao Tomé island].
Type material: M. melvilli, holotype in NMW (1955-158-1094), represented in Gofas & Fernandes
(1988: figs. 4a, 4b, 5a, 5b); M. liparozona, lectotype in NWM (1955-158-1093) (Fig. 61), also repre¬
sented in Gofas & Fernandes (1988: figs. 6a, 6b).
Other material examined: Principe: 10 sp, Ponta das Burras, 2-5 m (CSG); 1 s, Ponta Viro Viro, inside
of the Baia de Santo Antonio, 01°38'406"N, Q7°27'329"E, 16 m (CSG); 4 sp, Ilheu Bom Bom, 10 m
(CSG).
Description : See Tomlim & Shackle¬
ford (1913) and Gofas & Fernandes
(1988).
Remarks : These taxa (M. melvilli and
M. liparozona ) are admitted by Gofas &
Fernandes (1988) as possible synonyms
because the difference of the pink bands
is not important in this group.
The material collected and presented
in this work has more colour and a more
marked pattern than the holotype, but
the latter could be from a light form
(like M. liparozona ). At present the status
of these taxa awaits more information
and provisionally we keep them as dif¬
ferent from M. gemma.
Marginella chalmersi Tomlin & Shackleford, 1912 (Figure 7 A-K)
Marginella chalmersi Tomlin & Shackleford, 1912: 320, pl.4, figs 3-4. [Type locality: Sao Tomé].
Type material: Holotype (Figs. 7B-C) in NHMUK (7124).
Other material examined: Sao Tome: 12 sp, Sete Pedras, 10 m (CSG); 2 sp, Lagoa Azul-Morro Car-
regado, 12 m (CSG); 55 sp, Lagoa Azul "Fundáo" at 15-36 m, collected during the night (CSG).
Description : See Tomlin & Shackle¬
ford (1912) and Gofas & Fernandes
(1988). The new population found is as
follows: Shell ovoid elongate, robust,
smooth and shiny, with a high spire and
rounded apex. Suture indistinct. Aperture
narro w and elongate, larger than 1/2 of
the height size. Outer lip thickened, with
about 10-14 denticles inside. The coloura-
tion is cream-white and dark brown: two
bands of this last colour are visible on the
last whorl: the upper one is larger, sepa¬
ra ted from the suture by a light band, and
it is formed by two parts, the upper one
9
Iherus , 32 (1), 2014
Figure 6. A-F: Marginella gemma Adams, 1850. A, B: shell with a similar shape to the holotype, 10.9
mm, Ponta das Burras, Sao Tomé (CSG). C, D: shells with a dubious pattern from Pedra Galé, 13.1
and 1 1.5 mm (CSG); E, F: shell with typical light pattern, 14.8 and 12.4 mm, Bonné de Joquei,
Principe (CSG). G, H: live collected specimens, showing their soft parts. I-M: Marginella cf. melvilli
Tomlin & Shackleford, 1913; I: lectotype of M. liparozona , 7.4 mm (NMW); J-M: shells considered
as forms of M. melvilli , 1 1.0, 1 1.0, 11.1, 10.1 mm, Ilheu Bombom, Principe (CSG).
Figura 6 A-F: Marginella gemma Adams, 1850. A, B: concha con forma similar al holotipo, 10,9 mm,
Ponta das Burras, Sao Tomé ( CSG). C, D: conchas con un patrón de color dudoso, de Pedra Galé, 13, 1 y
11,5 mm (CSG); E, F: concha con patrón claro típico, 14,8 y 12,4 mm, Bonné de Joquei, Principe
( CSG). G, H: ejemplares recolectados vivos, mostrando sus partes blandas. I-M: Marginella cf. melvilli
Tomlin & Shackleford, 1913; I: lectotipo de M. liparozona, 7,4 mm (NMW); J-M: conchas conside¬
radas como formas de M. melvilli, 11,0; 11,0; 11,1; 10,1 mm, Ilheu Bombom, Principe (CSG).
10
ROLÁN & GORI: New information on the marginellids of Sao Tomé and Principe
Figure 7. Marginella chalmersi Tomlin & Shackleford, 1912. A, B: holotype (NHMUK 7124); C-
F: shells, 5.9, 5.5, 6.1, 5.1 mm, from Sete Pedras, Sao Tome, 10 m (CSG); G-K: shells 6.9, 7.8,
7.8, 7.5, 7.9 mm, Lagoa Azul “Fundao”, Sao Tomé (CSG).
Figura 7. Marginella chalmersi Tomlin & Shackleford ' 1912. A, B: holotipo (NHMUK 7124); C-F:
conchas, 5,9; 5,5; 6,1; 5,1 mm, de Sete Pedras, Sao Tome, 10 m (CSG); G-K: conchas 6,9; 7,8; 7,8;
7,5; 7,9 mm, Lagoa Azul “Fundao”, Sao Tomé (CSG).
with numerous axial irregular lines; the
lower one formed by large isolated
blotches, usually with oblique extensions
and sometimos with two of them fused;
the lower dark band is present around the
middle of the last whorls and is almost
formed by axial irregular lines. The spaces
between them are white with cream •
orange lines or small, very variable
blotches.
The soft parts, described by Gofas &
Fern andes (1988: pl. la) are whitish.
Dimensions: the holotype is 6 mm,
and the population has a similar size;
larger specimens of the new found po¬
pulation reach 8.0 mm.
Remarks: The species is represented
and described in Gofas & Fernandes
(1988). This description is coincident
with the original description and the
holotype. The population from Lagoa
Azul has shells which are a little larger
and lighter in colour but probably
within the variability of the species.
Marginella spinacia Gofas & Fernandos, 1988 (Figure 8 A-M)
Marginella spinacia Gofas & Fernandes, 1988: [Type locaiity: Esprahinha].
Type material: Holotype in MNHN (represented in Gofas & Fernandes, 1988: figs. 2a, 2b); paratypes
in many collections.
11
Iberus , 32 (1), 2014
Other material examined: Sao Tomé: 120 sp, 10-15 m, Lagoa Azul-Morro Carregado (CSG); 107 sp,
Esprainha, 2-10 m (MHNS); 87 sp, Ponta de Diogo Vaz, 12 m (CSG); 56 sp, Lagoa Azul-Morro Car-
regado, 11 m (CSG); 12 sp, Lagoa Azul, South side, 12 m (CSG); 30 sp, 8 s, Kia Reef, 15 m (CSG); 14
sp, Baia Ana Chaves, 22 m (CSG); 2 sp, Pedra Conchas, 20 m (CSG); 10 sp, Sao Tomé Capital, 10 m
(CSG); 10 sp, 20 s, Lagoa Azul "Fundao", 30 m (SC); 10 sp, 25 s, Lagoa Azul "Fundao", 36 m (CSG);
1 sp, Ilheu Santana, 20 m (CSG); 7 sp, 8 s, Minerio 43 m (CSG); 250 sp, Lagoa Azul-Morro Car-
regado, 8-12 m (MHNS). Principe: 75 sp, Baia das Agulhas, 10 m (CSG).
Description : See Gofas & Fernandes
(1988).
Dimensions: Usually between 7.0-8.0
mm, ordy exceptionally the largest
shells reach up to: 11.2 mm
Remarks : This species was described
from Sao Tomé (Gofas & Fernandes,
1988) and it was presented as a dark
shell, with typical pattern and scarce
variability, and collected between 2-5 m
in depth. In recent collections by the
second author (SG) the species was
found down to 30 m, the deep popula-
tions being lighter in colour (Figures
7G-K). It was collected also from
Principe Island recently in larger quan-
tity (Figures 8M). This species was
recorded in this island for the first time
by Rolán & Fernandes (1992).
We show here the high variability of
this species, so that those who are prone
to describe new species with such great
ease understand that it is preferable to
know better the variability of popula-
tions before creating new and needless
taxa erroneously.
Marginella michelae Cossignani, 2012 (Figure 9 A-J)
Marginella michelae Cossignani, 2012. Malacologia, 75: 30. [Type locality: "Isola de Principe, Sao
Tomé, Tennosa Pequeña"].
Type material: Holotype in MMM, Cupra Marittima.
Other material examined: Sao Tomé: 3 sp, 1 s, East Sete Pedras, on spheroid concretions of the
coralline alga Lithothamnion coralliodes, 30 m (CSG).
Description : See Cossignani (2012).
The soft parts are milk white;under
magnification it can be seen that the
background is a little transparent and
there are many very small milk white
dots.
Distribution and remarks: Cossignani
(2012) described this species based on
scarce material. Furthermore, the
description of Marginella michelae has an
important error, because the author
indicated as type locality "Isola di
Principe, Sao Tomé, Tennosa (sic)
Pequeña". The second author of the
present work (SG) has dived extensively
in Principe, in Sao Tomé and in Tinhosa
Pequeña (this is the exact ñame). The
present species has been found only in a
small area around Sete Pedras, South-
east of Sao Tomé Island, at 27-30 m, col-
lected on spheroid concretions of the
coralline alga Lithothamnion coralliodes.
This must be considered the real type
locality.
Marginella colombi Cossignani, 2012 (Figures 10 A-L, 11 A-K)
Marginella colombi Cossignani, 2012: 31, figs. without number. [Type locality: stated as "Isola de
Principe, Sao Tomé, Tennosa Pequeña", see remarks].
Type material: Holotype in MMM.
Other material examined: Principe Island: 10 sp, Ponta Cabra, Baia de Santo Antonio, 12 m (CSG);
2 sp, Sete Brazas, 21 m (CSG); 10 sp, 1 j, Tinhosa Grande, 15 m (CSG); 6 sp, Tinhosa Grande, 16 m
(CSG); 6 sp, 2 s, Tinhosa Grande, 20 m (CSG); 5 sp, 2 s, Tinhosa Grande, 12 m (CSG); 1 sp, Tinhosa
12
ROLÁN & GORI: New Information on the marginellids of Sao Tomé and Principe
Figure 8. Marginelía spinacia Gofas & Fernaedes, 1988. A, B: 7.0 and 7.1 mm, typical pattern»
Lagoa Azul, 15 m (GSG); C, D: 7.2, 8.1 mm, light pattern, Lagoa Azul, 15 m (CSG); E-L: 9.0,
10.2, 8.5, 8.6, 8.8, 8.3, 9.0 mm, variable patterns, Lagoa Azul, 28 m (CSG); M: lot of shells from
Principe, Baia das Agulhas (CSG).
Figura 8. Marginelía spinacia Gofas & Fernandez, 1988. A , B: 7,0 y 7,1 mm, patrón de color típico,
Lagoa Tlzul, 15 m (CSG); C, D: 7,2 y 8,1 mm, patrón claro, Lagoa Azul, 15 m (CSG); E-L: 9,0;
10,2; 8,5; 8,6; 8,8; 8,3; 9,0 mm, patrones variables, Lagoa Azul, 28 m (CSG); M: lote de conchas de
Principe, Baia das Agulhas ( CSG).
13
Iberus , 32 (1), 2014
Figure 9. Marginella michelae Cossignani, 2012. A-H: shells. A, B: 10.3 mm; C, D: 9.6 mm; E, F:
9.6 mm; G: 9.6 mm; H: 9.7 mm; all from CSG. I: view of the apex; J: specimen of the Figures E
and F, with soft parts.
Figura 9. Marginella michelae Cossignani, 2012. A-H: conchas. A, B: 10,3 mm; C, D: 9,6 mm; E, F:
9.6 mm; G: 9,6 mm; H: 9,7 mm; todas de CSG. I: vista apical; J: ejemplar de las Figuras E y F, con
partes blandas.
14
ROLÁN & GORI: New information on the marginellids of Sao Tomé and Principe
Figure 10. Marginella coiombi Cossignani, 2012, shells with variable pattern. A: 6.3 mm (CSG);
B-C: 6.5 mm; D-E: 6.5 mm; F-G: 6.6 mm; H: 6.5 mm: all from Sete Pedras, Principe (CSG); I-J:
shell, 6.5 mm, Sete Brazas, Principe (CSG); K-L: shells, 6.3 mm and 6.6 mm, Ponta da Gra$a,
Principe Island (CSG).
Figura 10. Marginella coiombi Cossignani, 2012, conchas con patrón de color variable. A: 6,3 mm
( CSG); B-C: 6,5 mm; D-E: 6,5 mm; F-G: 6,6 mm; H: 6,5 mm: todas de Sete Pedras, Principe (CSG);
I-J: concha, 6,5 mm, Sete Brazas, Principe ( CSG); K-L: conchas, 6,3 mm y 6,6 mm, Ponta da Graga,
isla de Principe ( CSG).
15
Iberus, 32 (1), 2014
Grande, 25 m (CSG); 1 sp, Tinhosa Grande, 25 m (CSG); 10 sp, Sete Brazas, 20 m (CSG); 6 sp, Sete
Brazas, 21 m (CSG); 7 sp, Sete Brazas, 20 m (CSG); 9 sp, west Sete Pedras, 18 m (CSG); 8 sp, Pedra
Galé, 35 m (CSG); 36 sp, Pedra do Kaki, 16 m (CSG); 3 sp, off Ilheu dos Mosteiros, 20 m (CSG); 1
sp, Ponta da Graga, 16 m (CSG); 2 sp, Ponta da Graga, 17 m (CSG); 1 sp, Praia Evora, Santo Antonio,
7 m (CSG); 2 sp, Praia Evora, Santo Antonio, 9 m (CSG); 1 sp, west Tinhosa Pequeña, 42 m (CSG);
2 sp, Tinhosa Pequeña, 25 m (CSG); 6 sp, Tinhosa Pequeña, 20 m (CSG); 4 sp, Bonné de Joquei, 12
m (CSG); 3 sp, 10 m (CSG); 1 sp, Pedra Galé, 10 m (CSG); 3 sp, Pedra Galé, 35 m (CSG); 1 sp, 1 s,
Ilheu Santana, Santo Antonio, 10 m (CSG); 32 sp, 3 j, Baia das Agulhas, 10 m (CSG).
Remarks : This is one of the species
we had under description and was pre-
empted by the paper of Cossignani
(2012) based on only three shells. The
type locality must be corrected to the
exact ñame Tinhosa Pequeña.
Cossignani (2012) indicated as type
locality "Isola di Principe, Sao Tomé,
Tennosa (sic) Pequeña", making the
same mistake as for M. michelae. Mar-
ginella colombi has never been found in
Sao Tomé, but only in Principe and the
small islets around.
We have found two specimens that
are completely white.
The variability of the shell can be
shown in the following Figures 10 and
11.
Marginell a britoi spec. nov. (Figure 12 A-K)
Marginella festiva Kiener: Sowerby, 1847: 382, pl.75, figs. 72-73.
Type material: Holotype (Figs. 12A-B) MNHN (26624, sp); paratypes: MNCN (15.05/60086, 1 sp,
Figs. 12C-D); MPíNS (100603, 1 sp, Fig. 12E); CSG (4 sp); all from the type locality. Other paratypes:
CSG (1 sp, Lagoa Azul "Fundao", 26 m, Figs. 12G-H) (1 sp, Pedra Branca, 30 m); CFD (1 sp. type
locality; CPR (1 sp, Lagoa Azul "Fundao", 21 m); CFB (1 sp, Lagoa Azul "Fundao", 36 m); CSG (1
sp, Lagoa Azul "Fundao", Sao Tomé, 33 m, Fig. 121); CJR (1 sp, Lagoa Azul "Fundao", 30 m.
Other material studied: Sao Tomé: 2 s, eroded, from Type locality (CSG); 2 s, 1 s eroded, Lagoa
Azul "Fundao", 30-34 m (CSG). Principe: 1 j, Tinhosa Pequeña, 20 m (CSG).
Type locality: Ponta de Diogo Vaz, 12-21 m, 0Q°19T59"N, 06°29'633"E, Sao Tomé Island.
Etymology: The specific ñame is after the native diver Eider "Dede" Brito, who helped during
diving collections.
Description : Shell elongate with
máximum dimensions varying in differ-
ent populations, solid, shiny with a
prominent spire. The colour is dark
except for one band in the middle of the
last whorl which has irregular borders
and may be variable in width; the dark
part of the shell has brown and red
alternating, this last colour in subsutural
position. Darker lines reach the thicken-
ing of the external lip which is whitish
or pink-cream and form there red-
brown spots, which are more numerous
on the anterior part.
Aperture narrow, a little wider in its
lower part; columella with four plaits;
external lip thickened and minutely
denticulate interiorly (15-18 teeth).
Soft parts: animal totally white, with
small milk white spots and some very
small grey ones, and some elongate yel-
lowish stripes on the caudal part.
Dimensions: The holotype is 12.2
mm. One shell reaches 20.2 mm. Other
shells from Lagoa Azul and from
Principe, are smaller than 10 mm.
Distribution: Known from Sao Tomé
Island; few shells with similar pattern
were coilected in Principe I., in Tinhosa
Pequeña.
Habitat : The specimens from the type
locality were coilected under large
stones, in 12-24 m. Flowever those from
Lagoa Azul were coilected washing the
dead coral Tubastrea aurea (Quoy &
Gaimard, 1830) from deeper water.
Remarks: We consider populations
from the type locality (Ponta de Diogo
Vaz, Sao Tomé) and those from Lagoa
Azul (Sao Tome) and Tinhosa Pequeña
16
ROLÁN & GORI: New Information on the marginellids of Sao Tomé and Principe
Figure 11. Marginella colomhi Cossignani, 2012, shells with variable pattern. A, B: 6.2 and 6.7
mm, Sete Brazas, Principe (CSG); C-E: shells, 5.7, 5.5, 5.6 mm, Sete Brazas (CSG); F, G: shells,
8.1 and 7.4 mm, Tinhosa Grande, Principe (CSG); H: shell, 6.1 mm, Ponta da Gra£a, Principe
(CSG); I: shell, 5.8 mm, Tinhosa Grande, Principe (CSG); J, K: shell, 8.8 mm, Bonné de Joquei,
Principe (CSG).
Figura 11. Marginella colombi Cossignani ', 2012, conchas con patrón de color variable. A, B: 6,2 y
6,7 mm, Sete Brazas, Principe (CSG); C-E: conchas, 5,7; 5,5; 5,6 mm, Sete Brazas (CSG); F, G:
conchas, 8,1 y 7,4 mm, Tinhosa Grande, Principe (CSG); H: concha, 6,1 mm, Ponta da Graga,
Principe (CSG); I: concha, 5,8 mm, Tinhosa Grande, Principe (CSG); J, K: concha, 8,8 mm, Bonné de
Joquei, Principe ( CSG).
17
IberuSy 32 (1), 2014
(Principe), to be conspecific, notwith-
standing the fact that the latter are evi-
dently smaller. They are equal in other
characters.
The differences of M. britoi with
other known species from Sao Tome and
Principe Islands are found in the shell
characters and the colon i of the soft
parts:
- Margineila gemma Adama, 1850 is
very variable in pattern and colouration:
most of their shell, s have 2 dark spiral
bands bordered by narrower líghter
bands which occasionally can be wider.
M. britoi has a red iine between the
bands and two more on the last whorl,
A white band is almost usual between
the dark ones, M. gemma can also have
wider cream or white areas but it lacks
red spiral bands. Margineila britoi is reía -
tívely narrower in shape.
- Margineila chalmersi Tomlín &
Shackleford., 1912 has a shell with very
typical axial irregular lines ín 2-3 bands.
The animal is whitish with yellow spots
ín the tentacles.
- Margineila spinacia Gofas & Fernan¬
dos,. 1988, is also dark, but smaller ín
size, reaching up to 8 mm, the pattern
presents numerous spiral lines of
minute spots, and the soft parts show
numerous yellow and wine coloured
oval spots.
- Margineila michelae Cossígnani,
2012 has a shell with a deep red colour
and is more bíconicaL
Margineila tyermani Marra!. 1876 (Figure 13 A-H)
Margineila tyermani Marrat, 1876: [Type localíty: Coriseo, Gabon].
Margineila eveleighi Tomlin & Shackleford, 1913: 11, pl. 1, figs 5-6. [Type locality: Sao Tomé
Island].
Type material: Holotype of Margineila tyermani probably in Liverpool Museum: not examined.
Holotype of Margineila eveleighi represented in Gofas & Fernandes (1988, figs. 12a, 12b)
Other material examined: Principe: 1 sp, Ponta Viro Viro, (CSG); 11 sp, Ponta da Mina, Santo
Antonio, 10 m (CSG); 6 sp, Ponta da Mina, Santo Antonio, 6 m (CSG); 5 sp, 1 j, Ilheu dos Mostei-
ros, 16 m (CSG); 1 s. Chímalo and Hospital Velho (CSG). Gabon: 12 sp, 1 j, Pointe Deras, near mouth
Io river, south Boolkomo Estuary, 0.5 m (CCS).
Description: See Marrat (1876). The
animal (Figs. 6E-G) has a transparent
foot with numerous spots and rounded
blotches of mílky-yellowish colour. In
the tentacles there is a line of spots. The
siphon is yellowish and not transparent.
Distribution: Coriseo, Gabon.
Habitat: On fíne mud and muddy
sand, 5-8 m.
Remarles: The most similar species
are M. mirandai and M. hernandezi (see
below).
Margineila mirandai spec. nov. (Figure 14 A-I)
Type material: Holotype (Figs. 14 A - C) in MNHN (26625). Paratypes in the following collections:
CSG (3 sp, Figs. 14D, 14F-G); CCS (1 sp, 1 j), from type locality; CSG (2 sp, 1 j), MHNS (100604, 1
sp, Fig. 14E), NMW (1 sp), CPR (1 s, 1 j), CJR (2 j) all from Praia Campana, NE of Principe, 15 m.
Other material examined: Principe: 2 juv, Ponta da Gra^a, Bala de Santo Antonio, 14 m (CSG); 1
juv, Praia Campanha, NE of the island, 15 m (CSG);
Type locality: Ponta da Graca, 01°37'856"N, 07°27'697"E, Baia de Santo Antonio, on sand, 14 m.
Etymology: The specific ñame is after Alberto Miranda, director of the Center Club Maxell of Sao
Tomé for his help with the diving and collecting.
Description: Shell elongate with background colour and black lines. The
poínted spire, solíd, shiny with a white protoconch is rounded, whitish, not
18
RoláN & GORI: New information on the marginellids of Sao Tomé and Principe
Figure 12. Marginella britoi spec. nov. A, B: holotype, 12.2 mm (MNHN); C, D: paratype, 12.8
mm (MNCN); E, F: paratype, 11.9 mm (MHNS); G, H: paratype, 10.8 mm (CSG), all from
type locality; I: paratype, 9.4 mm, Lagoa Azul, Sao Tomé (CSG); J, K: specimens with soft parts.
Figura 12. Marginella britoi spec. nov. A, B: holotipo, 12,2 mm (MNHN); C, D: paratipo , 12,8 mm
(MNCN); E, F: paratipo, 11,9 mm (MHNS); G, H: paratipo, 10,8 mm (CSG), todos de la localidad
tipo ; I: paratipo, 9,4 mm, Lagoa Azul, Sao Tomé (CSG); J, K: ejemplares con partes blandas.
19
Iberus, 32 (1), 2014
Figure 13. Marginella tyermani Marrat, 1876. A, B: shells, 8.3 and 7.8 mm, from Ponta da Mina,
Sao Tomé (CSG); C, D: Shells, 8.3, 7.3 mm, Gabon (CSG); E-G: living specimens from Ponta da
Mina, Sao Tomé; H: lot of specimens from the same locality (CSG).
Figura 13. Marginella tyermani Marrat , 1876. A, B: shells, 8,3 y 7,8 mm, de Ponta da Mina, Sao
Tomé (CSG); Conchas, C, D: 8,3 y 7,3 mm, Gabon (CSG); E-G: Ejemplares vivos de Ponta da Mina,
Sao Tomé; H: lote de ejemplares de la misma localidad ( CSG).
20
RoláN & GORI: New information on the marginellids of Sao Tomé and Principe
Figure 14: Marginella mirandai spec. nov. A-C: holotype, 6.7 mm (MNHN); D: paratype, 6.1
mm, from Baia da Gra^a (CSG); E: paratype, 5.5 mm (MHNS); F, G: paratype, 6.1 mm (CSG);
H, I: the same specimen with soft parts on dark and light background.
Figure 14: Marginella mirandai spec . nov. A-C: hoiotipo, 6,7 mm (MNHN); D: paratipo, 6,1 mm, de
Baia da Graga (CSG); E: paratipo, 5,5 mm (MHNS); F, G: paratipo, 6,1 mm (CSG); H, I: El mismo
ejemplar con partes blandas ; fotografiado sobre fondo oscuro y claro.
21
Iherus , 32 (1), 2014
clearly separated from the teleoconch.
This has about 3 % whorls, which
increase quickly. The last whorl is very
prominent and represents more than
about 3A of the height. Aperture elon-
gate, a little wider in its lower part;
external lip white ,very widened; col-
umella opisthocline, with 4 separated
and very prominent folds, of which the
three lower ones are very oblique; the
portion without folds represents about
30-40% of its total length. The external
pattern appears grey in general, with 6-7
fine spiral black lines on the last whorl,
and only one on each of the first whorls.
These lines are irregularly separated
(the wider space is usually the subsu¬
tural) and between them appear short
axial and very variable lines or blotches,
frequently inclined in different direc-
tions.
Dimensions: the holotype is 6.7 mm.
The paratypes are slightly smaller.
Soft parts whitish, transparent with
numerous milk white spots and
blotches, those situated anteriorly being
smaller and the lateral and caudal ones
larger. These milk white spots appear
also on the tentacles and are more
numerous on the siphon.
Distrihution : NE of Principe Island.
Remarks : This species at first glance
may show some similarity to M. tyermani,
but the latter species is larger, with a spire
that is more pointed, and more numerous
spiral dark lines. The axial lines and
blotches in M. mirandai are usually limited
to one interspace between spiral lines and
the colour of the blotches on the soft parts
is white instead of yellowish.
For the differences with M. hernan-
dezi spec. nov. see below.
Marginella hernandezi spec. nov. (Figure 15A-K)
Type material: Holotype (Figs. 15A-B) in MNHN (26626). Paratypes: CSG (1 s, Praia das Burras,
18-20 m, Fig. 15D); CSG (3 s, 4 j, Baia das Agulhas, Fig. 15C); CCS (2 s) from the same locality; CSG
(1 s, Fig. 15G); CSG (1 s, Fig. 151), from Ponta da Graga,; MHNS (100605, 1 s, Fig. 15F), CPR (1 sp,
Fig. 15H), and CJR (1 s, Fig. 15E), from Praia das Burras: all from Principe island.
Type locality: Principe I.: Baia das Agulhas, 01°36'06"N, 07°20'55"E, 15 m, on rocky bottom.
Etymology: The specific ñame is after José María Hernández, a Spanish malacologist, who passed
away some time ago, visited Sao Tomé Islands several times and collected there with the authors.
Description : Shell elongate with
pointed spire, solid, shiny with a grey
colour. The protoconch is rounded,
whitish, not clearly separated from the
teleoconch. This has 3 3á whorls, which
increase quickly. The last whorl is very
prominent and represents 3á of the
height. On the dorsum, some shells can
have very short and scarcely prominent
axial ribs, not present in other shells.
Aperture elongate, slightly wider in its
lower part; external lip white, very
widened; columella opisthocline, with 4
separated and very prominent folds
which are very oblique, specially the
lower ones and which take up about
60% of the length. The external pattern
appears grey in general, with fine, spiral
lines of dark grey, one on each of the
first whorls and 10 on the last one, not
regularly separated and crossed by
irregular zigzag axial lines, which form
two light bands with axial blotches.
The animal is transparent with small
white to scarcely yellowish blotches on
the foot, the siphon is light yellow and
the tentacles transparent with small
light yellow dots.
Dimensions: The holotype is 9.4 mm.
The larger paratype is 9.8 mm.
Distribution : This species was found
in Baia das Agulhas, diving, dredged at
Praia das Burras and night diving at
Ponta da Graga, Baia de Santo Antonio.
Habitat: Live material was found
under sand, 3-15 m and under a thin
layer of sand on fíat rocks; also dredged
at 20 m in dark sand mixed with spher-
oid concretions of Lithothamnion coral-
lioides.
Remarks : This new species is differ¬
ent from any other of the islands species
22
RoláN & GORI: New information on the marginellids of Sao Tomé and Principe
Figure 15. Marginella hernandezi spec. nov. A, B: holotype, 9.4 mm (MNHN); C: paratype, 9.3
mm, type locality (CSG); D: paratype, 8.5 mm, off Praia das Burras (CSG); E: paratype, 8.4 mm,
same locality (CJR); F: paratype, 7.4 mm (MHNS); G: paratype, 8.6 mm, Baia das Agulhas
(CSG); H: paratype, 7.4 mm off Praia das Burras (CPR); I: paratype, 9.8 mm, Baia de Santo
Antonio (CSG); J, K: living specimens on dark bottom.
Figure 15. Marginella hernandezi spec. nov. A, B: holotipo, 9,4 mm (MNHN); C: paratipo, 9,3 mm,
de la localidad tipo ( CSG); D: paratipo, 8,5 mm, frente a Praia das Burras (CSG); E: paratipo, 8,4
mm, misma localidad (CJR); F: paratipo, 7,4 mm (MHNS); G: paratipo, 8,6 mm, Baia das Agulhas
(CSG); H: paratipo, 7,4 mm frente a Praia das Burras (CPR); I: paratipo, 9,8 mm, Baia de Santo
Antonio ( CSG); J, K: ejemplares vivos sobre fondo oscuro.
23
Iherus , 32 (1), 2014
by its gray colour pattern, its wider
aperture, wider thickened outer lip and
prominent columellar folds. Some shells
lack spiral lines, but we consider that
they are the same species ,being equal in
other characters.
Marginella tyermani Marrat, 1876 from
Congo, Gabon and Sao Tomé has some
remóte similarity but this latter species
has a shell which is strongly ribbed, with
a greenish-yellowish colour, is more
solid, relatively wider, the apex is wider,
the columellar folds larger, the spiral
lines are more marked, and the fine axial
zigzag lines are almost always wanting.
The animal is similar but the blotches on
the foot are larger and more yellowish in
M. tyermani.
CONCLUSIONS
For the present work approximately
1500 specimens and shells of marginel-
lids from Sao Tomé and Principe were
examined, and several trips to both
islands were necessary for collecting
material, photographs and information.
In the present work 4 new species are
described, giving information on others
previously known.
It seems evident that the West
African coast is a paradise for the
genera Marginella , Volvarina , Granulina,
etc. with numerous endemic species
living within short distances. Of these
ACKNOWLEDGEMENTS
The optical photographs were made
in the Department of Genetics of the
University of Vigo. The SEM photo of
Plesiocystiscus violáceas was made in the
Centro de Apoyo Científico y Tec¬
nológico a la Investigación (CACTI) in
the same University by Jesús Méndez.
Some photographs of living animáis
were made by the second author after
the collecting.
The second author thanks all the
staff of Club Maxel diving center of Sao
Tomé: Jean Louis Testori and Nicolás
Cartrom (the oíd and the new owner).
M. mirandai spec. nov. is smaller, it
has about 6 more marked spiral lines on
the last whorl instead of those 10 fine
lines in M. hernandezi; furthermore the
pattern is different: M. hernandezi has ir¬
regular zigzag with axial lines more
marked than the spiral ones, that are pro-
longed over the middle of the whorls,
sometimes covering the complete shell;
on the contrary, the pattern of M. mi¬
randai has more predominant spiral lines,
while the axial lines are not in zigzag but
frequently shorter, rarely Crossing the spi¬
ral ones. Both species live sympatrically.
Marginella fumígala Gofas & Fernan-
des, 1994, from Angola, is larger (13-16
mm), the two first whorls are smooth and
the subsequent ones show axial ribs.
areas, some have a marked personality,
such as those from Cape Verde Archi-
pelago, Senegal, Ghana, Sao Tomé and
Principe Islands, and Angola, which are
the best recognized. In these areas,
many species have been described
during the last years of the last century
and the beginning of this one and new
papers are informing us continuously
about new records and new species
described. However many Coastal areas
are unexplored even now and so it is
probable that further new species shall
be described in a near future.
Edmilson Augusto and Alberto Miranda
(the oíd and the new director), and the
divers: Helder "Pede" Brito, Eslander
"Danger" Campos, Apolo Pires,
Armando Pires; Lucas Narciso Tabares;
Rodhesio dos Anjos, the motorist Joao
Lopes Antonio. Thanks to José Rosado
and Cedric Simbile for their cooperation
in the research and observations in
Principe and for the photos of the live
animáis. To Peter Ryall for his notes and
to Jennifer Callighan, assistant curator
of Mollusca, National Museum of Wales
for her help and information.
24
ROLÁN & GORI: New Information on the marginellids of Sao Tomé and Principe
BIBLIOGRAPHY
Clover P.W. 1990. Due nuove specie di Mar¬
ginella. La Conchiglia,2 53-255: 21-23.
CossiGNANl T. 2012. Tre nuova marginelle da
Sao Tomé (Gastropoda: Prosobranchia, Mar-
ginellidae). Malacologia, 75: 29-31.
Fernandes F. & Rolán E. 1992. The Margin-
ellidae (Mollusca, Gastropoda) of Principe
Islands (República de Sao Tome e
Principe). Journal of Conchology, 34(2): 85-
90, pl. 1.
Gofas S. & Fernandes F. 1988. The marginel¬
lids of Sao Tomé, West Africa. Journal of Con¬
chology, 33(1): 1-30.
Nobre A. 1886. Exploragáo scientifica da ilha
de S. Thomé: conchas terrestres e marinhas
recolhidas pelo Sr. Adolpho Moller. Boletim
da Sociedad Geographyca, Lisboa , 4 (6): 1-15.
Nobre A. 1887. Remarques sur la faune mala
cologique marine des possesions portugaises
de 1 Afrique Occidentale. Jornal de Sciencias
Mathematicas, Physicas e Naturaes, Lisboa , (46):
107-120.
Kiener L.C. 1834. Specie s General et Iconogra-
phie des Coquilles Vivantes . Columellaires,
Genre Marginelle.
Rang S. 1832. Marginelle. Marginella Lam. M.
helmatine. M. helmatina, Rang. Magasin de
Zoologie, 2: pl. 5 (and unpaginated text).
Reeve L.A., 1866. Conchologia Iconica. Vol.XV,
Marginella.
Tomlin J.R. le B. & Shackleford L.J. 1912.
Bescriptions of two new species oí Marginella
from San Thome island. Journal of Conchology,
13:319-320
Tomlin J.R. le B. & Shackleford L.J. 1913a. De-
scriptions of two new species of Marginella
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14: 11 [pl.l published with 1913b]
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scriptions of new species of Marginella and
Mucronalia from Sao Thomé. Journal of Con¬
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Marginellidae, Olividae, Columbellidae.
25
.
.
© Sociedad Española de Malacolagía
Iberus, 32 (1): 27-34, 2014
Wm
An undescribed species of Cryptosaccus (Gastropoda:
Hygromiidae) from the south- west of the province of León,
NW Spain
Una nueva especie de Cryptosaccus (Gastropoda: Hygromiidae) del
sur de Province León, NO España
David T. HOLYOAK* and Geraldine A. HOLYOAK*
Recibido el 25-VIII-20 13. Aceptado el 4-X1-2013
ABSTRACT
An unknown species of the family Hygromiidae, discovered among calcium-poor quartzite
rocks at high altitude (ca 1 879 m) on a mountain ridge in the Sierra de la Cabrera on the
Southern border of the province of León, is named here as Cryptosaccus cabrerensis. The
sheli characters and genital anatomy show strong similarities to Cryptosaccus asturiensis
Prieto & Puente, 1 994, as well as some clear differences. The new species has the acces-
sory sac adjoining the dart sac visible externally during dissection, unlike C. asturiensis.
Possible affinities of Cryptosaccus with other genera are discussed.
RESUMEN
Se describe una especie desconocida de la familia Hygromiidae, descubierta entre roque¬
dos de cuarcita pobre en calcio, a gran altura (ca 1 879 m) en una montaña en la Sierra
de la Cabrera en la frontera sur de la provincia de León, bajo el nombre Cryptosaccus
cabrerensis. Los caracteres de la concha y la anatomía del aparato genital muestran una
fuerte similitud con Cryptosaccus asturiensis Prieto y Puente, 1 994, así como algunas dife¬
rencias claras. La nueva especie tiene el saco accesorio ¡unto al saco de dardo visible
externamente durante la disección, a diferencia de C. asturiensis. Se discuten las posibles
afinidades de Cryptosaccus con otros géneros.
INTRODUCTION
Cryptosaccus asturiensis was named
in a new monotypic genus by Prieto &
Puente (1994) from rocky limestone
habitats in the Somiedo Valley of Astu¬
rias province, NW. Spain, where it
appears to be a localised endemic. In
July 2013 an unknown species of Hygro¬
miidae was fourid by the authors among
calcium-poor quartzite rocks at high
altitude (ca 1879 m) on a mountain ridge
of the Sierra de la Cabrera on the Sout¬
hern border of León province. This did
not match any of the species listed for
León province or adjoining Zamora pro¬
vince in the comprehensive review by
Hermida (1992). Investigation of its
sheli characters and genital anatomy
have revealed strong similarities to
Cryptosaccus asturiensis as well as some
clear differences, so it is described here
and named as a second species of the
genus, C. cabrerensis.
* Quinta da Cachopa, Barcoila, 6100-014 Cabezudo, Portugal, e-mail: holyoak9187@gmail.com
27
Iherus , 32 (1), 2014
MATERIAL AND METHODS
Grid references of localities were
obtained using hand-held Garmin GPS
devices, in 2007 using a GPS 12 accurate
to <10 m horizontally but considerably
less accurate for altitudinal data, in 2013
using an Etrex High Sensitivity instru-
ment accurate to <5 m both horizontally
and vertically. Living snails were
drowned in water overnight, then pre-
served in 80% Industrial Methylated
Spirit.
Shells of the new species described
below are weakly calcified, so they are
remarkably thin, flexible and fragile.
Henee, many of the empty shells are
broken or distorted and measurement
data are partly incomplete for that
reason. Other shell measurements are
incomplete because they are based on
whole drowned specimens retained in
spirit, with which the presence of the
extended body prevenís measurements
of the aperture and umbilicus. It proved
to be impossible to pulí intact bodies
from the shells of this taxon without
destroying the shell, so that the anato-
mical material studied was limited to
foreparts of four bodies that could be
extracted, in three of them by making a
hole in the underside of the penultimate
TAXONOMIC PART & RESULTS
shell whorl in order to cut through the
body. Other specimens were left intact
in spirit to allow fu ture studies. Extrac-
tion of more of the body from the more
calcified shells of C. asturiensis did not
present any difficulty. Shell descriptions
and dissections were carried out using a
Meiji RZ Series stereo-microscope, dra-
wings were made with assistance from a
Meiji drawing tube, shell photographs
with an Infinityl camera and shell mea¬
surements were made with an eye-piece
graticule. Shell whorls were counted
using the method described by Kerney
& Cameron (1979). Proximal and distal
refer to the position in relation to the
ovotestis.
Abbreviations: AH: máximum
height of shell aperture; AW: máximum
width of shell aperture; B: shell breadth;
CGAH: Collection of G.A. and D.T.
Holyoak; DTH: D.T. Holyoak; GAH:
G.A. Holyoak; H: shell height; IMS:
Industrial Methylated Spirit; MNCN:
Museum Nacional de Ciencias Natura¬
les, Madrid; n: sample size; NHMUK:
The Natural History Museum, London;
U: máximum width of umbilicus. The
map references for localities are based
on the LJ.T.M. grid. The material descri¬
bed is in CGAH, MNCN and NHMUK.
Hygromiidae Tryon, 1866
Cryptosaccus Prieto & Puente, 1994; type species C. asturiensis Prieto & Puente, 1994 by original
designation
Cryptosaccus asturiensis Prieto & Puente, 1994
Cryptosaccus asturiensis Prieto & Puente, 1994, Arch. Molí., 123: 112. [Type locality: 2 km al sur de
Pola de Somiedo, en la base de la pared de un contrafuerte calcáreo. Prieto, Puente, Altonaga
& Gómez-Moliner (2011: 878) pointed out that this was " entre Caunedo y Gúa" and that the
UTM coordinates originally given as 29T QH 234 772 do not represent the type locality].
Material examined: Specimens all from Spain, in CGAH: (a) by AS227 N. of Pola de Somiedo, Prov.
Asturias, 29T QH 232 769, E.-facing limestone crag and slopes, ca 655 m alt., 23 May 2007, GAH &
DTH site E23, 1 dry shell & body in IMS, 13 specimens in IMS; (b) just E. of AS227 at ca 3 km S. of
Santa María del Puerto, León province, 29T QH 261 655, limestone rocks and grassland on W.-
facing slope, ca 1410 m alt., 25 & 26 May 2007, DTH site E27, dry shell & body in IMS, 2 specimens
in IMS, 13 shells (dead when collected).
28
HOLYOAK AND HOLYOAK: An undescribed species of Cryptosaccus from the NW Spain
Figure 1. Shells of Cryptosaccus cabrerensis sp. nov. A-C: holotype (shell breadth 7.8 mm). D-F: paraty-
pes from type locality. D: apical whorls to show microsculpture; E: “scaly” microsculpture on perip-
hery of body whorl near aperture (to left of photograph); F: living animal aestivating on underside of
quartzite boulder (shell breadth ca 7.5 mm).
Figura 1. Conchas de Cryptosaccus cabrerensis spec nov. A-C: holotipo (ancho de la concha 7,8 mm);
D-F: paratipos de la localidad tipo. D: vueltas apicales mostrando la microescultura; E: microescultura
“escamosa” en la periferia de la última vuelta i cerca de la apertura (a la izquierda de la fotografía); F:
animal vivo en estivación en la parte inferior de un bloque de cuarcita (ancho de la concha, unos 7,5
mm).
When the species was named it was
known only from the type locality, from
steep limestone crags. Prieto et al. (2011:
878) added three more localities, all within
the Somiedo Valley, including a limestone
wall and a road cutting. A "Vulnerable"
Red List category is assigned to the species
(Gómez, 2011; Prieto et al., 2011). Our site
(b) listed above apparently represents a sig¬
nifican! extensión to the published data on
the geographical and altitudinal ranges
and recorded habitat of the species and the
first record of it (just) outside the Somiedo
Valley and (just) in León province.
Cryptosaccus cabrerensis G.A. Holyoak & D.T. Holyoak sp. nov.
Type material: Holotype (Figs 1A-C; reg. no. MNCN15.05/60087) from type locality, collected 1
July 2013 by G.A. Holyoak and D.T. Holyoak at site E334; forepart of body in spirit (IMS) and dry
shell kept separately.
Paratypes: All from type locality with same collection data as holotype; 3 shells (dry) and incom¬
plete bodies (foreparts in IMS), 5 specimens in IMS, 18 shells (dead when collected, mainly in poor
condition) in CGAH; 1 specimen in absolute ethanol in NHMUK.
Type locality: ca 1 km W. of pass at Alto del Peñón (Sierra de la Cabrera, León province, Spain), 29T
QG 0178 7555, ca 1879 m alt., unshaded quartzite scree adjoining low scrub high on N.-facing slope.
Etymology: Prieto & Puente (1994: 110) stated that the generic ñame Cryptosaccus was derived
from the greek kryptos = oculto (hidden) and the latin saccus = saco (sac). Our species epiphet cabre¬
rensis is an adjective derived from Sierra de la Cabrera, the massif around the type locality. Saccus
is a masculine noun and the latin suffix -ensis is in agreement with the generic ñame for the epip-
hets cabrerensis and asturiensis.
29
Iberus , 32 (1), 2014
Figure 2. Anatomy of distal genitalia in Cryptosaccus cabrerensis sp. nov. A: holotype; B: partly sche-
matic diagram of penis, vagina and dart sac complex in a paratype (showing longitudinal sections
of dart sac complex and distal two-thirds of penis; the vagina and genital atrium opened and flatte-
ned to illustrate internal ridges). Abbreviations, as: accessory sac; be: bursa copulatrix; dbc: duct of
bursa copulatrix; ds: dart sac; e: epiphallus; f: flagellum; fo: free oviduct; ga: genital atrium; gp:
genital pore; mg: mucus gland; p: penis; prm: penial retractor muscle; ror: right ommatophore
retractor muscle; sod: spermoviduct; v: vagina; vd: vas deferens; ve: verge.
Figura 2. Anatomía de los órganos genitales distales en Cryptosaccus cabrerensis spec. nov. A: holotipo;
B: diagrama semiesquemático del pene, la vagina y complejo del saco del dardo en una paratipo ( mos¬
trando secciones longitudinales de complejo del saco del dardo y de los dos tercios distales de pene; la
vagina y el atrio genital abiertos y aplastados para mostrar las crestas internas). Abreviaturas, as: saco
accesorio; be: bolsa copulatriz; dbc: conducto de la bolsa copulatriz; ds: saco del dardo; e: epifalo; f:
flagelo; fo: oviducto libre; ga: atrio genital; gp: poro genital; mg: glándula del mucus; p: pene; prm:
músculo retractor del pene; ror: músculo retractor del ommatoforo derecho; sod: spermoviduct; v: vagina;
vd: vaso deferente; ve: verga.
Description : Adult shell (Figs 1A-F)
very depressed cortical with flatter apex
and somewhat flattened below. Breadth
at maturity 6. 9-7.9 mm (n = 10), height
3.44.5 mm (n = 9), H/B 0.49-0.61 (n - 9).
Whorls 4.5-4. 8 (n = 9), expanding regu-
larly, rounded at periphery, distinctly
flattened above, with moderately deep
sutures. Umbilicus narrow, máximum
width 0.7-11 mm (n = 4) (U/B 0.10-0.14,
n = 4), deep and symmetrical, exposing
inner edges of uppermost whorls. Aper-
ture rounded-oval except where inte-
rrupted by penultimate whorl, slightly
flattened below, máximum width 3.0-3.8
mm (n = 5), máximum height 2.7-35 mm
(n = 6) (AW/AH 1.10-1.18, n = 5). Aper-
ture lacking any internal rib; peristome
thin throughout, plañe except at umbili¬
cus where narro wly reflected. Shell very
thin, uncalcified, flexible when fresh,
translucent, light brownish-green.
Shell surface with waxy lustre. Pro-
toconch nearly smooth. Whorl 1.0-1. 5
with distinct, regular, spiral ridges (Fig.
ID), which become much less obvious
amongst other sculpture on later
whorls. Irregular low transverse ribs or
30
Holyoak AND HOLYOAK: An undescribed species of Cryptosaccus from the NW Spain
striae (growth ridges) ± prominent from
whorl 1.3 to shell aperture, these
tending to be more strongly developed
on top of shell in narrow band below
each suture. Penultimate and body
whorls with microsculpture on perios-
tracum of closely spaced, shallowly
crescentic transverse ridges giving a
scaly pattern (their sides concave
towards shell aperture: Fig. 1E), arran-
ged in somewhat irregular spiral rows;
these are absent from umbilicus and a
broad zone around it on underside of
shell. Shell hairs apparently lacking, but
small immatures not seen.
Exterior of body with foreparts light
grey above and towards foot-fringe, sides
and solé of foot whitish; mantle whitish
with irregular blotches and large spots of
dark brown, these markings larger and
darker (blackish) near right-hand edge,
forming irregular lines. The dark mar¬
kings are visible on living animáis
through the translucent shell (Fig. 1F).
Genital anatomy (Fig. 2), based on
dissection of distal parts of bodies of
four mature individuáis: Genital pore lo-
cated cióse behind lower tentacle on
right-hand side of forepart of body at
about mid-height. Genital atrium short,
its proximal end of similar width to vagi¬
na, narrowing distally towards genital
pore. Right ommatophore retractor mus-
ele passes between penis and vagina. Pe¬
nis moderately long, cylindrical, with
penial retractor muscle inserted at its
proximal end, the muscle short and wide
to narrower and moderately long (two-
thirds length of epiphallus), attached to
body wall. Interior of penis studied in
one specimen: distal two-thirds of penis
comprised of an outer sheath that joins
genital atrium at its narrower distal end;
most of interior of sheath occupied by
long-conical verge that tapers to a
bluntly pointed tip at distal end, which
has a large pore. Epiphallus long, the
proximal part coiled. Flagellum long (up
to 1.5x length of epiphallus), conical-cy-
lindrical, in three individuáis coiled on
top of epiphallus when in situ , in fourth
individual extended proximally inside
body wall. Vas deferens a thin tube, en-
ding at junction of epiphallus and flage¬
llum, passing distally closely attached
alongside epiphallus and penis to pe-
nial-vaginal angle (beneath right omma¬
tophore retractor muscle), then returning
proximally alongside vagina (beneath
two mucus glands) and free oviduct. Va¬
gina ± cylindrical, of similar length to
penis but often somewhat wider, with
large dart sac complex inserting exter-
nally at point about one-third below its
proximal end. Inner wall of vagina (one
examined) with nine high ridges that
continué distally to middle of vagina; se-
veral ridges coalesce in distal one-third
of vagina, leaving five or six ridges at its
distal end, two of which continué along
interior of outer part of genital atrium.
The dart sac complex clearly a double
structure when viewed externally, with
outer part (dart sac) 2.5x width and of
greater length than inner part (accessory
sac). Dart sac muscular; one examined
internally containing small dart (two
broken white cylindrical pieces seen: ca
0.25 and 0.2 mm long, both of ca 0.05 mm
diameter); narrow central canal of sac
opens into vagina in a longitudinally
elongate pore. The accessory sac not fu-
sed to adj acent wall of vagina for much
of its length and free from dart sac for
about one-half of its length, also muscu¬
lar, with narrow central cavity (empty in
one dissected) that opens as pore into va¬
gina. Six, se ven or eight mucus glands in
four groups insert near proximal end of
vagina; in one individual two groups
each of two pairs arise on opposite sides
of vagina, in other individuáis one or
two of glands not paired and their arran-
gement around vagina rather variable.
All mucus glands short, narrowed at in-
sertion and widening to a swollen bul-
bous proximal end. Free oviduct cylin¬
drical, similar in thickness to vagina but
about one-third its length. Duct of bursa
copulatrix cylindrical, moderately wide
throughout, nearly twice length of vagi¬
na and nearly as wide, coiled in situ. Bur¬
sa copulatrix thin-walled, closely appres-
sed to and resting within curve of sper-
moviduct, in three individuáis D-shaped
with duct inserting obliquely on the
rounded edge, in one individual oval but
tapering into duct.
31
Iberus, 32 (1), 2014
DISCUSSION
C. cabrerensis differs conspicuously
from C. asturiensis in shell characters. Its
shell is smaller (breadth up to 7.9 mm
compared to 10 mm), with fewer whorls
(up to 4.8 compared to 5. 5-6), much
thinner and more translucent, and the
sculpture of projecting triangular scales
on the periphery of the body whorl is
much lower and less developed. Their
genital anatomy is generally similar, dif-
fering most obviously in the dart sac of
C. cabrerensis being accompanied by a
smaller accessory sac that is visible
externally duririg dissection, whereas in
C. asturiensis the accessory sac is not
visible externally, this character having
formed the basis of the generic ñame
Cryptosaccus. Also, the bursa copulatrix
is much more elongate in C. asturiensis
(length >5x width) than in C. cabrerensis
(length l-2x width).
Although the specific distinctness of
C. cabrerensis from C. asturiensis can
hardly be doubted, they nevertheless
appear to be more closely related to
each other than to any other Hygromii-
dae. Thus, they share a shell of similar
general form, narrowly umbilicated
with a simple peristome and with dis-
tinctive scale-like sculpture on the
periphery of the body whorl. Among
characters of the genital anatomy, they
share a single dart sac with a small
accessory sac joined along its inner
edge, six to eight mucus glands inser-
ting proximally to the insertion of the
dart sac arranged approximately as two
pairs each on opposite sides of the
vagina, a long slender epiphallus and a
long slender flagellum. They are also
both rupestral species occurring in adja-
cent regions of NW. Spain, albeit separa-
te d by virtually the full width of León
province (Fig. 3).
Prieto & Puente (1994) pointed out
the considerable biogeographic and eco-
logical similarities between Cryptosaccus
asturiensis and Pyrenaearia spp. However,
they did not emphasise the cióse simila-
rity between them in most characters of
their genital anatomy, perhaps because
Pyrenaearia differs in having the acces¬
sory sac visibly sepárate from the dart sac
externally, often having the mucus
glands inserting closer to the insertion of
the dart sac, in addition to differences in
shell sculpture (lack of scaly microsculp-
ture; juveniles often with shell hairs).
Nevertheless, since C. cabrerensis has an
accessory sac that is visible externally, it
prompts reconsideration of possible affi-
nities with Pyrenaearia. A recent molecu-
lar-phylogenetic study of Pyrenaearia
(Elejalde, Madeira, Prieto, Backeljau
& Gómez-Moliner, 2009) did not consi-
der Cryptosaccus, but its detailed analyti-
cal data should now provide an excellent
basis for future comparisons. Since Zeno-
biella subrufescens (Miller, 1822) also has
similar genital anatomy (Schileyko,
2005: 1961), a "scaly" pattern of shell
microsculpture and a range extending to
N. Spain it should also be compared, alt¬
hough its shell differs in having a very
small umbilicus.
C. cabrerensis is known only from the
type locality, on an exposed, gently N.-
facing montane slope at ca 1879 m alt.,
cióse below a rocky ridge that forms a
major watershed which represents the
border between Province León and Pro¬
vince Zamora to the south (Fig. 3). The
snails were concealed beneath the
surface of an unshaded scree composed
of boulders and large blocks of quartzite
(Fig. 4). Most of them were cióse to the
upper edge of the scree adjacent to
dense low scrub (<0.5 m high), compri-
sed of Genista sanabrensis Valdés Berm.,
Castrov. & Casaseca (the commonest
bush), Vaccinium myrtillus L. and Festuca
sp. with Erica australis L. ca 1 m high
nearby. The flora and rock type indicate
extremely calcium-poor conditions, with
acidic soils.
Living C. cabrerensis occurred at low
density, aestivating in the hot dry sunny
weather mainly on the undersides of
rocks (Fig. 1F); they had a thin membra-
nous and translucent epiphragm inside
the shell aperture. The only other
mollusc we found there was a single
dead shell of Oxychilus cf. alliarius
(Miller, 1822).
32
HOLYOAK AND HOLYOAK: An undescribed species of Cryptosaccus from the NW Spain
v
Figure 3. Map of north-western Spain showing distribution of Cryptosaccus asturiensis and C. cabre-
rensis sp. nov. in ten kilometre squares of the U.T.M. grid. Figure 4. Habitat of Cryptosaccus cabre-
rensis spec. nov. at the type locality.
Figura 3. Mapa del noroeste de España indicando la distribución de Cryptosaccus asturiensis y de C.
cabrerensis sp. nov. en cuadrículas U.T.M. de 10 km. Figura 4. Hábitat de Cryptosaccus cabrerensis
spec. nov. en la localidad tipo.
33
Iberus , 32 (1), 2014
Discovery of an undescribed species of
Cryptosaccus among acidic rocks at such
high elevation was unexpected. More
widespread searching in the Sierra de la
Cabrera and beyond will be needed to
establish its range and conservation status.
BIBLIOGRAPHY
Elejalde M.A., Madeira M.J., Prieto CE.,
Backeljau T. & Gómez-Moliner B.J. 2009.
Molecular phylogeny, taxonomy, and evo-
lution of the land snail genus Pyrenaearia
(Gastropoda, Helicoidea). American Malaco-
logical Bulletin, 27 (1/2): 69-81.
Gómez B. 2011. Cryptosaccus asturiensis. In:
IUCN 2013. IUCN Red List of Threatened
Species. Versión 2013.1. <www.iucn. red-
list.org>. Downloaded 21 August 2013.
Hermida J. 1992. Estudios faunísticos y ecológi¬
cos de los moluscos gasterópodos terrestres de
Asturias , León , Zamora y Salamanca. Unpu-
blished doctoral thesis, Departamento de
Bioloxia Animal, Universidade de Santiago
de Compostela. 302 pp.
Kerney M.P. & Cameron R.A.D. 1979. Land
snails ofBritain & north-west Europe. London:
HarperCollins. 288 pp., 24 pls.
ACKNOWLEDGEMENTS
Thanks are due to Rui da Costa
Mendes for help with literature. Fig. 3
was prepared using the DMAP software
written by Dr A.J. Morton.
Prieto C.E. & Puente A.1. 1994. Un nuevo Hy-
gromiinae del Norte de la Península Ibérica,
Cryptosaccus asturiensis n. gen., n. sp. Archiv
für Molluskenkunde, 123 (1/6): 109-122.
Prieto C.E., Puente A.I., Altonaga K. & Gó-
mez-Moliner B.J. 2011. Cryptosaccus astu¬
riensis Prieto y Puente, 1994. In: Verdú J.R.
Atlas y Libro Rojo de los invertebrados amena¬
zadas de España. Especies Vulnerables. Volu¬
men 2: Moluscos. Madrid: Editorial Ministe¬
rio de Medio Ambiente Rural y Marino, pp.
877-880.
Schileyko A.A. 2005. Treatise on recent te-
rrestrial pulmonate molluscs. Part 14. Heli-
codontidae, Ciliellidae, Hygromiidae. Rut-
henica, Suppl., Moscow, 2: 1907-2047.
34
© Sociedad Española de Malacología
Iberas , 32 (1): 33-43, 2014
'FffíjZs
Estado de conservación de la náyade Margaritifera
margaritifera (Linnaeus 1758) en el curso bajo del río
Mandeo (A Coruña)
Status of the freshwater pearl mussel Margaritifera margaritifera
(Linnaeus 1758) in the lower course of the river Mandeo (A Coruña)
Alvaro BARROS*
Recibido el 2-VII-20 13. Aceptado el 13-1-2014
RESUMEN
Se estudian las densidades, la estructura por tamaños y la clasificación por edades de una
población de la náyade Margaritifera margaritifera en el curso bajo del río Mandeo (A Coruña).
Se seleccionaron tres parcelas de estudio en el año 2000 que fueron nuevamente estudiadas
en el año 2010, año en el que además se añadió una cuarta parcela. Las densidades no
difirieron de manera significativa entre los dos años de control, mientras que la distribución
por tamaños mostró un claro dominio de los ejemplares adultos en todas las parcelas en los
dos años considerados. En la parcela controlada únicamente en 2010 se registró una talla
media superior a la de las otras zonas estudiadas, lo que unido al hecho de que sólo se
hayan localizado animales jóvenes en puntos muy concretos del río, podría indicar una
distribución no homogénea de los distintos grupos de edad. Aunque la fracción mayoritaria
de la población está constituida por animales de entre 2 1 y 40 años de vida, la localización
de ejemplares inmaduros apunta a que existe cierto grado de reclutamiento pobiacional en
el tramo estudiado del río Mandeo.
ABSTRACT
In this study the population density, size distribution and age classes of the freshwater
pearl mussel Margaritifera margaritifera in the low stretch of the Mandeo river (A Coruña)
are analysed. Three sampling plots were selected in 2000, which were studied again in
2010, when a fourth plot was added. Differences in density were not significant between
years, and most sampled animáis were adult in all plots and in both years. Freshwater
pearl mussels found in the plot studied ¡ust in 2010 were significantly larger, and juvenile
mussels were only found on certain oreas of the river, suggesting a not homogenous
distribution of different age classes along the river. Although most freshwater pearl mussels
were 21-40 years oíd, some juveniles were also recorded, revealing some degree of
current recruitment in the studied stretch of the river Mandeo.
INTRODUCCIÓN
La náyade Margaritifera margaritifera
(Linnaeus 1758) se distribuye por las
fachadas costeras atlánticas de Norte¬
américa y Europa, incluida la Rusia
europea, en ríos árticos y atlánticos
(Smith, 2001). En Europa, Geist (2010)
* Departamento de Ecoloxía e Bioloxía Animal, Edificio Ciencias Experimentáis, Universidade de Vigo, As
Lagoas, 36310, Pontevedra, alvaro.barros@uvigo.es
35
Iberus , 32 (1), 2014
cita su presencia en diecinueve países.
Sin embargo, a pesar de esta extensa área
de distribución, se trata de una especie en
clara regresión a nivel mundial, actual¬
mente catalogada como "en peligro" por
la IUCN (2013). En Europa, las alteracio¬
nes provocadas por el hombre en sus
hábitats han causado el envejecimiento
de muchas poblaciones por falta de reclu¬
tamiento, cuando no su completa desapa¬
rición (Verdú y Galante, 2009). Entre
los impactos más importantes cabe citar
la construcción de obras hidráulicas (par¬
ticularmente embalses) en los ríos; estas
infraestructuras dificultan o impiden los
movimientos de los salmónidos, especies
sin las cuales la náyade no puede com¬
pletar su ciclo vital, al ser las larvas (glo-
quidios) parásitos obligados de las
mismas (Young y Williams, 1984;
Bauer, 1987a, 1987b; Beasley y Roberts,
1999). Al mismo tiempo, los embalses
provocan un cambio radical en el hábitat
de la especie al disminuir la velocidad de
la corriente y modificar los patrones de
deposición de los materiales sólidos en
suspensión (Moorkens, 2011). Otra
causa importante de desaparición es la
contaminación de las aguas, particular¬
mente la de origen orgánico, que afecta a
la especie fundamentalmente en su
estado juvenil (Bauer, 1988; Young,
1991; Young, Cosgrove y Hastie, 2001).
En la Península Ibérica, esta náyade
está presente actualmente únicamente en
el cuadrante noroccidental. En Portugal,
donde se consideró extinta durante
muchos años (Young et al., 2001), ha
sido redescubierta en seis ríos de la
cuenca del Duero, y al norte de esta en
los ríos Cávado y Neiva (Reís, 2003;
Varandas, Lopes-Lima, Teixeira, Hinz-
mann. Reís, Cortes, Machado y Sousa,
2013). En España, hoy en día se encuentra
únicamente en un curso de Asturias (río
Narcea; Álvarez-Claudio, García-
Rovés, Ocharan, Cabal, Ocharan y
Álvarez, 2000), en seis de Castilla y León
(ríos Bibey, Negro, Tera, Castro, Águeda
y Alberche; Velasco, Araujo, Bueno y
Laguna 2002; Morales, Negro, Lizana,
Martínez y Palacios, 2004; Velasco,
Araujo, Balset, Toledo y Machordom,
2006; Araujo, 2006) y en Galicia. Precisa¬
mente en esta última región es donde
mantiene su área de distribución más
amplia y continua en la Península, ocu¬
pando un buen número de cursos que
vierten sus aguas a las fachadas atlántica
y cantábrica (Araujo, 2008). En una
reciente revisión del estatus de la especie
en Galicia, Lois, Ondina, Outeiro,
Amaro y San Miguel (2013) citan su pre¬
sencia en un total de 55 ríos, encontrando
ejemplares juveniles en 11 de los mismos,
lo que pone de relieve la importancia
crucial de las poblaciones gallegas en la
conservación de la especie en Europa. En
el río Mandeo (A Coruña) hay referencias
históricas de su presencia (Macho, 1878),
mientras que la existencia de recluta¬
miento fue confirmada hace décadas
(Bauer, 1986). Según el trabajo de Lois et
al. (2013), el Mandeo sería el noveno río
con mayor población estimada de la
especie en Galicia (del orden de los 7.500
ejemplares), habiendo aún presencia de
ejemplares juveniles, aunque en escaso
número.
En el presente trabajo se aporta
información sobre el estatus de la
náyade de río en el curso bajo del río
Mandeo, analizándose los cambios
habidos en las densidades y en la estruc¬
tura por tamaños y edades de una
población estudiada con un lapso de
diez años. Además, se discute la viabili¬
dad de la población y las amenazas a las
que actualmente se enfrenta la especie
en este río.
MATERIAL Y MÉTODOS
Área de estudio
Con una longitud de 56 km, el río
Mandeo nace en la sierra de A Cova da
Serpe, a unos 700 m de altitud, y desem¬
boca en la ría de Betanzos, en la provin¬
cia de A Coruña. Durante una serie de
prospecciones realizadas en el año 1999
en los tramos medio y bajo del río, se de¬
terminó que la náyade de río estaba pre¬
sente en la mayor parte de los puntos
muestrados, si bien resultó ser especial¬
mente abundante en los últimos kilóme¬
tros del curso (Figura 1). Así, en agosto
de 2000 se procedió a delimitar tres par-
36
BARROS: Estado de conservación de Margaritifera margaritifera en el río Mandeo
Figura 1 . Área de estudio, indicando la localización aproximada de las parcelas estudiadas.
Figure 1. Study area, showing approximate situation oftbe sampled ureas.
celas de estudio en el citado tramo, a las
cuales se unió una cuarta parcela en
agosto de 2010 (Figura 1). La elección de
estos puntos de muestreo se realizó aten¬
diendo a la heterogeneidad de biotopos
que ocupa la especie en los ríos y que es¬
tán caracterizados fundamentalmente
por el tipo de sustrato, la profundidad y
la cobertura de sombra proyectada por
los árboles de ribera en el agua (Git-
tings, O'Keefe, Gallagher, Finn y
O'Mahony, 1998; Hastie, Philip y
Young, 2000) (Tabla I). En el primer año
de trabajo la posición de cada parcela fue
georeferenciada mediante un GPS, reali¬
zándose además distintos croquis que
permitiesen su localización exacta en
posteriores visitas.
Densidad y estructura de la pobla¬
ción por tamaños
El censo de ejemplares presentes en
cada parcela se realizó de manera visual
mediante la realización de transectos
longitudinales sistemáticos, cubriendo
toda la extensión de los sectores del río
delimitados. Cada transecto abarcó una
banda de censo de un metro de ancho a
cada lado del observador. Para evitar
conteos múltiples de un mismo ejemplar
se colocó una pequeña piedra de color
claro, cogida en el propio lecho del río,
al lado de cada náyade. El censo se dio
por concluido cuando en sucesivas repe¬
ticiones de los transectos no se localiza¬
ron animales sin dicha piedra anexa.
Cuando la visibilidad del fondo, por la
luz o por la corriente, fue mala, se
empleó un visor de fondo. En 2010 se
obtuvo además un dato de densidad
máxima en cada parcela, con el fin de
comparar esta medida con la existente
para otros ríos ibéricos. Este dato se
obtuvo contando los ejemplares que
quedaban englobados en un cuadrado
metálico de un metro de lado que fue
situado sobre las zonas con mayores
agrupamientos de náyades. Todos los
37
Iberus , 32 (1), 2014
Tabla I. Características físicas de las parcelas de estudio seleccionadas en el río Mandeo. Diámetros
de las partículas del sustrato: piedra grande >20 cm; piedra pequeña 2-20 cm; grava 2-20 mm;
arena < 2 mm.
Table I. Physical characteristics of sampled plots in the river Mandeo . Bottom partióles diameters: large
stone >20 cm; small stone 2-20 cm; gravel 2-20 mm; sand < 2 mm.
datos de densidades se presentan como
ejemplares/m2 (ej/m2).En cada parcela
se procedió a tomar la medida de la lon¬
gitud máxima de la concha del mayor
número posible de ejemplares con un
calibre de 0,05 mm de precisión. Una
vez medidas, las náyades fueron deposi¬
tadas en su ubicación exacta y las
piedras colocadas para su localización
fueron retiradas.
Estructura de la población por
edades
Los cálculos de las edades que
correspondieron a una determinada lon¬
gitud de la concha se realizaron emple¬
ando la fórmula de crecimiento no lineal
de von Bertalanffy (1938), expresada
como
L(t) = Lint [l-e'ht'V]
donde t representa la edad, L(t) la
longitud de la náyade a la edad t, Lmf es
una constante que representa la longi¬
tud asintótica o la longitud teórica
máxima que la náyade puede alcanzar
cuando t tiende a infinito, K representa
la constante de crecimiento y to es la
edad teórica a la cual la longitud de la
náyade debe ser cero. En nuestro caso,
los valores de Lmf, K y to fueron obteni¬
dos de los cálculos realizados por San
Miguel, Monserrat, Fernández,
Amaro, Hermida, Ondina y Altaba
(2004) para el propio río Mandeo.
Análisis estadístico
Los cambios en la densidad de las
tres parcelas entre los dos años de
estudio fueron analizados mediante el
test de Wilcoxon. Las longitudes de las
conchas se ajustaron a los supuestos de
normalidad y homocedasticidad
(prueba de Kolmogorov-Smirnov, a>
0,12 en todas las muestras), por lo que
se analizaron mediante análisis de la va-
rianza (ANOVA unifactorial). Final¬
mente, para analizar posibles varia¬
ciones en las frecuencias por categorías
de tamaño entre parcelas y años se
empleó el test X2. El nivel de significa¬
ción para todos los análisis se fijó en
0,05.
RESULTADOS
Densidad poblacional
En el año 2000 se contabilizó un total
de 168 ejemplares en el conjunto de las
tres parcelas estudiadas, frente a 183 en
2010; la parcela 4, controlada única¬
mente en 2010, sumó 370 individuos
(Tabla II). Atendiendo a las tres parcelas
para las que hay datos de los dos años
de estudio, las densidades observadas
no difirieron de manera significativa en
ninguna de ellas comparando los datos
de 2000 y 2010 (Wilcoxon, W = 3, P = 1).
Así, en 2000 la densidad fue máxima en
la parcela 2, media en la 3 y mínima en
la 1, situación que se repitió diez años
después. La única diferencia destacable
se dio en la parcela 2, donde en 2010 se
detectó un aumento del 21% frente al
número de ejemplares presentes en
2000. La mayor densidad de población
se registró en la parcela 4, con 6,17 ej/m2
y una densidad máxima de 50 ej/m2.
38
BARROS: Estado de conservación de Margaritifem margaritifera en el río Mandeo
Tabla II. Tamaño de la población, densidad (ejemplares/m2) y tasa de cambio en las tres parcelas
controladas en 2000 y 2010.
Table II. Population size, density (individuals/m2) and population change rate recorded on the three
studied plots in 2000 and 2010.
Distribución de tamaños
En el año 2000 se midieron 106 de
los 168 ejemplares contabilizados (63%),
mientras que en 2010 se midieron todos
los ejemplares encontrados en las parce¬
las maestreadas en 2000 (n = 183) más
una muestra de 100 ejemplares localiza¬
dos en la parcela 4, lo que sumó un total
de 282 ejemplares. Las longitudes en las
parcelas 1-3 fueron similares entre sí
tanto en 2000 (ANOVA unifactorial,
p2,io3 = 0,87; P = 0,42) como en 2010
(F2,i80 = 0,81; P = 0,45). Así mismo, no
difirieron en el conjunto de estas tres
parcelas entre años (Fi,287 = 0,85; P =
0,36). Por el contrario, los ejemplares de
la parcela 4 resultaron ser sensiblemente
mayores que los de las parcelas 1-3
(ANOVA unifactorial para varianzas
desiguales, F275,8 = 40,33; P < 0,0001),
mientras que 4 de los 5 únicos juveniles
(longitud inferior a los 65 mm; Hastie
et al., 2000; Young et al., 2001)
medidos entre los dos años de estudio
se encontraron en la parcela 2.
Tanto en 2000 como en 2010 y en las
cuatro parcelas consideradas la gran
mayoría de los ejemplares alcanzaron
una talla de 80-100 mm (Tabla III, Figura
2). En el año 2000, las frecuencias por
categorías de tamaño en la parcela 1,
considerando sólo tres categorías en
función de los tamaños de muestra (<80,
80-90 y >90 mm), no difirieron de las de
la parcela 2 (X2 = 4,17; g.l. = 2, P = 0,12; n
= 68) ni de la 3 (X2 = 2,03; g.l. = 2 ,P =
0,36; n = 50), mientras que estas dos
últimas tuvieron poblaciones estructura¬
das de manera similar (X2 = 0,77; g.l. = 2,
P = 0,68; n = 94). De las categorías de
tamaño consideradas, la mayoritaria en
todas las parcelas fue la de 80-90 mm.
En el año 2010 aumentó considerable¬
mente el número de náyades incluidas
en la categoría 90-100 mm en el conjunto
de la población (Figura 2). Una vez más,
las frecuencias no variaron entre las dis¬
tintas parcelas: parcela 1 vs. parcela 2:
X2= 1,47; g.l. = 2, P = 0,48; n = 131;
parcela 1 vs. parcela 3: X2 = 1,27; g.l. = 2,
P = 0,53; n = 62; parcela 2 vs. parcela 3:
X2 = 0,87; g.l. = 2, P = 0,65; n = 171.
Estructura de la población por
edades y longevidad
La porción mayoritaria de la pobla¬
ción medida en el río Mandeo en el año
2010, con una longitud de la concha de
80-100 mm, se corresponde con una
edad de 21 a 40 años, siendo por tanto
adultos reproductores; la población
juvenil, formada por animales de <65
mm (Figura 3), representaría única¬
mente el 1% de la población. Entre las
389 náyades medidas en este estudio, la
de mayor longitud, localizada en la
parcela 4, alcanzó los 123,55 mm, corres¬
pondiéndole una edad próxima al
máximo descrito para Galicia (65 años,
según San Miguel et al., 2004).
DISCUSIÓN
Cambios en la densidad y en la
estructura de la población por tamaños
Las variaciones registradas en la
densidad de las tres parcelas controladas
en 2000 y 2010 no fueron significativas, lo
que sugiere que la población se ha
39
Iherus , 32 (1), 2014
Tabla III. Tamaño de muestra, media longitudinal y desviación típica de las poblaciones
muestreadas en 2000 y 2010.
Table III. Sample size, average length and standard deviation of populations studied in 2000 and
2010.
mantenido estable en el período de diez
años considerado. Puesto que durante la
realización del trabajo de campo no se
detectaron transformaciones importantes
en el hábitat de las parcelas estudiadas, ni
la recolección directa de ejemplares, las
variaciones observadas en el número de
náyades presente en cada parcela
podrían deberse a fenómenos de mortali¬
dad natural (parcelas 1 y 3) o a la llegada
de nuevos ejemplares arrastrados por la
corriente en períodos de aguas caudalo¬
sas (parcela 2). El aumento registrado en
2010 del número de animales incluidos
en la categoría 90-100 mm podría estar
explicado simplemente por el creci¬
miento natural de los ejemplares, de unas
décimas de milímetro anuales (San
Miguel et al., 2004), en el período consi¬
derado.
Las frecuencias de las distintas cate¬
gorías de tamaño fueron similares entre
parcelas en los dos años de estudio. Sin
embargo, teniendo en cuenta el mayor
tamaño de los ejemplares de la parcela 4
y que casi todos los ejemplares juveniles
se encontraron en la parcela 2, es posible
que la distribución de los distintos
tamaños en el río no sea homogénea, si
no que distintas zonas del río podrían
tener condiciones adecuadas para el
asentamiento de unas clases de tamaño
determinadas (Duarte y Diefenbach
1994; Geist y Auerswald, 2007).
Sobre la viabilidad de la población
Las densidades observadas en el
curso bajo del río Mandeo en el presente
estudio (0,27-6,17 ej/m2) son superiores
a las estimadas para el conjunto del río
(0,008-3,7 ej/im; Lois et al., 2013), lo
que indicaría que el tramo estudiado
presenta condiciones particularmente
favorables para la especie. La gran
mayoría de los ejemplares localizados
fueron adultos, siendo claramente mino¬
ritaria la fracción de la población
formada por juveniles, que estuvo muy
lejos del 25% estimado como necesario
para que una población pueda ser consi¬
derada viable (Hastie y Cosgrove,
2002). La abundancia de juveniles es
posible que haya sido subestimada, ya
que durante esta fase las náyades per¬
manecen preferentemente enterradas y
para localizarlas es preciso tamizar el
lecho del río (Ziuganov, Zotin, Nezlin,
Tretiakov, 1994; Hastie et al., 2000;
Hastie y Cosgrove, 2002), lo que no se
ha realizado en el presente estudio. Sin
embargo, otros autores que sí realizaron
búsquedas específicas de juveniles ente¬
rrados (Lois et al., 2013), citan la pre¬
sencia de estos en un porcentaje bajo
(5%) sobre el total de las unidades de
muestreo en el Mandeo, destacando
además que únicamente el 27% de las
náyades juveniles en Galicia se encuen¬
tran enterradas en el sustrato. Por consi-
40
BARROS: Estado de conservación de Margaritifera margaritifera en el río Mandeo
mm
mm
Figura 2. Número de náyades por intervalos de tamaño (mm) medidas en el conjunto de las parce¬
las 1-3 en 2000 (n = 106) (arriba) y 2010 (n = 183) (abajo).
Figure 2. Number offreshwater pearl mussels on each class size (mm) measured on plots 1-3 in 2000 (n
= 106) (above) and in 2010 (n = 183) (beiow).
Figura 3. Ejemplar adulto de Margaritifera margaritifera de unos 100 mm de longitud junto a dos
inmaduros de unos 43 mm, fotografiados durante la realización del presente estudio en el río
Mandeo. En estos últimos son visibles a simple vista seis anillos de crecimiento anual.
Figure 3. An adult Margaritifera margaritifera about 100 mm length along with two immatures about
45 mm length found during the present study on the river Mandeo. Six anual growth rings are easy
visible in the latter.
41
Iberas , 32 (1), 2014
guíente, la información actualmente dis¬
ponible apunta a que la presencia de
juveniles en el río Mandeo es escasa,
encontrándonos ante una población en
claro proceso de envejecimiento.
Como conclusión, el hecho de que las
densidades de náyades se hayan mante¬
nido prácticamente invariables en el
período de diez años considerado,
sugiere que el curso bajo del río Mandeo
mantiene un hábitat favorable para la
especie. En este sentido, es de destacar
que las densidades de salmónidos (parti¬
cularmente de trucha común Salmo trutta
fario ) observadas en el Mandeo están
muy próximas a las que registran
algunos de los ríos con las poblaciones de
náyades más importantes de Galicia,
como el Eo o el Masma (Hervella y
Caballero, 1999; Lois et al., 2013). Sin
embargo, es preocupante la escasez de
náyades juveniles, lo que supone una
amenaza el mantenimiento a largo plazo
de la población. Durante la realización
del presente estudio se detectaron distin¬
tas amenazas que se ciernen de manera
preocupante sobre la población de
náyades estudiada, pese a encontrarse
incluida en el LIC Betanzos-Mandeo.
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A José A. de Souza, Manuel Aira,
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42
BARROS: Estado de conservación de Margaritifera margaritifera en el río Mandeo
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a
© Sociedad Española de Malacología
Iberas , 32 (1): 45-51, 2014
Wm
A new Aforia (Gastropoda: Conoidea: Cochlespiridae) from
Galicia Bank (NW Iberian Península)
Una nueva Aforia (Gastropoda: Conoidea: Cochlespiridae) del banco
de Galicia (NO Península Ibérica)
Serge GOFAS*, Yuri KANTOR** and Ángel A. LUQUE ***
Recibido el 1 l-X-2013. Aceptado el 13-1-2014
ABSTRACT
A gastropod coliected ¡n 1720 m depth at the foot of the Galicia Bank ¡s described as a
new species in the genus Aforia (Cochlespiridae). The family placement is supported by
the characters of the radula. The disjunct distribution (Northern Pacific, Antarctic and
subantarctic areas, and here NE Atlantic) of species currently assigned to the genus Aforia
is discussed.
RESUMEN
Se describe un gasterópodo recogido a 1720 m de profundidad al pie del banco de
Galicia como una nueva especie del género Aforia (Cochlespiridae). Su inclusión en esta
familia es acorde con los caracteres de la rádula. Se discute la distribución discontinua
(Pacífico Norte, Antártida y áreas subantárticas, y aquí Atlántico NE) de las especies
actualmente asignadas al género Aforia.
INTRODUCTION
The exploration of the deep-sea
benthos of Western Europe has been
active since the end of the XIX century
and has increased spectacularly since
the last two decades of the XX century.
Nevertheless, contrary to the Coastal
area where the finding of a new species
is now exceptional, there is still much to
be discovered even among large and
spectacular benthic animáis of the
bathyal and abyssal zones.
The Galicia Bank is a seamount
located about 200 km west of the NW
Iberian Península, culmina ting at ca. 600
m depth on a large (ca. 6250 km2)
plateau, and separated from the main-
land shelf by depths of ca. 2500 m in the
Galicia Inner Basin. It is irregularly
shaped, with the N-NW part gently
sloping down to the abyssal plain and
the eastern edge forming a cliff down to
ca. 1800 m. The substrate is composed of
basaltic lavas and oceanic crust, covered
by sediments which are largely of pelagic
origin (Ercilla, Casas, Vázquez, Igle¬
sias, Somoza, Juan, Medialdea, León,
Estrada, García-Gil, Farrán, Bohoyo,
García & Maestro, 2011). Regarding
^Departamento de Biología Animal, Facultad de Ciencias, Universidad de Málaga, Campus de Teatinos -
29071 Málaga, Spain . E-mail: sgofas@uma.es
** A. N. Severtzov Institute of Ecology and Evoíution, Russian Academy of Sciences, Leninski prosp. 33,
Moscow 119071, Russia. - E-mail: kantor.yuril956@gmail.com
*** Laboratorio de Biología Marina, Departamento de Biología, c/ Darwin, 2, Universidad Autónoma, 28049
Madrid, Spain - E-mail angel.luque@uam.es
45
Iberus , 32 (1), 2014
water masses in contact with the sea
bottom (Fiúza, Hamann, Ambar, Díaz
del Río, González & Cabanas, 1998),
the bank is reached from the South by
quite diluted Mediterranean Outflow
Water at two depth intervals, the upper
core centred around 780 m and the lower,
most saline core around 1100-1200 m. In
the deeper part, North Atlantic Deep
Water is present regionally with a core
situated ca. 2500-3000 m, but a wedge of
the fresher and colder Labrador Sea
Water (LSW) situated around 1800 m and
moving southwards, locally accentuates
the gradient with the lower Mediter¬
ranean outflow.
The information about the bank and
its ecosystems, and the few scientific
surveys have highlighted the high con-
centrations of nutrients and the presence
of deep-sea coráis in the shallower part of
the bank (Duineveld, Lavaleye &
Berghuis, 2004). The bank has been
visited by several expeditions in the past,
among which the French Campaign
SEAMOUNT 1 (Muséum National d'His-
toire Naturelle, París, 1987), in which 12
stations were sampled (Bouchet &
Métivier, 1988). In 2009, the Diva-Artabria
11-09 campaign was conducted within the
project "Latitudinal gradients of biodi-
versity in the deep sea of the Atlantic
Ocean" and sampled Galicia Bank. Also
in 2009, the Spanish Institute of Oceanog-
raphy (IEO) conducted the ECOMARG
programme on several sites of the north-
em Iberian margin including Galicia Bank.
Although quite abundant material was
obtained, no comprehensive list of mol-
luscs was ever published and the only
reports regard isolated species (e.g. Rolán
Mosquera & Pedrosa, 1981), often in the
context of broader taxonomic work (e.g.
Bouchet & Warén, 1993; Gofas, 2007).
INDEMARES is an EU-funded LIFE+
project aimed at documenting ten off-
shore areas of Spanish waters as
prospects for marine Natura 2000 areas,
thereby fulfilling the commitments of the
Marine Strategy Framework Directive of
the EU. Galicia Bank is one of these ten
areas and was the target of the cruise
INDEMARES 0711 from July, 18 to
August, 10 2011, with the R/V "Miguel
Oliver" (of Secretaría General de Pesca,
Spanish Ministry of Agriculture, Food
and Environment), under the expedition
leader Dr. Alberto Serrano (Instituto
Español de Oceanografía). This expedi¬
tion made operations with rock dredge
(15 hauls, 779-1697 m depth), beam trawl
(11 hauls, 744-1720 m depth) and otter
trawl (9 hauls, 751-1764 m depth), along
with box core samples, suprabenthic sled
hauls, pelagic net operations, underwater
video recording and CTD measurements.
Here we present the description of a
large conoidean gastropod collected
during the INDEMARES 0711 cruise.
The unique specimen comes from the
deepest beam trawl operation, situated
in the transition zone between Galicia
Bank proper and the Galicia Inner Basin,
near the foot of the eastern cliff of the
bank.
MATERIAL AND METHODS
During INDEMARES 0711 cruise,
the beam trawl was towed for 15
minutes after settling on the bottom, at a
speed of two knots. The mesh was 10
mm but consistently brought up a finer
fraction as it got clogged by the sedi-
ment. The samples were sieved immedi-
ately on board, in sea water, on a set of
10, 5, 2 and 0.5 mm meshes; the two
coarser fractions were sorted also on
board and immediately fixed in 96°
ethanol, and the specimen studied here
was processed in this way. The finer
fractions were mostly fixed in totality
for posterior sorting in the lab.
The sample
Haul VIO was a successful beam trawl
operation which brought on board over
10,000 macroscopic specimens totalizing
over 9,600 grams (for the fractions sorted
on board). The sediment was a pteropod
ooze, in which shells of Cavolinia inflexa
(Lesueur, 1813) were the most conspicu-
ous component. The bottom sea-water
temperature at the corresponding depth
on the nearest CTD profile was 6.36 °C,
and the salinity 35.3824 %o. From the
nearest box core (BC16, 42° 43.536' N - 11°
46
GOFAS ETAL.: A new Aforia (Gastropoda: Conoidea: Cochlespiridae) from Galicia Bank
28.128' W, 1751 m), sediment was a poorly
sorted very fine sand, with 2.411% over
500 pm, 51.280 % 63-500 pm and 46.309 <
63 pm, and had an organic matter conten!
of 3.69%.
Most abundant species of the mac-
robenthos in terms of biomass were the
deep-sea eel Synaphobranchus kaupii
Johnson, 1862 (80 specimens, 1510 g)
and the holothurian Benthogone rosea
Koehler, 1895 (27 specimens, 1880 g).
Other abundant elements collected alive
were unidentified ophiuroids, the
bivalves Limopsis eristata Jeffreys, 1876
(most abundant species overall, 702
specimens) and Parvamussium propin-
quum (Smith, 1885) (570 specimens), the
large scaphopod Fissidentalium capillo-
sum (Jeffreys, 1877) (150 specimens), the
gastropods Kryptos koehleri (Locard,
1896) (129 specimens), Amphissa acute-
costata (Philippi, 1844) (87 specimens)
and Scaphander punctostriatus (Mighels
& Adams, 1842) (59 specimens). In total,
37 live-taken molluscan species were
identified in the sample (23 gastropods,
11 bivalves and 3 scaphopods). Shells of
Fissidentalium occupied by sipunculids
were also abundant. The haul also con-
tained a single specimen of a large gas-
tropod, which appeared to be a new
species and is described herein.
SYSTEM ATIC PART
Genus Aforia Dalí, 1889
Type species: Pleurotoma circinata Dalí, 1873 [Recent, Bering and Okhotsk seas], by original desig-
nation.
Aforia serranoi spec. rtov. (Figs. 1-6)
Type material: Holotype, live taken specimen (MNCN15.05/60099)), BANGAL 0711, haul VIO
(R/V "Miguel Oliver", 08 08 2011).
Type locality: East of Galicia Bank (42° 41.875' N, 11° 26.708' W, 1,720 m to 42° 42.36' N, 11° 26.93'
W, 1,723 m).
Etymology: The species ñame is dedicated to the expedition leader Dr. Alberto Serrano, of Insti¬
tuto Español de Oceanografía.
Description : Shell fusiform, comprising
eight whorls, white, rather opaque. Pro-
toconch comprising about 1.75 whorls,
smooth and globose, corresponding only
to protoconch I, with a máximum diame-
ter of 1.2 mm, height 1.45 mrri. Protoconch-
teleoconch transition indistinct, indicated
by the onset of a deep anal sinus and a
keel. Teleoconch sculpture dominated by
a distinct keel running near the middle of
the whorls, slightly closer to the abapical
suture, and a second keel, much less pro-
nounced, running on the body whorl in
continuation of the suture. Subsutural
ramp very slightly concave just below the
suture, then fíat down to the main keel,
covered by very faint, fíat spiral cordlets
and by flexuous growth lines reflecting
the former positions of the anal sinus.
Sculpture between keels and on the abapi¬
cal part of the body whorl, of fine, fíat and
rounded spiral cords, much more definite
than on the subsutural ramp. Subsutural
ramp practically smooth on the three
upper spire whorls, which have two low
cords below the keel. Cords becoming
more distinct on the subsutural ramp start-
ing on fourth whorl, from 7 on the 4th
whorl to 9 on the penultimate and last
ones. Cords closely spaced, with variable
width and interspaces variable from
slightly broader than the cords to half the
width of the cords. Number of cords below
the keel increasing from 2 to 4 on fourth
whorl and to 6 on penultimate whorl; six
spiral cords between the keels on the last
whorl and 23 on the shell base and canal.
Body whorl prolonged by a tapering
siphonal canal, which is not separated
from the rest of the whorl by any discon-
47
Iberus , 32 (1), 2014
tinuity of the profile. Aper ture elongate-
pyriform, tapering gradually towards the
siphonal canal. Colurnellar and parietal
edge continuous, lined with a very thin
callus. Outer lip simple, rather thin, with
a broad parabolic shaped anal sinus
between the suture and the peripheral keel.
Dimensions: shell length 33.6 mm,
width 11.2 mm; body whorl length 22.5
mm (measured along the axis on Fig. 1),
aperture length (with canal) 17.9 mm,
aperture length (without canal) 10.5 mm.
Animal rather stout, with a small
head provided with cylindrical tenta-
cles, blind. Operculum large, 8 mm in
length, oval, spiral, with nucleus
strongly shifted to left, semitransparent
and brownish.
Radula relatively short, consisting of
ca. 30 transverse rows of teeth, 10
nascent, 1.39 mm in length (13% of aper¬
ture length without canal). Radula com-
prising three teeth per row permanently
attached to the basal membrane. Central
tooth wide, weak, with arched posterior
margin, with slightly elevated edges of
posterior and lateral margins, and ante¬
rior margin indistinct, fused with the
subradular membrane. A single small tri¬
angular cusp emanating from posterior
margin. Lateral teeth small (length
around 180 pm or 1.7% of aperture length
without canal), dúplex, rather fíat, with
pointed major limb with sharp edges,
and large accessory limb of same length.
Systematic placement and species com-
parisons : Aforia is currently considered
an extremely broadly distributed genus
comprising 18 Recent species found
from upper boreal waters in the Pacific
to Antárctica (WoRMS, 2013). The
integrity of the group has never been
studied molecularly (only a single
Antarctic species, Aforia magnifica
(Strebel, 1908) can be found in
Genbank). Therefore, the genus is cur¬
rently defined mostly on rather loose
conchological grounds and, to a lesser
extent, on radular characters. Seven
species of Aforia from north Pacific and
three from Antarctic and sub-Antarctic
waters have been studied in this respect,
and all these have rather similar
radulae, including A. serranoi spec. nov.
However, the radula of all known
Cochlespiridae species is very similar
and hardly can be an ultimate proof that
boreal, Antarctic and subantarctic
species of Aforia are congeneric. Thus,
pending additional molecular data on
different Aforia species, we tentatively
attribute the new species to the genus
Aforia as broadly conceived on the
grounds of shell and radula similarities
with the other known species.
The type species of the genus, Aforia
circinata (Dalí, 1873) described from
Alaska (Nateekin Bay, Unalashka, Aleut-
ian Islands) is twice as large (the holotype
measures 7.35 cm with a similar number
of whorls) as Aforia serranoi spec. nov.
and lacks the abapical keel, having
instead the base of the body whorl regu-
larly rounded. Similarly, the abapical keel
is absent in all other species of Aforia
described from the north Pacific
(although most of them, except A. kincaidi
(Dalí, 1919) are currently considered as
synonyms of A. circinata).
Other species of Aforia are known
from Antarctic waters (Dell, 1990),
among which A. magnifica (Strebel, 1908)
and A. multispiralis Dell, 1990 most
resemble the species described here.
They share the configuration of spiral
keels with fíat spiral cords in between,
but are twice as large and with a less
acute spire in the former and more
prominent keels in the latter. Aforia sta-
minea (Watson, 1881), from Marión and
Prince Edward and Kerguelen Islands is
also larger than A. serranoi spec. nov.,
and lacks the abapical spiral keel
(Watson, 1886). Aforia goniodes (Watson,
1881), described from Atlantic waters
(Río de La Plata, Argentina), is smaller
and stouter and also lacks the abapical
keel (Watson, 1886).
Sysoev and Kantor (1987) revie-
wed on conchological and anatomical
grounds the Pacific species of Aforia and
cióse genera and described three new
subgenera, one exclusively fossil. They
also followed the opinión of Powell
(1966) on the synonymy of Irenosyrinx
Dalí, 1908 (type species Pleurotoma (. Leu -
cosyrinx) goodei Dalí, 1890, type locality
north-western Patagonia, 1920 m) with
48
GOFAS ETAL.: A new Aforia (Gastropoda: Conoidea: Cochlespiridae) from Galicia Bank
Figures 1-6. Aforia serranoi spec. nov. Holotype, MNCN1 5.05/60099, East of Galicia Bank (42°
41.87’ N, 11° 26.71’ W, 1720 m to 42° 42.36’ N, 11° 26.93’ W, 1723 m). 1, 2: ventral and
dorsal view of shell, actual size 33.6 mm; 3: detail of protoconch; 4: operculum, largest diameter 8
mm; 5: partial view of the radula; 6: detail of marginal teeth.
Figuras 1-6. Aforia serranoi spec. nov. Holotipo, MNCN 15.05/60099, Este del banco de Galicia (42°
41,87’ N, 11° 26,71’ W, 1 720 m, a 42° 42,36’ N, 11° 26,93’ W, 1723 m). 1, 2: vista ventral y dorsal
de la concha, tamaño real 33, 6 mm; 3: detalle de la protoconcha; 4: opérenlo, diámetro mayor 8 mm;
5: vista parcial de la rádula; 6: detalle de los dientes marginales.
Aforia (Aforia). The generic status of
Irenosyrinx is not yet finally accepted,
although in the latest classification
(BOUCHET, KANTOR, SYSOEV & PUILLAN-
dre, 2011) the genus is listed question-
ably as a subgenus of Aforia.
Bouchet and Warén (1980) treated
Irenosyrinx as a sepárate genus, and
included one species described from the
Western Atlantic, L hypomela (Dalí, 1889)
(= Surcula tenerrima Locard, 1897) and
distributed also in the central Atlantic. It
is readily distinguished from A. serranoi
spec. nov. by a very weak upper keel
and the absence of the second abapical
one and much stronger spiral elements
on the subsutural ramp.
A large shell (25 mm) figured by
Bouchet and Warén (1980) under the
ñame Ancistrosyrinx clytotropis (Sykes,
1906) is possibly congeneric with Aforia
and it is the known species that comes
morphologically closest to the species
described here, at least in the European
realm. This shell comes from the north-
ern coast of Spain (700-1120 m) and
differs in being considerably stouter,
with a more pronounced, protruding
adapical keel and has the abapical keel
marked by a duplicated cord. Sykes7
49
Iherus , 32 (1), 2014
holotype collected off the West coast of
Portugal ("Porcupine" Sta. 17, 39° 42' N,
09° 43' W, 1980 m) is considerably
smaller (8 mm), but maintains the same
profile as the early whorls of the shell
figured by Bouchet and Warén (1980).
It should be mentioned that Bouchet
and Warén (1980) think that all extant
material of A. clytotropis including the
type is probably fossil.
Remarles on distribution : The present
report is the first record of Aforia in the
North Atlantic Ocean. The geographic
distribution of Aforia, if the generic
assignment of the new species is correct,
is intriguing. Given that Ancistrosyrinx
clytotropis (Sykes, 1906) is the morpho-
logically closest species to Aforia serranoi
spec. nov., it would turn out that mor-
phology suggests they are congeneric.
The paucispiral protoconch of all
species in this genus clearly indicates a
lecithotrophic (probably entirely intra-
capsular) development, which would
preelude long-distance pelagic dispersal
ACKNOWLEDGEMENTS
SG and AL are grateful to Alberto
Serrano, of Instituto Español de Oceano¬
grafía (Santander), leader of the Galicia
Bank workpackage of INDEMARES, for
the invitation to particípate in the 0711
BIBLIOGRAPHY
Bouchet P., Kantor Y.I., Sysoev A. & Pui-
llandre N. 2011. A new operational classi-
fication of the Conoidea (Gastropoda). Jour¬
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Bouchet P. & Métivier B. 1988. Campagne
Océanographique «SEAMOUNT 1». Compte
rendu et liste des stations. 29 pp. 1MSU /CMRS
(unpublished cruise report).
Bouchet P. & Warén A. 1980. Revisión of
the North-East Atlantic bathyal and abys-
sal Turridae (Mollusca, Gastropoda). Jour¬
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119.
Bouchet P. & Warén A. 1993. Revisión of
the Northeast Atlantic bathyal and abyssal
Mesogastropoda. Bollettino Malacologico
Supplemento 3: 577-840.
in the larval stage. There are however
more instances of this kind of distribu¬
tion among gastropods with intracapsu-
lar development, the most noteworthy
and best documented being that of a
group of closely related genera of the
muricid subfamily Trophoninae (Rara-
SEWYCH, 1984).
Powell (1951: 63-66) discussed bipo-
larity but pointed out that the distribu¬
tion of Aforia achieves continuity with the
presence of species such as A. persimilis
(Dalí, 1889) off the Pacific coast of Central
and South America. In the Atlantic, there
are no relevant data along the West
African slope, but A. serranoi spec. nov.
was found in the context of water masses
flowing from the North. Taking into
account that the slope fauna of the NW
Atlantic is relatively well known, such a
source area for this species is unlikely
and therefore its occurrence off Western
Europe remains unexplained and could
have originated in a different paleoceano-
graphic setting.
cruise and for handing over the material
for study; to shipboard companions and
crew of R/V "Miguel Oliver" for effi-
cient and pleasant collaboration during
the cruise.
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to a seamount cold-water coral community
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R., Estrada F., García-Gil S., Farrán M.,
Bohoyo F., García M. & Maestro A. 2011.
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poda) from northeast Atlantic Seamounts.
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tomy of four primitive muricacean gastro-
pods. Implications for Trophoninae phylo-
geny. American Malacological Bulletin, 3: 11-
26.
Powell A.W.B. 1951. Antarctic and Suban-
tarctic Mollusca: Pelecypoda and Gastro¬
poda. Discovery Reports, 26: 47-196, pl. 5-
10.
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Speightiidae and Turridae. An evaluation
of the valid taxa, both Recent and fossil,
with lists of characteristic species. Bulletin
of the Auckland Institute and Museum, 5: 5-
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functional morphology of the digestive Sys¬
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51
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■
© Sociedad Española de Malacología — Iherus , 32 (1): 53-64, 2014
Wm
Kaloplocamus ramosus (Cantraine, 1835) (Gastropoda:
Polyceridae): new records in the Bay of Biscay, with notes
on distribution and food
Kaloplocamus ramosus (Cantraine, 1835) (Gastropoda: Polyceridae):
nuevos sitios en el Golfo de Vizcaya, con datos sobre su distribución
y su dieta
Alex VANHAELEN* **, Claude MASSIN*, Jocelyne MARTIN” and Pascal
LAFFARGUE”
Recibido el 10-IX-2013. Aceptado el 16-1-2014
ABSTRACT
Many specimens of Kaloplocamus ramosus (Cantraine, 1 835) were collected ¡n the Bay of
Biscay during Ifremer's EVHOE scientific cruises in 2009 and 2010 (October - Novem-
ber). This species is new to the French Atlantic coast, which is its most northern known
occurrence, and a map of its distribution in the Bay of Biscay is provided. Two prey-
species (Bryozoa) were observed from gut contents. One of these, Cellaria salicornioides,
is an addition to the known food-list of K. ramosus. Data on the taxonomy are given and
Doris fimbñata delle Chiaje, 1841 is added to the synonymy of K. ramosus.
RÉSUMÉ
De nombreux spécimens de Kaloplocamus ramosus (Cantraine, 1835) ont été récoltés
dans le golfe de Gascogne au cours des campagnes scientifiques Ifremer EVHOE de
2009 et 2010 (octobre - novembre). L'espéce est nouvelle pour les cotes atlantiques
franpaises qui représentent sa latitude la plus nordique, et une carte de sa distribution
dans le Golfe de Gascogne est présentée. Deux espéces de Bryozoa ont été trouvées dans
le contenu stomacal. l'une d'entre elles, Cellaria salicornioides, est mentionnée pour la
premiére fois comme nourriture pour K. ramosus. Des informations sur la taxonomie sont
fournies et Doris fimbriata delle Chiaje, 1 841 est ajoutée á la synonymie de K. ramosus.
RESUMEN
Numerosos ejemplares de Kaloplocamus ramosus (Cantraine, 1 835) fueron recogidos en el
Golfo de Vizcaya durante las campañas científicas Ifremer EVHOE en 2009 y 2010 (entre
octubre y noviembre). La especie es nueva para la costa Atlántica francesa, lo cual repre¬
senta la cita más septentrional conocida, y se presenta un mapa de su distribución en el Golfo
de Vizcaya. Dos especies-presa (Bryozoa) fueron observadas en el contenido estomacal. Una
de ellas, Cellaria salicornioides, se añade a la lista de presas conocidas de esta especie. Se
incluye un mapa de distribución en el Goifo de Vizcaya así como también datos sobre la
taxonomía, añadiendo Doris fimbriata Delle Chiaje, 1841 a la sinonimia de K. ramosus.
* Royal Belgian Institute of Natural Sciences, (RBINS), Department of Recent Invertebrates, Section
Malacology. Vautier Street, 29, B-1000 Brussels, Belgium.
** Instituí franjáis de recherches pour l’exploitation de la mer (Ifremer), Unité Écologie et Modeles pour
l’Halieutique, Ifremer Centre de Nantes, rué de file d’Yeu, B.P. 21105. France.
Corresponding author: alex.vanhaelen@naturalsciences.be
53
Iherus , 32 (1), 2014
Figure 1. Map 1. EVHOE 2009-2010. Sampling area and stations. Some symbols of stations
(01016/N0817, 00991/N0779, 00992/N0780) overlap, see Table I. Stars represent localities
where Kaloplocamus ramosus was found.
Figura 1. Mapa 1. EVHOE 2009-2010. Area y estaciones de maestreo. Algunos símbolos de estaciones
(01016/N0817, O099HN0779, 00992/N0780) se confunden, véase la Tabla I. Las estrellas represen¬
tan localidades en las que se encontró Kaloplocamus ramosus.
INTRODUCTION
Opisthobranchs are well recorded in
the Southern part of the Bay of Biscay
(García and Bertsch 2009) as well as in
the Celtic Sea and the English Channel
(Hayward and Ryland 1990). However,
they have been less well investigated in
the northern part of the Bay of Biscay,
which is the northern limit of the Lusi-
tanian Province. During recent EVHOE
(Evaluation Halieutique pour FOuest de
l'Europe) scientific cruises in this area,
whose main purpose was to assess the
state of the stock of commercially valu-
able species (fishes, cephalopods, crus¬
tácea), numerous specimens of the opistho-
branch Kaloplocamus ramosus (Cantraine,
1835) were collected for the first time.
K. ramosus is a widespread species,
living mainly in températe waters, but
also reported in tropical waters ( e.g . East
coast of South Africa (Bergh 1907) and
the Hawaiian Islands (Pittman and
Fiene 2010)).
The present paper reports on geo-
graphic distribution, taxonomy, anato-
my, food and habitat of K. ramosus.
54
VANHAELEN ET AL.: Kaloplocamus ramosus new records in the Bay of Biscay
Table I. Kaloplocamus ramosus (Cantraine, 1835). Sample data. All specimens collected by Jocelyne
Martin and Pascal Laffargue/Ifremer, determined by Alex Vanhaelen/RBINS. Cruise ref. nbr:
cruise reference number; Station ref. nbr: station reference number; Station references: N= 2009,
0= 2010; qty= quantity. Total stations: 18. Total specimens: 89 (87 at RBINS, 2 at Ifremer).
Tabla I. Kaloplocamus ramosus ( Cantraine , 1835). Datos de muestro. Todos los ejemplares recolectados
por Jocelyne Martin y Pascal Lajf argüe/ Ifremer, identificados por Alex Vanhaelen/RBINS. Cruise refi
nbr: número de referencia de la campaña; Station ref nbr: número de referencia de la estación; Datos
de la estación: N= 2009, 0= 2010; qty= cantidad. Total de estaciones: 18. Especímenes totales: 89 (87
de RBINS , 2 de Ifremer).
MATERIALS AND METHODS
The specimens were collected at 18
stations off Southern Brittany (Fig. 1,
Table I) during the EVHOE surveys on
board R/V Thalassa. During October
and November of 2009 and 2010, 142
bottom trawls (stations) were performed
each year.
The sampling method ineludes
diurnal standardized 30 minutes bottom
trawls at a speed of 4 knots, using a
GOV-36/47 otter trawl, 20 m wide, 4 m
high with a 20 mm mesh codend. This
gear is not adapted to capture small ben-
thic species (<3-4 cm) such as K. ramosus.
Therefore the absence of this species at
some stations could be an artefact. For
the same reason, the population density
could be underestimated.
Geographical coordinates, depths, sea
water temperatures and nature of the
bottom were recorded during the hauls.
All data pertaining to a given station
were uploaded in a database at Ifremer.
Contents of the trawl's pocket were
sorted on board. Unidentified opistho-
branchia were sorted, photographed
when in good condition and properly
labeled prior to fixation, usually in a 70%
ethanol solution or deep-frozen. Speci¬
mens were sent to RBINS for Identifica¬
tion and transfered to a 70% ethanol,
borax-buffered, (pH 8.0) solution.
The radula of specimen N0799/07
was dissected, critical point dried and
55
Iberus , 32 (1), 2014
gold-coated to perform SEM pho-
tographs with RBIN's FEI-QUANTA-
200 ESEM (Environmental Scanning
Electron Microscope) (Fig. 7).
Ten specimens (one per station, except
for station 0-1016) were dissected for Iden¬
tification of gut contents. Those stations
were selected along a line joining the most
southeasterly station (0-0961) to the most
northwesterly station (0-1026), at the most
regular possible intervals. Stations con¬
cerned were N-0796, N-0817, N-0822, O-
0954, 0-0961, 0-0922, 0-1016 (2 speci¬
mens), 0-1025 and 0-1026. Gut content
was immersed during 10 minutes in com-
mercial bleach, and then rinsed several
times in tap water. The bryozoa were iden-
tified according to De Blauwe (2009).
Except for two specimens kept at
Ifremer/Nantes (Ref. EVHOE2009-
N0817/02 and EVHOE2010-O962), the col-
lection was deposited at RBINS (general
catalogue nurnher IG-31.621) and assigned
an individual Recent Invertebrates Depart¬
ment code nurnber in the series INV.98.001
to INV.98.028
Two specimens from sample NOS 17
were photographed in a sea water filled
container, alive, relaxed and extended,
prior to fixation in ethanol (Fig. 2).
Photographs of preserved material
were taken through the ocular of a dis-
secting microscope with a handheld
camera.
For nomenclature we chose to follow
the CLEMAM taxonomic database.
RESULTS
Class Gastropoda Cuvier, 1797
Family Polyceridae Alder & Hancock, 1845
Genus Kaloplocamus Bergh, 1880 [1879]
Kaloplocamus ramosus (Cantraine, 1835)
Doris ramosa Cantraine, 1835: Bull. Acad. R. Belg., 2: 383; 1841, Nouv. Mém. Acad. R. Sci. Bruxelles,
13: 54, pl. 3 fig. 7. [type locality: Split (as Spalato), Croatia, type specimen lost].
Euplocamus croceus Philippi, 1836, Enum. Molí. Sicil., 1: 103, pl. 7 fig. 1 [Palermo (as Panormi),
Italy], — Bergh, 1880, Verh. zool.-bot. Ges. Wien, 29: 625
Euplocamus frondosas Philippi, 1839, Arch. Naturgesch., 5 (1): 114, pl. 3 fig. 1 [Sicily, Italy].
Doris fimbriata delle Chiaje, 1841, Descr. Anim. s. Vert., 2: 21; 5: 77; 6: pl. 44, Figs 7-10. [Napoli,
Italy] — new synonym.
Triopa yatesi Angas, 1864, }. Conchyl., París, 12: 60, col. pl. v, Fig. 8. [Watson Bay, S.E. Australia].
Euplocamus ramosus — Bergh, 1880, Verh. zool.-bot. Ges. Wien, 29: 625, in synonymy of Euplocamus
croceus PhiL, erroneously credited to "Phil. Malacol. médit. 1840. p. 54. pl. 3. Fig. 7.". This is
clearly confusión with Cantraine's 1841 publication, cited with a wrong date.
Euplocamus capensis ; Bartsch, 1915, Bull. U.S. natn. Mus. 91: 239, 212.
Kaloplocamus aureus Odhner, 1932, Ark. Zool., 23. A. (14): 41, fig. 11-12. [Bahía del Confital (juv.)
Gran Canaria].
Caloplocamus ramosus — Pruvot-Fol, 1951, Arch. Zool. exp. gén., 88 (1): 35, pl. 2 Figs 3, 16. [juvenile
from Banyuls, France] (generic ñame misspelled).
Euplocamus plumosus Schultz — Pruvot-Fol, 1954, Faune Fr., 58: 323. (not Euplocamus plumosus W.
Thompson, 1840, a júnior synonym of Limada clavigera (O. F. Müller, 1776)). [Sicily, Italy] (this
ñame could not be traced elsewhere).
Kaloplocamus filosus Cattaneo-Vietti and Sordi, 1988, Basteria, 52: 50, fig. 1-12. [Tuscany, Italy].
Thirty nine specimens of K. ramosus
were collected during the 2009 cruise
and fifty during the 2010 cruise.
Anatomy: The body of live specimens
(Fig. 2) is slug-like and about 3 times
longer than broad. Extended lengths are
between 29 and 33 mm. Contracted
bodies measure from 16 to 28 mm long.
The buccal mass is more or less evagi-
nated in most specimens. Transversal
section at mid-body is quadrangular.
The solé is smooth. Anterior edge of
56
Vanhaelen ET AL.: Kaloplocamus ramosus new records in the Bay of Biscay
Figures 2-5. Kaloplocamus ramosus. 2: dorsal view of two living specimens relaxed in sea water
(sample NOS 17); 3: branched processes on oral veil (arrows); 4: oral tentacle (arrow); 5: eyes on
dorsum of specimen N0822 (arrows). Figure 6. Bryozoa extracted from gut of specimen
N0796/08. A: Cellaria salicornioid.es: B: Caberea boryi.
Figuras 2-5. Kaloplocamus ramosus. 2: vista dorsal de dos ejemplares vivos relajados en agua de mar
( muestro NOS 17); 3: papillas ramosas del velo oral (flechas); 4: tentáculo oral (flecha); 5: ojos en el
dorso del espécimen N0822 (flechas). Figura 6. Briozoos extraídos del estómago del espécimen
N 0796/ 08. A: Cellaria sálico rnioides, B: Caberea boryi.
propodium is horizontally bilamellated.
No propodial tentacles are noticed.
Lateral edges are thick, narrow and
undulating.
In front, the oral veil is slightly
rounded, with 6-7 velar processes, here-
after named 'branched processes',
bearing short secondary papillae on
their distal third. (Fig. 3).
The dorsum is lined by a longitudi¬
nal row of 5 similar but larger branching
processes, the first in the row starting
57
Iberus , 32 (1), 2014
Table II. Kaloplocamus ramosus. Known prey-species.
Tabla II. Kaloplocamus ramosus. Especies de presas conocidas.
just behind and under the level of the
rhinophores, the others on the dorsum.
A typical doridian branchial plume
is present on the dorsal median, at two
thirds of posterior body length.
Basic colour of head, dorsum,
metapodium, stalk of rhinophores, stalk
of velar and dorsal processes is creamy-
yellow. Surface of dorsum and flanks
can be speckled (not in all specimens)
with small, randomly dispersed, whitish
warts (or minute tubercles) of variable
size and number.
Clavus of rhinophores is greenish-
yellow, somewhat darker on one of the
specimens. Branchial plume has a some¬
what darker yellow tinge than the body
base colour. Colour of all specimens is
uniform light beige, with no pigment
observed. Head, dorsum, metapodium,
velar and dorsal processes are speckled
with dark-reddish to brown spots in
variable densities and blots.
Oral tentacle on each side of head,
under oral veil, is fíat, rectangular, with
the two short sides somewhat rounded,
and is afixed to the body wall by one of
its longer sides. Ratio length to width =
2.5 to 3. (Fig. 4).
Rhinophore stalks are generally
retracted in mantle pocket; clavus (lamel-
lated cone) is protruding 50 to 100% from
the dorsum and bears between 14 and 31
thin complete or partial lamellae.
The observed number of lamellae is
dependent on the integrity (intact, flat-
tened) and degree of contraction and
retraction of the clavus.
One sample (N0796/07) shows a
circle-shaped ripple around the
rhinophore's base.
On the dorsum sides, a faint pallial
ridge can sometimes be seen, starting
behind the rhinophores and fading away
besides the branchial plume. The first
pair of the 5 dorsal branched processes is
located just under and slightly behind the
rhinophore basis; pairs 2 and 3 (generally
the largest) stand on the pallial ridge,
between the rhinophores and branchial
plume; pair 4 is near the plume and pair
5 is at the rear, between the plume and
metapodium.
The non-retracted branchial plume is
composed of 5 tripinnate gills arranged
in a horseshoe around the anal papilla.
A large genital pore is located on the
right side, under and between the first
and second dorsal branched processes.
Three specimens (N0796/02,
N0796/03 and N0822) show a pair of
tiny symmetrical dark spots embedded
in the dorsal epidermis behind the
rhinophores (Fig. 5).
The apparent absence of some char-
acters (i.e. the absence of the minute
rhinophoral sheath) can be an artefact
resulting from the preserving technique.
58
VANHAELEN ET AL.: Kaloplocamus ramosus new records in the Bay of Biscay
Figure 7. Kaloplocamus ramosus . Radula of specimen N0799/7. A: full view; B: hooked 3rd lateral
tooth (arrow) and dish-like rectangular 4th to n-th lateral teeth; C: three first lateral hooked teeth
(arrows a, b, c). SEM photos by J. Cellis and L. Despontin.
Figura 7. Kaloplocamus ramosus. Rádula del espécimen N0799/7. A: vista completa; B: tercer diente
lateral ganchudo (flecha) y dientes rectangulares y aplanados desde los cuartos; C: los tres primeros
dientes laterales ganchudos (flechas a, b, c). Fotos al MEB por J. Cellis y L. Despontin.
59
Iberus , 32 (1), 2014
The radula (Fig. 7) presents 17 rows of
denticles. In a typical row, from axis to the
outer side, the center (rachis) is free of den¬
ticles; the first to 3rd lateral denticles are
long, narrow hooks; the 4th to the 15,16
or 17th laterals are shallow rectangular
dish-like platelets, giving the radular
formula 17 x [ (15-17)3.0.3. (17-15)].
Diet : Large pieces (from 1.5 to 2.2
mm long) of Bryozoa were found in the
anterior part of the digestive tract of
most dissected specimens (Table II).
Some specimens showed only one
prey-species: Cellaria salicornioides Lam-
ouroux, 1816 (in N0822, 00954, 00961
and 00922) or Caberea boryi (Audouin,
1826) (in 01026) (Fig. 6). One specimen
(N0796) contained the two above-men-
tioned prey-species. The digestive tract
was found empty in four of the dis¬
sected specimens.
DISCUSSION
Taxonomy
According to literature, K. ramosus
has been allocated to 6 different genera
(Caloplocamus, Doris, Euplocamus , Idalia,
Kaloplocamus and Triopa) and cited
under 14 different species ñames
(aureus, capensis, croceus , croceus var.
capensis, filosas , fimbriata, frondosas,
japonicus, longicornis, orientalis, plumosas,
principiswalliae, tristis and yatesi), indi-
cating a highly variable species with a
very large distribution area.
For an extensive review of its taxon¬
omy, we refer to Valles and Gosliner
(2006) and Edmunds (2010).
Anatomy
General shape, body size, colour, dor¬
sal appendages, smooth solé, number of
rhinophore lamellae, 15 to 18 branched
processes, oral tentacles and radula of ob-
served specimens fit well with K. ramosus
(see Cantraine 1835, 1841; Philippi 1836,
1844; delle Chiaje 1841; Bergh 1880
[1879], 1883, 1892, 1899; Vayssiére 1901,
1913; Pruvot-Fol 1951, 1954; Macnae
1958; SCHMEKEL AND PORTMANN 1982;
Cattaneo-Vietti and Sordi 1988; Baba
1989; Orte a-Rato, Moro- Abad and Ca-
Distribution and ecology : Sampling
stations of the Bay of Biscay where
K. ramosus specimens were collected
cover an area of about 350 km long and
70 km wide, oriented along a SE-NW
axis, some 120 km southwards off Brit-
tany's south western coast, between 45°
12' 39.5" N - 02° 32' 29.2" W and 47°
23' 51.5" N - 05° 14' 28.2" W (Fig. 1 and
Table I).
Collected specimens were trawled
on the Bay of Biscay and Celtic Sea con¬
tinental shelves from depths ranging
from 128 m (station 01025) to 185 m
(station N0820) on soft bottoms (ranging
from sand to muddy sand) (Fig. 1).
Bottom temperatures during collection
ranged from 11.19 to 11.85° C. Whatever
the season, mean bottom temperatures
in that area are rather stable, ranging
from 10.8 to 11.9° C.
baller-Gutiérrez 2001 and Valles and
Gosliner 2006).
The number of pallial and velar
processes might be size-dependent:
Vayssiére (1901) mentions 3-4 processes
per side for "juveniles", 5 processes for
25-38 mm long specimens and 6 for the
"largest".
Taxonomic confusión of K. ramosas is
certainly related, at least partly, to the
large variability of its colour patterns,
going from creamy to brick red.
However, there is apparently no re-
lationship between colour patterns
and distribution. The colour range of
K. ramosus varies widely from translu-
cent white with yellow-orange spots
(Rudman, 2010a in Hong Kong) to palé
grey with light orange-red spots
(Edmunds 2010: Fig. 1-F, in Ghana) to
palé yellow cream with orange vari-
ously dispersed speckles (Baba 1949: pl.
XIII, Figs 46-47, in Japan) or with dark-
brown speckles (our specimens in
Atlantic France, Fig. 2) to almost
uniform bright red with minute light
yellow-orange spots (Wirtz and Debe-
lius 2003: 214, in the Azores). No partic¬
ular colour pattern seems to be locality-
60
Vanhaelen ET AL.: Kaloplocamus ramosus new records in the Bay of Biscay
dependent and different patterns are
sometimes observed simultaneously at
the same location.
Other Kaloplocamus species (e.g.
K. acutus ) also present a large colour
variability with stable external morpho-
logical characters such as the branching
processes (Baba 1955; Bolland 2006a, b
and Rudman 2010a, b).
The two tiny dark spots embedded
in the epidermis of the dorsum, behind
the rhinophores, are probably rudimen-
tary eyes. They have been mentioned by
DELLE Chiaje (1841 (2): 29) (as Dorisfim-
briata), Bergh (1883: 142) and Vayssiére
(1901: 69, pl. V, Fig. 2) and drawn by
Baba (1949 pl. XIII Figs 46, 47). Several
photographs of live specimens (not col¬
lected) showing the eyes can be viewed
on Pitmann and Fienne's, and on
Johnson'' s websites.
Diet
The diet of K. ramosus is usually
broadly referred to as "bryozoans".
Eight bryozoans have been identified
by different authors to species level
(Table II), all from the Mediterranean
Sea. Caberea boryi (Audouin, 1826) is
also found in our specimen (N0796/08).
Cellaria salicornioides Lamouroux, 1816
is a new known prey in the diet of
K. ramosus. Our observations and data
from literature Usted in Table II confirm
that K. ramosus feeds mostly on branch¬
ing Bryozoans.
The restricted number of prey-
species known so far is almost certainly
an artefact, being only reported from the
Mediterranean Sea and the Bay of
Biscay.
Distribution
K. ramosus is recorded for the first
time in the Bay of Biscay. The large area
where specimens were collected during
the EVOHE 2009 and 2010 cruises and
their large number (89) indicates that we
are dealing with a well established pop-
ulation. The Bay of Biscay extends the
distribution of K. ramosus, a températe,
sometimes tropical, species to its most
northerly known latitude, at the north-
ern limit of the Lusitanian Province.
According to literature and websites,
the species is nearly cosmopolitan but
not yet mentioned from the Americas
and the Philippines.
The present worldwide distribution
of K. ramosus and its potential taxo-
nomic confusión (14 synonyms recog-
nized) leads us to suspect that we are
dealing either with a highly variable
species or with a species complex. As
already suggested by Valles and
Gosliner (2006), Rudman (several
SEASLUGFORUM messages) and
Edmunds (2010), these hypotheses
should be tested via DNA analysis of
material proceeding from the different
areas.
Ecology
Collecting depths (128-185 m) of our
material are in agreement with those
reported in the literature (5-400 m)
(Wirtz 1998: 12; Edmunds 2010: 296;
Koutsoubas, Tselepides and Elefthe-
riou 2000: 91; Zsilavecz 2007).
Kaloplocamus ramosus is known from
soft bottoms (Pruvot-Fol 1951; Wirz-
Mangold and Wyss 1958; Rós 1975;
Koutsoubas et al. 2000; Doménech,
Avila and Ballesteros 2006; present
study) and hard bottoms (Avíla,
Azevedo, Gonqalves, Fontes and
Cardigos 1998; García-Gómez 2002;
Cervera, Calado, Gavaia, Malaquias,
Templado, Ballesteros, García-
Gómez and Megina 2004; Malaquias,
Calado, Padula and Cervera 2009;
Rudman 2010a; Bolland 2006;
Pittman AND Fiene 2010; SURG 2007;
Wirtz and Debelius 2003). This differ-
ence is mainly related to the collecting
methods. Specimens collected by divers
have been found on rocky bottom or
wrecks, or in caves whereas dredging
and bottom trawling is performed on
soft bottom. The species is reported
from moderately to highly exposed
rocky habitats (Pittman and Fiene
2010) as well as from modérate current
areas (our study). K. ramosus is active
day and night, under ledges or stones
(Johnson 2005, Wirtz and Debelius,
2003, Pittman and Fiene 2010, Mis-
senden 2005).
61
Iberus , 32 (1), 2014
AKN O WLEDGEMENTS
We are grateful to the crews of the r/v
Thalassa who helped in the data collection
during the EVHOE surveys. One of us
(AV) wishes to thank Therry Backeljau
and Rose Sablón (both at RBINS) for their
support, access to literature and lab facili-
ties; William Rudman (Australian
Museum) for his early comments on the
Identification of K. ramosus; Frédéric
André (DORIS - Données d'Observations
pour la Reconnaissance et l'Identification
de la faune et de la flore Subaquatiques,
Fédération Frangaise d'Etudes et de
Sports Sous-Marins) for references about
the prey of K. ramosus ; Bernard Picton for
improving the English, Rocío Lopez-
Alonzo and Jesús Troncoso for improving
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64
© Sociedad Española de Malacología
Iberus , 32 (1): 65-85, 2014
Nomenclatura! notes on some European marine bivalve
species
Apuntes nomenclaturales sobre algunas especies de bivalvos de
Europa
Rudo von COSEL*, Serge GOFAS** & Jean-Maurice POUTIERS*
Recibido el 6-XI-2013. Aceptado el 17-1-2014
ABSTRACT
Some nomenclatura! ¡ssues affecting European species are discussed. The following taxa
are treated under 1CZN Art. 23.9, with the required references provided:
- Mytilus variabilis Krauss, 1 848 (currently Brachidontes variobilis (Krauss, 1 848)) is
declared nomen pmtectum against the sénior homonym Mytilus variabilis Fischer von
Waldheim, 1 807, declared nomen oblitum. The still earlier ñame Brachidontes ustulatus
(Lamarck, 1819), currently used as the valid ñame for a native species of Western Aus¬
tralia, should take precedence over 8. variabilis (Krauss, 1 848) were it demonstrated that
it is the same biological species, but in the current state of knowledge it is proposed to
keep them sepárate.
- Modiola nigra Cray, 1 824 (currently Musculus niger (Cray, 1 824)) is declared nomen
protectum against the sénior synonym Mytilus discors svecicus Fabricius, 1788, declared
nomen oblitum.
- Ostrea flexuosa Poli, 1795 (currently Flexopecten flexuosus (Poli, 1795)) is declared
nomen protectum against the sénior synonym Ostrea coarctata Born, 1778, declared
nomen oblitum.
- Chama aculeata Poli, 1795 (currently Centrocardita aculeata (Poli, 1795)) is declared
nomen protectum against the sénior homonym Chama aculeata Stróm, 1768, declared
nomen oblitum , thereby making valid the current usage and avoiding the need for using
the júnior synonym Centrocardita elegans (Requien, 1 848).
- Solen marginatus Pulteney, 1799 is declared nomen protectum against the sénior syn-
onyms Hypogaea tentaculata Poli, 1791, Solen rotundatus Spengler, 1794 and Solen gla-
dius Róding, 1798, all declared nomina oblita.
The following cases are discussed but do not meet the conditions for application of ICZN
Art. 23.9:
- Anadara polii (Mayer, 1868) is in prevailing usage over Anadara gibbosa (Reeve,
1 844) but the latter ñame has been employed once in the XX century so must be used.
- Chama gryphina Lamarck, 1819 (currently Pseudochama gryphina (Lamarck, 1819)), is
not a júnior synonym of Chama cornuta Dillwyn, 1817. The latter ñame is made an objec-
tive synonym of Chama bicornis Linnaeus, 1758 by lectotype designation.
- Venus rhomboides Pennant, 1777 (currently Polititapes rhomboides (Pennant, 1777))
should not be superseded by Venus virgínea Linnaeus, 1767. The extant types of Venus
* Muséum National d’Histoire Naturelle, Département Systématique et Evolution (Case Póstale 51), 55, Rué
Buffon, F-75231 Paris Cedex 05, France.
** Departamento de Biología Animal, Facultad de Ciencias, Universidad de Málaga, Campus de Teatinos - E-
29071 Málaga, Spain. E-mail: sgofas@uma.es
65
Iberus , 32 (1), 2014
virgínea are here figured. Two of them belong to the species currently known as Venerupis
geographica (Gmelin, 1791) and one to Venerupis aurea (Gmelin, 1791). Therefore, the
suppression of the ñame virgínea seems the best outcome.
- Ensis arcuatus (Jeffreys, 1 865) and Salen ensis majar Chenu 1 843 are both held as
júnior synonyms of Ensis magnus Schumacher, 1817, therefore the ñame Solen siliqua var.
arcuata Jeffreys, 1 865 does not need to be declared nomen protectum.
- Neaera bicarinata Jeffreys, 1 882 does not need to be declared a nomen protectum
against Neaera striata var. bicarinata Jeffreys, 1876, which is a nomen nudum , therefore
not available.
RESUMEN
Se discuten algunas cuestiones de nomenclatura que afectan a especies europeas. A los
siguientes taxones se aplican las disposiciones del Art. 23.9 del CINZ, aportando las
referencias requeridas:
- Mytilus variabilis Krauss, 1 848 (actualmente Brachidontes variabilis (Krauss, 1 848)) se
declara nomen protectum frente al homónimo más antiguo Mytilus variabilis Fischer von
Waldheim, 1 807, declarado nomen oblitum. El nombre aún anterior Brachidontes ustu/a-
tus (Lamarck, 1819), actualmente en uso como el nombre válido para una especie de Aus¬
tralia Occidental, debe prevalecer sobre 8. variabilis (Krauss, 1848) siempre que se
demuestre que se trata de la misma especie biológica. En el estado actual de los conoci¬
mientos, se propone mantenerlas separadas.
- Modiola nigra Gray, 1 824 (actualmente Musculus niger (Gray, 1 824)) se declara nomen
protectum frente a Mytilus discors svecicus Fabricius, 1788, declarado nomen oblitum.
- Ostrea flexuosa Poli, 1 795 (actualmente Flexopecten flexuosus (Poli, 1 795)) se declara
nomen protectum frente al sinónimo más antiguo Ostrea coarctata Born, 1778, declarado
nomen oblitum.
- Chama aculeata Poli, 1 795 (actualmente Centrocardita aculeata (Poli, 1 795)) se declara
nomen protectum frente al homónimo más antiguo Chama aculeata Stróm, 1768, decla¬
rado nomen oblitum, con lo cual se convalida el uso actual y se evita la necesidad de utili¬
zar el sinónimo más reciente Centrocardita elegans (Requien, 1 848).
- Solen marginatus Pulteney, 1799 es declarado nomen protectum frente a los sinónimos
más antiguos Solen rotundatus Spengler, 1794 y Solen gladius Róding, 1798, todos ellos
declarados nomina oblita.
Los siguientes casos se discuten, pero no cumplen los requisitos de aplicación del art 23.9
del CINZ:
- Anadara poli i (Mayer, 1 868) está en uso predominante sobre Anadara gibbosa (Reeve,
1844), pero este último nombre se ha utilizado una vez en el siglo XX y debe ser utili¬
zado.
- Chama gryphina Lamarck, 1819 (actualmente Pseudochama gryphina (Lamarck, 1 819)),
no es un sinónimo más reciente de Chama cornuta Dillwyn, 1817. Este último se hace
sinónimo objetivo de Chama bicornis Linnaeus, 1758 por designación de un lectotipo.
- Venus rhomboides Pennant, 1777 (en la actualidad Polititapes rhomboides (Pennant,
1777)) no debe ser sustituido por Venus virgínea Linnaeus, 1767. Se figuran los tipos
existentes de Venus virgínea. Dos de ellos pertenecen a la especie actualmente conocida
como Venerupis geographica (Gmelin, 1791) y uno es Venerupis aurea (Gmelin, 1791).
Por lo tanto, la supresión del nombre virgínea parece lo más apropiado.
- Ensis arcuatus (Jeffreys, 1 865) y Solen ensis major Chenu 1 843 se consideran como
sinónimos más recientes de Ensis magnus Schumacher, 1817. Por lo tanto, no es necesa¬
ria la declaración de Solen siliqua var. arcuata Jeffreys, 1 865 como nomen protectum.
- Neaera bicarinata Jeffreys, 1 882 no necesita ser declarado nomen protectum frente a
Neaera striata var. bicarinata Jeffreys, 1 876, que es un nomen nudum, por lo tanto, no
está disponible.
66
COSEL ET AL.: Nomenclatura! notes on some European marine bivalve species
INTRODUCTION
The rules of zoological nomenclature
regarding priority and homonymy are
intended for preserving the stability of
ñames, and in most cases are instrumen¬
tal in doing so. There are nevertheless
some instances where undesirable
changes may be mandatory when oven
looked sénior synonyms or homonyms
are brought to attention. Digging out
forgotten ñames was legitímate under
the previous editions of the Code, and
one of the most troublesome cases in the
European marine fauna has been the
revival of limpet ñames Patella nigra, P.
ulyssiponensis and P. rustica against
respectively Patella safiana, P. aspera and
P.lusitanica all universally in usage at
that time (Christiaens, 1973). Fortu-
nately, the 1999 edition of the Code
(ICZN, 1999, Art. 23.9) has provisions
for the conservation of usage when
certain conditions are met.
The recent publication of the "Com-
pendium of Bivalves" (Huber, 2010) has
been a landmark in malacology, and
provides for the first time a global
overview of species-level taxonomy in
this class. Precisely because it is compre¬
hensivo, it is also influential and, for
instance, has been taken as the default
standard in the World Register of
Marine Species, the leading database for
marine biodiversity data. Some changes
in nomenclature are suggested therein
on the grounds that earlier ñames which
interfere as synonyms or homonyms
have been overlooked so far and should
be used.
The purpose of this paper is to docu-
ment the usage which can lead to the
application of Art. 23.9, and to discuss
some addítional cases for which,
although the requirements of this article
are not met, a conservativo treatment is
considered desirable for the stability of
nomenclature. Marshall & Spencer
(2013), in a similar approach, discussed
the nomenclatura! changes which are
derived from Huber (2010) affecting the
New Zealand fauna and propose the
rebuttal of some of them, but without
declaring any nomen oblitum or nomen
protectum. Huber (2010) declared eight
ñames as nomina protecta and their
sénior synonyms or homonyms nomina
oblita but these actíons are invalid
because they lack citation of the refer-
ences of usage required by ICZN Art.
23.9; three of these ( Solen arcuatus Jef-
freys, 1865, Chama gryphina Lamarck,
1819 and Cuspidaria bicarinata Jeffreys,
1882) regard the European fauna and
will be discussed herein.
Literature was searched for possible
instances of usage using both the Zoo¬
logical Record and the advanced search
function of Google Books <http: / /
books.google.com/> and Google
Scholar <http: / / scholar.google.com/ >,
eventually checked against the origináis
(digital or paper). We have strived to
inelude references from different fields,
including palaeontology or archaeology,
fisheries, environmental studies as well
as checklists or taxonomic papers.
We have made the option to ignore
the usage by authors after 2000, includ¬
ing Huber (2010) for the ñames to be
dismissed, as the Code reads (our
italics) "23.9.2. An author who discov-
ers that both the conditions of 23.9.1 are
met should cite the two ñames together
and state explicitly that the younger
ñame is valid...". This interpretation is
supported by ICZN Art. 23.9.6. which
reads "The delibérate use of a ñame
contrary to Article 23.9.1 (...) must not
be taken into account in determining
usage".
We have systematically reported the
cases in which species considered here
are the type of a genus-group ñame.
This is not formally an argument in
favour of conserving ñames but still
makes it desirable whenever possible.
Article 23.9 of ICZN is here repro-
duced for guidance :
23.9.1. prevailing usage must be main-
tained when the following conditions are
both met:
23.9.1.1. the sénior synonym or
homonym has not been used as a valid ñame
after 1899, and
67
Iberus , 32 (1), 2014
23.9.1.2. the júnior synonym or
homonym has been used for a particular
taxon, as its presumed valid ñame, in at
least 25 works, published by at least 10
authors in the immediately preceding 50
years and encompassing a span of not less
than 10 years. [i.e. 1963-2012]
23.9.2. An author who discovers that both
the conditions of 23.9.1 are met should cite the
tzvo ñames together and State explicitly that
the younger ñame is valid , and that the action
is taken in accordance zvith this Ardele; at the
same time the author must give evidence that
the conditions of Ardele 23.9.1.2 are met, and
also state that, to his or her knowledge, the
condidon in Ardele 23.9.1.1 applies. From the
date ofpublicadon ofthat act the younger ñame
has precedence over the older ñame. When
cited, the younger but valid ñame may be qual-
ified by the term nomen protectum and the
invalid, but older, ñame by the term nomen
oblitum. In the case ofsubjecdve synonymy,
whenever the ñames are not regarded as syn-
onyms the older ñame may be used as valid.
CASES TREATED UNDER ICZN ART. 23.9
Brachidontes variabilis (Krauss, 1848) vs. Brachidontes pharaonis (Fischer, 1870)
and Brachidontes ustulatus (Lamarck, 1819)
Ñames involved:
Mytilus variabilis Fischer von Waldheim, 1807, Muséum Demidoff, vol. 3: 249.
Mytilus variabilis Krauss, 1848, Die Südafrikanischen Mollusken: 25-26, pl. 2 fig. 5.
Mytilus ustulatus Lamarck, 1819, Histoire naturelle des animaux sans vertebres, vol. 6 (1): 122.
Mytilus pharaonis P. Fischer, 1870, Journal de Conchyliologie, 18: 169.
A small mytilid species was known
from South Africa and also reported as
an invader in the Mediterranean Sea
under the ñame Brachidontes variabilis
(Krauss, 1848) until the mid 1990's,
when it was realized that this is a
primary homonym of Mytilus variabilis
Fischer von Waldheim, 1807. Under the
current code, Art. 23.9 applies since
Fischer von Waldheim's ñame (currently
treated as a júnior synonym of Mytilus
edulis Linnaeus, 1758) has never been
used otherwise than in synonymy. The
ñame variabilis was dismissed in Dekker
& Orlin's (2000) check-list and since
then, the júnior ñame B. pharaonis
(Fischer, 1870) is treated as synonym
and is universally used in the Red Sea
and Mediterranean realm (Engl, 1995;
Buzzurro & Greppi, 1996 and most
later authors). However Huber (2010)
proposed to use for this species a still
earlier ñame Brachidontes ustulatus
(Lamarck, 1819) originally described
from "Brazil".
Huber writes "Lamy (1936) (...) con-
sidered M. pharaonis from the Red Sea,
Suez and arabicus the same. Further-
more, Lamy analysed Lamarck's
MNHN type series of M. ustulatus,
described from Brazil and noted it iden-
tical to variabilis = arabicus= pharaonis."
Nevertheless, knowing this, Lamy still
used B. variabilis as the valid ñame for
this taxon. There is concurrently a con-
sistent usage of Brachidontes ustulatus
(Lamarck, 1819) as the valid ñame of a
taxon in Western Australia (Bindon,
Dortch & Kendrick, 1978; Jones, 2004;
Platell, Ang, Hesp & Potter, 2007;
Scott & Johnson, 1993; Semeniuk,
1997; Wells & Bryce, 1984; Wilson,
1998 among others) and therefore the
question of conserving the ñame M.
variabilis is a taxonomic, not nomenclat¬
ura! issue. The Brazilian type locality is
evidently wrong, as no candidate
species is reported there; many of the
Baudin expedition types are from King
George Sound, Western Australia, and
that is probably where B. ustulatus carne
from (F. Wells, pers. comm.).
A recent molecular study (Sirna Ter-
RANOVA, Lo BRUTTO, ARCULEO &
Mitton, 2007) reveáis that the Red Sea
and Mediterranean populations, the East
African populations and a population
from Hong Kong represent three cryptic
68
COSEL ETAL.: Nomenclatura! notes on some European marine bivalve species
species. This implies separating the Red
Sea B. pharaonis from the East African B.
variabilis and using both ñames as valid.
The final decisión on giving the ñame B.
ustulatus (Lamarck, 1819) precedence
over B. variabilis (Krauss, 1848) depends
on whether the Western Australian
species is the same as that living in the
eastern African coast.
In order to conserve both ñames
potentially valid, we hereby declare,
according to the provisions of ICZN Art.
23.9.2 the ñame Mytilus variabilis Krauss,
1848 as nomen protectum , and Mytilus
variabilis Fischer von Waldheim, 1807, as
nomen oblitum. Usage of Brachidontes
variabilis (Krauss, 1848) as a valid ñame
has been found in the following sources:
Arcidi acono & Di Gerónimo, 1976;
Atapattu, 1972; Barnard, 1964 (p. 395);
Barash & Danin, 1992 (p. 333-334);
Biggs, 1973 (p. 382); Britton, 1990 (p.
816-817); Cheung, Luk & Shin, 2006;
Davies, 1980; Felsenburg & Safriel,
1974; Lavee & Ritte, 1994; Lee &
Morton, 1985; Morton, 1988; Oliver,
1992 (p. 48, 224); Nakhlé, Cossa,
Khalaf & Beliaeff, 2006; Parenzan,
1974 (p. 59); Plaziat, Baltzer,
Choukri, Conchon, Freytet, Orszag-
Sperber, Raguideau & Reyss, 1998 (p.
551); Rajagopal, Venugopalan, Van
der Velde & Jenner, 2005; Safriel,
Felsenburg & Gilboa. 1980; Safriel,
Gilboa & Felsenburg, 1980; Safriel &
Sasson-Frostig, 1988; Sirna Terra-
nova et AL., 2007; Sloan, 1979; Stern &
Achituv, 1978; Tang. 1992; Tringali &
Villa, 1989 (p. 35); Ünsal, 1984; Valen-
tich-Scott, 2003 (p. 265, with addi-
tional references cited therein); van
Aartsen & Kinzelbach, 1990.
Musculus niger (J.E. Gray, 1824) vs. Musculus svecicus (Fabricius, 1788)
Ñames involved:
Mytilus discors Linnaeus — (3 svecicus Fabricius, 1788, Nye Samling af det Kongelige Danske Videns-
kabers Selskabs Skrifter, 3: 460.
Modiola nigra Gray, 1824, Supplement to Appendix, Parry's Voy age for the Discovery of a north-west
passage in the years 1819-1820, p. ccxliv.
This is a common boreal species,
widely distributed in the North Atlantic,
Arctic and North Pacific oceans, and of
importance in trophic webs. Fabricius
(1788) introduced the ñame in the form
Mytilus discors , L. — (3 svecicus , based on
figures 766-767 of Chemnitz, 1785 (vol.
8, p. 191-195, pl. 86 fig. 766-767, with
type locality in the Kattegat); he did not
use the term "variety" and a subspecies
concept was not elaborated at that time,
but it is clear that the ñame is available
with a subspecific rank in the sense of
ICZN. Modiola nigra Gray, 1824 is based
on the figure given by Montagu (1808:
pl. 26 fig. 4) of a specimen from Laskey,
collected in the Frith of Forth (Scotland)
and wrongly assigned to his Mytilus dis -
crepans Montagu, 1803. The ñame has
consistently been applied since its publi-
catión and is widely used in modern lit¬
era tu re. In the XX century, to our knowl-
edge only Dautzenberg & Fischer
(1912), and Lamy (1937) mentioned the
ñame svecicus but nevertheless used
Modiolaria nigra or Musculus niger, as
valid. In application of ICZN Art. 23.9.2,
we declare the ñame Mytilus discors sve¬
cicus Fabricius, 1788 as a nomen oblitum,
and Modiola nigra Gray, 1824, a nomen
protectum. Usage of the ñame Musculus
niger as valid was documented in the
following sources: Aitken & Gilbert,
1996; Bahr & Gulliksen, 2001; Cham-
bers, 2009 (p. 131); Denisenko, Rachor
& Denisenko, 2003; Fenchel, 1964;
Gilbert, 1977; Jewett & Feder, 1980;
Kendall, 1996.; K dra, Gromisz,
Jaskula, Lege y ska, Maciejewska,
Malec, Opanowski, Ostrowska, Wlo-
DARSKA-KOWALCZUK & W SLAWSKI,
2010; Króncke, 1994; Lande, 1975;
Larsen, Frischer, Rasmussen &
Hansen, 2005; McCormick-Ray,
Warwick & Ray, 2011; Mohlenberg &
Riisgárd, 1978; Muus, 1973; Nordsieck,
69
Iherus , 32 (1), 2014
1969 (p. 36); Norton, 1975; Pollock,
1998 (p. 157); Rózycki, 1992; Schiotte,
1989; Bífero & Welch, 1992; Strand,
Jacobsen, Pedersen & Granmo, 2003;
Tandberg, Schander & Pleijel, 2010;
Tebble, 1966 (p. 47).; Tyler, 1972; Vader
& Beehler, 1983; Wagner, 1977;
Wildish & Peer, 1983.
Flexopecten flexuosus (Poli, 1795) vs. Flexopecten coarctatus (Born, 1778)
Ñames involved
Ostrea coarctata Born, 1778, Index Rerum Naturalium Musei Caesarei Vindobonensis, 1: 90-91.
Ostrea flexuosa Poli, 1795, Testacea Utriusque Siciliae, vol. 2: 161, pl. 28 fig. 11.
Flexopecten flexuosus is a common
species found in the Mediterranean and
the Atlantic Ibero Moroccan area. Ostrea
flexuosa Poli, 1795 is the type species of
Flexopecten Sacco, 1897 by original desig¬
naron. To our knowledge, the ñame
coarctatus was used (in the original com-
bination Ostrea coarctata ) by Brocchi,
1814 (574, pl. 14 fig. 9) and later cited
under the combina tion Peden coarctatus
by Defrance (1824) but, possibly
because of the influence of Sacco's
(1897) monograph, we could not trace
any usage as the valid ñame in the XX
century. Usage of flexuosus is wide-
spread and we therefore consider that
the conditions are met for application of
ICZN Art. 23.9.2 and declare the ñame
Ostrea flexuosa Poli, 1791 as nomen protec-
tum, and Ostrea coarctata Born, 1778,
nomen oblitum. Dijkstra (2009) already
proposed this but did not provide the
references required by the Code.
Usage as valid ñame in the combina-
tion Flexopecten flexuosus (unless other-
wise noted) was found in the following
sources: Biagi & Corselli, 1978 (p. 11);
Dijkstra & Goud, 2002 (p. 60-61); Dijk¬
stra, 2009 (p. 105, 110-111); Giannuzzi-
Savelli, Pusateri, Palmeri, Ebreo,
Coppini, Margelli & Bogi, 2001 (p. 182-
183); Giribet & Peñas, 1997 (p. 57);
Gofas, 2011a (p. 578); Margus, 1991;
Nordsieck, 1969 (p. 53); Óztürk,
Buzurro & Benli, 2003 (p. 64); Paren-
zan, 1974 (p. 112-113); Peña, Canales,
Adsuara & Sos, 1996; Rico-García,
1988 (p. 113); Rolán, 2011 (p. 323); Rom-
BOUTS, COOMANS & VAN PEL, 1991 (p.
40); Rueda, Salas & Gofas, 2000 (p.
104); Schiaparelli, 2008; in the combi-
nation Chlamys flexuosa or Chlamys (Flex¬
opecten) flexuosa : Cachia, Mifsud &
Sammut, 2004 (p. 54); D' Angelo &
Gargiullo, 1978 (p. 178); Lucas, 1980
(p. 164); Montero Agüera, 1971 (p.
171); Poppe & Goto, 1993 (p. 61);
Pelorce, 2007 (p. 47); Pérés & Picard,
1964 (p. 85, 87); Salas, 1996 (p. 55);
Sileti , 2006; Zenetos, 1997 (p. 464).
Centrocardita aculeata (Poli, 1795) vs. Centrocardita elegans (Requien, 1848)
Ñames involved
Chama aculeata Poli, 1795: Testacea Utriusque Siciliae , vol. 2: 122, pl. 23 fig. 22.
Chama aculeata Stróm, 1768: Det Kongelige Norske Videnskabers Selskabs Skrifter, 4: 368, pl. 16 fig. 4.
Cardita elegans Requien, 1848: Catalogue des coquilles de Tile de Corsé: 27.
Chama aculeata Poli, 1795 is the type
species of genus Centrocardita Sacco,
1899. The ñame has been displaced for
being a primary homonym of Chama
aculeata Stróm, 1768 (synonym of
Hiatella árctica) which has never, to our
knowledge, been used later as the valid
ñame of a taxon. We therefore consider
that the conditions are met for applica¬
tion of ICZN Art. 23.9.2 and declare the
ñame Chama aculeata Poli, 1795 as nomen
protectum, and Chama aculeata Stróm,
1768, nomen oblitum. With this statement,
Cardita elegans Requien, 1848 is to be
treated as a subjective júnior synonym
of Centrocardita aculeata (Poli, 1795).
70
COSEL ET AL.: Nomenclatura! notes on some European marine bivalve species
There is a limited usage of the ñame
elegans (in the combination Glans
elegans) in the recent literature (Cachia,
Mifsud & Sammut, 1993: 59; Cachia,
1999; Óztürk et al., 2003: 65; Repetto,
Orlando & Arduxno, 2005; Cachia,
Mifsud & Sammut, 2004: 87) but this
does not interfere with restoring C.
aculeata , because the ñame declared as
nomen oblitum is the sénior homonym of
C aculeata (Poli, 1795), not the sénior
synonym Car dita elegans.
Usage of the ñame aculeata (in the
combination Glans aculeata unless other-
wise stated) has been found in the fol-
lowing sources: Albayrak, Balkis &
Balkis, 2004; Altimira, 1977 (as Car dita
(Glans) aculeata ); Bakir, Óztürk, Doan
& Ónen, 2012; Bernasconi & Stanley,
1997; Borg, Howege, Lanfranco,
Micallef, Mifsud & Schembri, 1998 (p.
19); Dhora, 2009; Giribet & Peñas, 1997
(p. 58); Gofas, 2011b (p. 595); Koulouri,
Dounas, Arvanitidis, Koutsoubas &
Eleftheriou, 2006; La Porta, Targusi,
Lattanzi, La Valle, Paganelli &
Nicoletti, 2009; M alaquias, Bentes,
Erzini & Borges, 2006; Manousis,
Mpardakis, Paraskevopoulos &
Galinou-Mitsoudi, 2010 (p. 166);
Marasti, 1973 (p. 106-107, as Glans
(Centrocardita) aculeata ); Meco Cabrera,
1982 (p. 102, as Cardita aculeata );
Montero Agüera, 1971 (p. 155, as
Beguina (Mytilicardita) aculeata ); Nord-
sieck, 1969 (p. 74, as Cardita (Glans)
aculeata ); Pancucci-Papadopoulou,
Simboura, Zenetos, Thessalou-Legaki
& Nicolaidou, 1999; Parenzan, 1974
(p. 153, as Cardita aculeata ); Pérés &
Picaro, 1964 (p. 94, as Cardita aculeata );
Salas, 1996 (p. 62); Schiaparelli, 2008;
Sileti , 2006 (p. 143); Tornaritis, 1987
(p. 147, as Cardita aculeata ); Trapani,
Scotti, Gianguzza, Chemello &
Riggio, 1999; Zenetos, 1996 (p. 125);
Zenetos, 1997 (as Glans (Centrocardita)
aculeata).
Solen marginatus Pulteney, 1799 vs. Solen rotundatus Spengler, 1794, Solen
gladius Roding, 1798 and Hypogaea tentaculata Poli, 1791
Ñames involved:
Solen marginatus Pulteney, 1799, Catalogue of the birds , shells , and some of the more rare plants of
Dorsetshire: 28.
Hypogaea tentaculata Poli, 1791, Testacea Utriusque Siciliae, vol. 1: 16-17.
Solen rotundatus Spengler, 1784, Skrivter af Naturhistorie-Selskabet, Kiobenhavn, 3 (2): 86-87.
Solen gladius Roding, 1798, Museum Boltenianum : 154.
The solé European representative of
the family Solenidae has been alterna-
tively known under the ñames Solen
vagina Linnaeus, 1758 or Solen margina¬
tus Pulteney, 1799. The former ñame has
been often used in the XIX century but
is now abandoned in the European liter¬
ature for taxonomic reasons. It has been
established that Linnaeus confused two
different species under the ñame vagina ,
which should apply to the Indo-Pacific
species, rather than to the European
Solen marginatus Pulteney, 1799.
Bucquoy, Dautzenberg & Dollfus
(1895: 498) give a complete listing of XIX
century usage.
There are however three other ñames
which predate Solen marginatus and have
a European type locality. Solen rotundatus
Spengler, 1794 is based on Lister (1692,
pl. 1056, fig. 5) and is stated as Mediter-
ranean. It was listed by Mórch (1871) as
being Solen vagina as understood at that
time. Other wise than through the State-
ment of origin, Lister 's figure could not
be definitely assigned to one of the
species involved. The ñame had defi¬
nitely fallen into oblivion when Huber
(2010) proposed that it should supersede
S. marginatus. Solen gladius Roding, 1798
is based on Chemnitz (1782, vol. 6, pl. 4
fig. 27) with Solen vagina given in syn-
onymy and, to our knowledge, has never
been used later as a valid ñame.
Poli (1791) used for this species the
ñame Solen vagina , as usual in his time.
71
Iherus, 32 (1), 2014
but, in his rather queer system of paral-
leí nomenclature for the soft parts and
the shells, named these soft parts
Hypogaea tentaculata . Such ñames are
here treated as ñames introduced in syn-
onymy, and therefore not available if
never used as valid which is the case
here (cited by Nordsieck, 1969: 145, in
synonymy of Solen marginatus). The
cardiid genus Cerastoderma, which was
introduced in the same conditions, is
widely used nowadays as valid.
An additional reason for preserving
the stability of the ñame Solen
marginatus is that this ñame is used in
local fisheries regulations (e.g. Ministé-
ríos da Defes a Nacional, das
Finanzas, da Justina, da Agricul¬
tura, Pescas e Alimentado, das
Obras Públicas, Transportes e Comu-
NICAQÓES, DA SaÚDE E DO COMÉRCIO E
Turismo, 1989; Consejería de Agricul¬
tura y Pesca, 2009; Consellería del
Medio Rural y del Mar, 2012). There¬
fore, the ñame Solen marginatus Pul-
teney, 1799 is declared nomen protectum
against Solen rotundatus Spengler, 1794,
Solen gladius Róding, 1798 and Hypogaea
tentaculata Poli, 1791, all declared
nomina oblita.
Usage of Solen marginatus as a valid
ñame was found in the following
sources: Bakir et al., 2012 (p. 180);
Cachi a, Mifsud & Sammut, 2004 (p.
105); Cosel, 1993; da Costa & Martínez-
Patiño, 2009; Eales, 196 (p. 164); Fer-
nández-Tajes & Méndez, 2007; Giribet
& Peñas, 1997 (p. 59); Gutiérrez
Zugasti, 2010 (p. 341); ITayward &
Ryland, 1995 (p. 616); Hodgson,1984;
López, Rodríguez & Carrasco, 2005,;
López-Flores, Garrido-Ramos, de la
Herran, Ruiz-Rejón, Ruiz-Rejón &
Navas, 2008; Manousis et al., 2010. (p.
167); Marina & Urra, 2011 (p. 690);
Meijer, 1993; Montero Agüera, 1971 (p.
224); Nordsieck, 1969 (p. 145, erro-
neously credited to Pennant, 1777);
Óztürk,Buzurro & Benli, 2003 (p. 65);
Parenzan, 1974 (p. 364); Pérés &
Picard, 1964 (p. 58, 113); Remacha-
Triviño, 2005; Riedl, 1983 (p. 372); Schi-
aparell, 2008; Tirado, Rodríguez,
Bruzón, López, Salas & Márquez,
2002; Vale & Sampayo, 2002.; Zenetos,
1996 (p. 125); Zenetos, 1997 (p. 466).
CASES WHICH CANNOT BE TREATED UNDER ART. 23.9 OF ICZN
Anadara gibbosa (Reeve, 1844) vs. Anadara polii (Mayer, 1868)
This is a moderately common
species occurring on the outer shelf of
the Mediterranean and the Eastern At¬
lantic from the Iberian Península to An¬
gola. Until the mid 1990s, most authors
(e.g. Nordsieck, 1969; Parenzan, 1974;
Poppe & Goto, 1993) misidentified this
species as Anadara diluvii (Lamarck,
1805) and therefore did not use the spe-
cific ñame polii. This usage persisted
even later (e.g. Zenetos, 1997; Giribet &
Peñas, 1997), although it is now widely
accepted (Oliver & Cosel, 1993; Huber,
2010) that A. diluvii is a different, extinct
species with a type locality in the
Miocene of the Loire basin, France (and
type species of Diluvarca Woodring,
1925 by original designation).
Arca gibbosa Reeve, 1844 was
described from unknown locality and
appears to be a sénior synonym of Arca
polii Mayer, 1868. There is one syntype
(reg. n° 1969216) in The Natural History
Museum, London, recorded by Steven-
son (1972) who did not, however, relate
it to Anadara polii.
There is unfortunately one instance
of a usage of the specific ñame gibbosa
Reeve in the XX century (van Beek,
1969), within an archaeological publica-
tion regarding South Arabia. The figures
given (his pl. 56a) would fit Anadara
natalensis (Krauss, 1848), a well-known
species ranging from the Red Sea and
Persian Gulf to South Africa and intro¬
duced in the Eastern Mediterranean,
rather than Anadara polii. Albeit a
misidentification, it remains that for
nomenclatura! purposes it preeludes the
application of ICZN Art. 23.9. We could
72
COSEL ET AL.: Nomenclatura! notes on some European marine bivalve species
document the 25 usages of the epithet
polii , but hardly any more, so that it is
not realistic to make a case for request-
ing the suppression of gibbosa. There-
fore, we endorse Huber's (2010) view of
using Anadara gibbosa.
Arca gibbosa Reeve, 1844 is a sénior
homonym of Arca gibbosa d'Orbigny, 1845
(Paléontologie Frangaise, Crétacé III, p.
224, pL 316 fig. 5-8), a Mesozoic fossil
species for which Nyst (1848) coined the
replacement ñame Arca sarthacensis.
Chama cornuta Dillwyn, 1817 vs. Chama bicornis Linnaeus, 1758 or Chama
gryphina Lamarck, 1819 (Figure 1)
Ñames involved:
Chama bicornis Linnaeus, 1758, Systema Naturae, ed. 10: 692.
Chama cornuta Chemnitz, 1784, Systematisches conchylien-Cabinet, vol. 7: 150, pl. 52 fig. 516-520
(unavailable nomenclaturally: ICZN Direction 1).
Chama cornuta Dillwyn, 1817, A descriptive catalogue of Recent shells: 222.
Chama gryphina Lamarck, 1819, Histoire naturelle des animaux sans vertebres, vol. 6 (1): 97.
The well-known ñame Chama
gryphina Lamarck, 1819, currently
Pseudochama gryphina (Lamarck, 1819),
has been declared, by Huber (2010),
nomen protectum against Chama cornuta
Dillwyn, 1817. Flowever this action was
backed by references cited by Lamy
(1928) whereas Art. 23.9.2 requires refer¬
ences spanning not less than ten years in
the immediately preceding 50 years (i.e.
since 1963), therefore it is invalid.
Consultation of the original refer¬
ences involved revealed a much more
complex issue, rendering pointless the
citation of 25 usages of the ñame
gryphina which we could easily find.
The ñame Chama cornuta Dillwyn, 1817
is based on a suite of references among
which Chama cornuta Chemnitz, 1784
(non-binominal, therefore unavailable)
and Chama bicornis Linnaeus, 1758. The
species is stated as inhabiting the
Mediterranean, reproducing Linnaeus'
(1758) indication for Chama bicornis.
Chemnitz's original figures show
chamids, some of which stated as col-
lected in the Indian Ocean, and all of
which have an upper valve coiling
counterclockwise. Therefore Chemnitz's
specimens are real Chama, not
Pseudochama, and Chama cornuta, if
based on those figures, cannot be a
sénior synonym of Pseudochama gryphina
(Lamarck, 1819).
Although not formally a replace¬
ment ñame, the epithet cornuta is meant
by Dillwyn (1817) to stand in place of
bicornis, considered improper because
only one valve, not both, have horn-like
projections. Both ñames are based on
shared cited references (see below), so
that we consider appropriate to treat
Chama cornuta Dillwyn, 1817 as an objec-
tive synonym of Chama bicornis Lin¬
naeus, 1758.
Chama bicornis Linnaeus, 1758 is
itself a problematic taxon. It is based on
three references to pre-Linnean authors:
Caput 19 [pp. 29-30] of Fabius
Columna (1616); pl. 214 fig. 49 of Lister
(1687) and p. 174 [misquoted "274" in
Linnaeus, 1758], pl. 12 fig. 87-88 of
Klein (1753), the two latter being copied
from Columna (1616). All these refer¬
ences are repeated by Dillwyn (1817) in
addition to the reference to Chama bicor¬
nis Linnaeus, 1758. In order to make the
two ñames formally objective syn-
onyms, the specimen figured by
Columna (1616) is here designated as
lectotype of Chama bicornis Linnaeus,
1758 and of Chama cornuta Dillwyn,
1817. The specimens labelled Chama
bicornis in the Linnean collection in
London are believed to be subsequent
(Hanley, 1850: 90; Dodge, 1952: 139)
and the latter author advocated the sup¬
pression of the ñame. There are no regis-
tered specimens of Chama bicornis in
Uppsala (Wallin, 2001). The fact that
Linnaeus coined the ñame " bicornis " on
the assumption that Columna (1616)
73
Iherus , 32 (1), 2014
Figure 1. The three figures cited by Linnaeus (1758) in support oí riis Chama bicornis . Above, in
Fabius COLUMNA (1616, caput 19, page 30, lower figure unnumbered); middle, in LlSTER (1687,
pl. 214 fig. 49); lower, in KLEIN (1753: pl. 12 fig. 87-88) (upper two figures, downloaded from
Google Books; lower figure from Biodiversity Heritage Library) .
Figura 1. Las tres figuras citadas por Linneo (1758), en apoyo de su Chama bicornis. Arriba, en
Fabius Columna (1616, caput 19, página 30, figura inferior sin numerar j, en medio, en LlSTER
(1687, pl2l4 Fig. 49); abajo, en Klein (1753: pl 12 fig 87-88) (dos figuras superiores, descargados de
Google Books, la de abajo desde Biodiversity Heritage Library).
figured both valves and not the inside
and outside of the same valve (see
Dollfus & Dautzenberg, 1932: 302) is
a good indication that he had only that
figure as a source.
The specimen figured by Fabius
Columna is a real Chama , not
Pseudochama, according to the coiling di
rection, and the figures in this book are
not "mirror image" of the original, as
happened in several XVII century books
due to the process of engraving. The pro-
truding item is stated as being a branch
of red coral, not part of the shell itself.
This supports strongly Linnaeus' indica¬
tion of a Mediterranean origin, and sug-
gests that the specimen was collected in
rather deep water. This fits the view ex-
pressed by Pallary (1919), Dautzen¬
berg (1927) and Dollfus and Dautzen¬
berg (1932) that Chama bicornis is an ear-
lier ñame for the Mediterranean Chama
circinata Monterosato, 1884 = C, nicolloni
Dautzenberg, 1392. Dollfus &
Dautzenberg (1932) nevertheless re •
frained from adopting the ñame bicornis
as valid, due to the confusión around it.
The ñame Chama bicornis cannot be
declared nomen oblitum because it has
been used as the valid ñame of a taxon
by Dautzenberg (1927: 304) as a sénior
synonym of his own Chama nicolloni ,
74
COSEL ET AL.: Nomenclatura! notes on some European marine bivalve species
and again by Nordsieck (1969: 96,
authorship erroneously credited to
Gmelin, 1790) in an identification guide.
The latter usage was followed by
Einsele, Elouard, Herm, Kógler &
Schwarz (1977: 11) for West Africa and
Zenetos (1996) for Greece. The outcome
is that the ñame is available, and in the
current stage of knowledge, either it
should be used as a sénior ñame for the
circalittoral Mediterranean species cur-
rently known as Chama circinata Mon-
terosato, 1884, or its suppression
requested to ICZN. There is a possibility
that C. circinata turns out to be a
synonym of C. gryphoides Linnaeus,
1758, which should in this case take
precedence over all those ñames.
Solen ensis major Chenu 1843 vs. Ensis arcuatus (Jeffreys, 1865) and
Ensis magnus Schumacher, 1817
Ñames involved *
Ensis magnus Schumacher, 1817, Es sai d'un nouveau susteme des habitations des vers testacés : 143, pl.
14 fig. la, b.
Solen ensis major Chenu, 1843, Illustrations conchyliologiques, vol. 2: pl. 3 fig. 2, 2a, b, e.
Solen siliqua var. arcuata Jeffreys, 1865, British Conchology, vol. 3: 19.
The taxonomic recognition of Ensis
arcuatus (Jeffreys, 1865) as a distinct
species has come relatively late but is
currently widely accepted. However the
still earlier ñame Ensis magnus Schu¬
macher, 1817, has been adopted as valid
by Cosel (2009), with E. arcuatus treated
as a synonym. The ñame magnus has
been used repeatedly in the XX century
(e.g. Bloomer, 1905; Nordsieck, 1969:
146; van Urk, 1982: 30; Seaward, 1990:
83; Poppe & Goto, 1993: 107) and is by
no means "oblitum". Ensis magnus is the
type species of Ensis Schumacher, 1817,
by monotypy. Huber (2010) proposed to
declare Solen ensis major Chenu 1843
nomen oblitum in order to protect Solen
arcuatus Jeffreys, 1865, but did not com¬
plete the action by providing the 25
instances of usage for S. arcuatus. We
consider however this unnecessary,
since Ensis magnus has precedence. In
the event that Ensis arcuatus should be
recognized as specifically distinct from
Ensis magnus, the latter ñame remains
valid because Solen ensis major would
become a júnior synonym of Ensis
magnus.
Polititapes rhomboides (Pennant, 1777) vs. Polititapes virgineus (Linnaeus,
1767) (Figure 2, Table I)
Ñames involved:
Venus virgínea Linnaeus, 1767, Systema Naturae, ed. 12: 1136.
Venus rhomboides Pennant, 1777: British Zoology, vol. 4: 97.
The ñame Venus virgínea Linnaeus,
1767 was used for a common European
species currently known as Polititapes
rhomboides (Pennant, 1777), by most
British authors in the XIX century as
summarized in Forbes & Hanley (1848:
vol. 1, p. 388, pl. 25 fig. 4, 6) and Jef¬
freys (1864, vol. 2, p. 353-355). Jeffreys
explicitly wrote that this was Tapes vir¬
gineus "of modern authors", probably
not of Linnaeus. Synonymy and reasons
for preferring rhomboides for the Euro-
pean species were thoroughly discussed
in Bucquoy, Dautzenberg & Dollfus
(1893: 397-400). Modern usage of rhom¬
boides as the valid specific ñame was
started by Petit (1851: 297, in the
binomen Pullastra rhomboides ) but was
triggered by the influential work of
Bucquoy, Dautzenberg & Dollfus
and became universal after being
adopted in Winckworth's (1932: 244)
75
Iberus , 32 (1), 2014
Figure 2. The Linnean types of Venus virgínea in the Museum of Evolution, Uppsala. Specimen
1347 (actual length: 21.9 mm) and 1348 (actual length: 18.0 mm) are Venerupis geographica
(Gmelin, 1791); Specimen 1349 is Polititapes aureus (Gmelin, 1791). The Swartz labels and the
modern label are shown below each specimen. Photos courtesy of Erica Mejlon, the Evolution
Museum, Uppsala.
Figura 2. Los tipos de Linneo de Venus virginea en el Museo de la Evolución, Uppsala. Espécimen 1347
(longitud: 21,9 mm) y 1348 (longitud: 18,0 mm) son Venerupis geographica (Gmelin, 1791); Espéci¬
men 1349 es Polititapes aureus (Gmelin, 1791). Las etiquetas de Swartz y la etiqueta moderna se mues¬
tran debajo de cada espécimen. Fotos cortesía de Erica Mejlon, el Museo de la Evolución, Uppsala.
list of British molluscs. Type material of
Venus rhomhoides is stated as still exist-
ing by Smith (1913).
The other taxon which has been
denoted as Venus virginea Linnaeus,
1767 in the XIX century literature is cur-
rently known as Murcia hiantina
(Lamarck, 1818) = flammiculata
(Lamarck, 1818) = rimularis (Lamarck,
1818) = vermiculosa (Lamarck, 1818), and
agrees with the type locality "in Indiis"
stated by Linnaeus (1767). The ñame
Venus virginea was used in this sense by
Philippi (1849: vol. 3, p. 22 (28), pl. 8
figure 2-4), who was very critical over
Lamarck's (1818) taxonomic treatment
of venerids but still distinguished Venus
hiantina Lamarck, 1818 from Venus vir¬
ginea Linnaeus, 1767 (Philippi stated
"not of authors") - rimularis Lamarck,
1818 = vermiculosa Lamarck, 1818 = flam ¬
miculata Lamarck, 1818, all currently
placed in synonymy of Marcia hiantina.
The interference with the European
species seems to originate from the
treatment of Venus virginea in Gmelin's
(1791) edition of the Systema Naturae ,
where the description (p. 3294) is differ-
ent from that of 1767 and a reference to
Chemnitz (1784: vol 7, p. 60, pl. 43 Fig.
457-458) is added. The latter figures
were published with the unavailable
ñame Venus edulis , and Chemnitz wrote
that the species is brought to the Vien-
nese markets from Trieste and Fiume
[now Rijeka, Croada] in the Adriatic sea.
76
COSEL ET AL.: Nomenclatura! notes on some European marine bivalve species
Table I. Number of hits for generic combinations of Venerupis decussata, V. aurea and V. rhomboides
in Google Books (G.B.) and Google Scholar (G.S.), accessed May 20, 2013.
Tabla I. Número de resultados para las combinaciones genéricas de Venerupis decussata, V. aurea y V.
rhomboides en Google Books (G.B.) y Google Scholar (G.S.), consultado el 20 de mayo de 2013.
The specimens on Figs. 457-458 are
unquestionably European, but are more
likely forms of Polititapes aureus
(Gmelin, 1791). Lamarck (1818: 600)
wrote "Les fig. de Chemnitz que cite
Gmelin, me paraissent étrangéres á cette
espéce". Anyway all this does not ínter-
fere nomenclaturally with Linnaeus'
species: either Gmelin's usage is a
misidentification, or if treated as a sepá¬
rate taxon it is a primary júnior
homonym.
There is a limited usage of Tapes vir-
gineus in XX century as valid ñame for a
European species (Newton, 1907; Har-
greaves, 1910; Ford, 1923; Fisher &
Templeton, 1935; Marres, 1947), which
preeludes application of Art. 23.9.
Nordsieck (1969: 118) States in syn-
onymy of Tapes rhomboides: “ Tapes vir¬
gíneas L., pars?" but does not use it.
Dodge (1952) stated an unmarked
specimen of Venus virgínea with no
proven type status in the Linnean
Society, London, which would agree
with the specimen figured by Reeve
(1864, pl. 4 sp. 17a) as Tapes virgínea.
Reeve's figure is clearly the same as
Venus rhomboides Pennant, 1777 but,
again, this has no bearing on the
Linnean species. This shell is not cur-
rently registered as a type on the
Linnean Society website
<http:/ /linnean-online.orgx Con-
versely there is a lot in University
Museum, Uppsala (# 1347-1349) labelled
as this species (Wallin, 2001). These
types are accompanied with the so-
called 'Swartz labels", printed using the
12th edition of "Systema Naturae" as a
model (Wallin, 2001). We consider
specimens 1347 and 1349 to belong to
the species currently known as Venerupis
geographica (Gmelin, 1791) and specimen
1349 to belong to the species currently
known as Polititapes aureus (Gmelin,
1791). The latter resembles Polititapes
rhomboides but is unequivocally dístin-
guished by the much more narrow and
less robust hinge píate, and the colour
pattern with yellowish inside and
brown streaks outside is more typical of
aureus than of rhomboides.
From the preceding discussion we
conclude that, would the ñame Venus
virgínea be revived, it should stand as a
sénior synonym of Polititapes aureus
(Gmelin, 1791) or Venerupis geographica
(Gmelin, 1791) if based on the extant
types, and is not Venus rhomboides
Pennant, 1777. Since both species ( Venus
hiantina and V. rhomboides ) for which
Linnaeus' ñame Venus virgínea has actu-
ally been used have another accepted
ñame with established usage, and since
so much confusión surrounds the
Linnean ñame, it is our intention to
request suppression of the latter from
the ICZN. In the meanwhile, we
propose to continué the accustomed
usage of the specific ñame Polititapes
rhomboides (Pennant, 1777) as valid.
Polititapes rhomboides is a commercial
species, albeit not of major importance,
its marketing is regulated under this
ñame in Spain and Portugal, and it is
known under this ñame in legal texts
regarding size and cióse season.
77
Iberus , 32 (1), 2014
Another issue related to this ñame
change is the generic placements for the
European species of Tapetinae, particu-
larly the option of ranking Ruditapes Chi-
amenti, 1900 (type species: Venus decus-
sata Linnaeus, 1758, subsequent designa
tion by Dall, 1902: 363) as a subgenus of
Venerupis but Polititapes Chiamenti, 1900
(type species: Venus aurea Gmelin, 1791,
subsequent designation by Dall, 1902:
363) as a genus. Without this being backed
by a novel phylogenetic analysis of the
Tapetinae, we feel it is preferable to follow
a conservaííve usage, particularly for these
ñames which apply to commercial species
and are used by many non-taxonomists
and also in legal documents. We can agree
that Tapes (based on Venus literata Lin-
naeus, 1758) is no i appropriate for the
species considered here but the two most
obvious options seem to be either to place
all in Venerupis and rank Polititapes also as
subgenus, or rank Ruditapes as full genus
as well.
Cuspidaria striata var. bicarinata (Jeffreys, 1876) vs. Cuspidaria
bicarinata (Jeffreys, 1882)
Ñames involved
Neaera striata var. bicarinata Jeffreys, 1876, Annals and Magazine of Natural History, (4) 18: 496.
Neaera bicarinata Jeffreys, 1882, Proceedings ofthe Zoological Society ofLondon (1881): 939, pl. 71 fig. 1.
Huber (2010: 792=793) declared
Neaera bicarinata Jeffreys, 1882 a nomen
protectum and Neaera striata var. bicari¬
nata Jeffreys, 1876, a nomen oblitum. The
species was not cited by Allen & Mor¬
gan (1981), is merely listed in Poutiers
& Bernard (1995), and has hardly ever
been cited any more in the XX century
literature, so that we are certain that the
requirements of Art. 23.9.1 are not met
Anyway Neaera bicarinata Jeffreys, 1882
is not preoccupied. Jeffreys (1876) men-
tioned a fragment of a species of Neaera
from 'Valorous' sta. 13, described it and
then wrote "It probably belongs to a
ACKNOWLEDGEMENTS
We are grateful to Erica Mejlon, the
Evolution Museum, Uppsala, Sweden, for
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sala University, Museum of Evolution, Zo-
ology Section (UEJZM), Uppsala.
Wells F.E. & Bryce C.W. 1984. A guide to the
common molluscs ofsouth-western Australian es¬
tuarios. Western Australian Museum, 112 p.
Wildish D. J. & Peer D. 1983. Tidal current
speed and production of benthic macrofauna
in the lower Bay of Fundy. Canadian Journal
ofFisheries and Aquatic Sciences, 40 (SI): s309-
s321.
Wilson B. 1998. Order Mytiloida. pp. 250-253.
In Beesley P.L., Ross G.J.B. & Wells A. (Eds.):
Mollusca: The Southern Synthesis. Fauna of Aus¬
tralia, Vol 5. CSIRO Publishing, Melbourne.
W inckworth R. 1932. The British marine mol-
lusca. Journal of Conchology, 19: 211-252.
Zenetos A. 1996. The marine Bivalvia (Mol-
lusca) of Greece. In Fauna Greciae, vol. VII (ed.
Hellenic Zoological Society and NCMR), pp.
1-319. Athens: NCMR.
Zenetos A. 1997. Diversity of marine Bivalvia
in Greek waters: effects of geography and en-
vironment. Journal of the Marine Biological As-
sociation ofthe United Kingdom, 77: 463-472.
85
'
NORMAS DE PUBLICACION
• La revista Iberus publica artículos de fondo, notas y monografías que versen sobre cualquiera de los aspectos
relacionados con la Malacología. Se entiende por artículo un trabajo de investigación de más de 5 páginas de
texto, incluidas láminas, gráficos y tablas. Las notas son trabajos de menor extensión. Las monografías son tra¬
bajos sobre un tema único, de extensión superior a las 50 páginas de la revista y que serán publicadas, si
procede, como un suplemento de Iberus. Los autores interesados en publicar monografías deberán ponerse
previamente en contacto con el Editor de Publicaciones. Se entiende que el contenido de los manuscritos no ha
sido publicado, ni enviado simultáneamente a otra revista para su consideración.
• Los manuscritos, así como toda la correspondencia relacionada con los mismos, deberán ser remitidos a:
Serge Gofas, Editor de Publicaciones, Departamento de Biología Animal, Universidad de Málaga, Campus de
Teatinos, s/n, 29071, Málaga, España y/o al correo electrónico <sgofas@uma.es>.
• El texto del trabajo podrá estar redactado en español, inglés, italiano, francés o portugués.
• Los artículos, notas y monografías deberán presentarse sobre DIN A-4, por una sola cara a doble espacio
(incluyendo referencias, pies de figura y tablas), con al menos 3 centímetros de margen por cada lado. Los tra¬
bajos se entregarán por triplicado (original y dos copias) y se incluirá una versión en un CD, o bien remitida
por correo electrónico, utilizando procesadores de texto en sus versiones de corrientes de Windows (pero no
en el formato .docx de Word 2007, el habitual de Windows Vista) o Macintosh. En caso de autoría compartida,
uno de los autores deberá hacerse responsable de toda la correspondencia.
• Junto con el trabajo debe incluirse una lista de al menos 4 posibles revisores del mismo, sin perjuicio de los
que el propio Editor considere oportunos.
• Los manuscritos se presentarán de acuerdo al siguiente esquema:
Primera página. Deberá incluir un título conciso, pero sugerente del contenido del trabajo, así como una traduc¬
ción al inglés del mismo (si el artículo no está escrito en inglés). Cuando sea preciso, deberá incluir referencia a
familia o taxones superiores. A continuación figurarán, por este orden, el nombre y apellidos completos del
autor o autores, las direcciones completas de los mismos, y un resumen del trabajo y su traducción al inglés.
Dicho resumen deberá sintetizar, en conjunción con el título, tanto los resultados como las conclusiones del
artículo; se sugiere una extensión de 100 a 200 palabras.
Páginas siguientes. Incluirán el resto del artículo, que debe dividirse en secciones precedidas por breves encabe¬
zamientos. Siempre que sea posible, se recomienda seguir el siguiente esquema: Introducción, Material y
métodos. Resultados, Discusión, Conclusiones, Agradecimientos y Bibliografía. Si se emplean abreviaturas no
habituales en el texto, deberán indicarse tras el apartado de Material y Métodos.
• Las notas breves deberán presentarse de la misma forma, pero sin resumen.
• Deberán evitarse notas a pie de página y referencias cruzadas. Deberán respetarse estrictamente los Códigos
Internacionales de Nomenclatura Zoológica y Botánica (últimas ediciones). Cuando un taxón aparezca por
primera vez deberá citarse su autor y fecha de su descripción. En el caso de artículos sistemáticos, cuando se
den las sinonimias de los taxones, éstas deberán citarse COMPLETAS, incluyendo en forma abreviada la
publicación donde fueron descritas, y la localidad tipo si es conocida entre corchetes, según el siguiente
esquema (préstese especial cuidado a la puntuación):
Dendrodoris limbata (Cuvier, 1804)
Sinonimias
Doris limbata Cuvier, 1804, Ann. Mus. Hist. Nat. Paris, 4 (24): 468-469 [Localidad tipo: Marsella].
Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop.-Car., 10: 275.
Dichas referencias no deberán incluirse en la lista de Bibliografía si es la única vez que se nombran en el texto.
Si se incluyen una lista completa de referencias de un taxón inmediatamente tras éste, deberá seguirse el
mismo esquema (sin incluir en Bibliografía las referencias que no se mencionen en otro lugar del texto).
• Sólo los nombres en latín y los de taxones genéricos y específicos deberán llevar subrayado sencillo o prefe¬
rentemente ir en cursiva. En ningún caso deberá escribirse una palabra totalmente en letras mayúsculas, ni
siquiera el Título. Las unidades a utilizar deberán pertenecer al Sistema Métrico Decimal, junto con sus correc¬
tas abreviaturas. En artículos escritos en castellano, en los números decimales sepárese la parte entera de la
decimal por una coma inferior (,), NUNCA por un punto (.) o coma superior (').
• Las referencias bibliográficas irán en el texto con minúsculas o versalitas: Fretter y Graham (1962) o Fretter
y Graham (1962). Si son más de dos autores se deberán citar todos la primera vez que aparecen en el texto
[Smith, Jones y Brown (1970)] empleándose et al. las siguientes veces [Smith et al. (1970)]. Si un autor ha publi¬
cado más de un trabajo en un año se citarán con letras: (Davis, 1989a; Davis, 1989b). No deberá emplearse op.
cit. La lista de referencias deberá incluir todas las citas del texto y sólo éstas, ordenadas alfabéticamente. Se
citarán los nombres de todos los autores de cada referencia, sea cual sea su número. Los nombres de los
autores deberán escribirse, en letras minúsculas o Versalitas. No deberán incluirse referencias a autores
cuando éstos aparezcan en el texto exclusivamente como autoridades de un taxón. Los nombres de las publica-
ciones periódicas deberán aparecer COMPLETOS, no abreviados. Cuando se citen libros, dése el título, editor,
lugar de publicación, n° de edición si no es la primera y número total de páginas. Deberán evitarse referencias
a Tesis Doctorales u otros documentos inéditos de difícil consulta. Síganse los siguientes ejemplos (préstese
atención a la puntuación):
Fretter V. y Graham A. 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp.
Ponder W.F. 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. En Ponder, W.F.
(Ed.): Prosobranch Phylogeny. Malacological Review, suppl. 4: 129-166.
Ros J. 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Mis¬
celánea Zoológica, 3 (5): 21-51.
• Las gráficas e ilustraciones deberán ser originales y presentarse preferentemente en formato electrónico al
formato de caja de la revista o proporcional a éste. Este formato es de 57 mm (una columna) o 121 mm (dos) de
anchura y hasta 196 mm de altura, si bien se recomienda utilizar el formato a dos columnas. Es importante que
todas las figuras sean remitidas en su formato original (por ejemplo, las fotografías en .jpg de alta calidad o .tif,
las gráficas en hojas de cálculo Excel o documentos de CorelDraw), puesto que las ilustraciones insertadas en
el manuscrito WORD son inservibles en la fase de imprenta. Las imágenes digitales deben ser formateadas en
su tamaño de impresión con una resolución mínima de 300 ppp para imágenes en color o escala de grises y de
600 ppp para las de blanco y negro. Considérese la reducción que será necesaria a la hora de decidir el tamaño
de las escalas o letras en las figuras, que no deberán bajar de los 2 mm. En figuras compuestas, cada parte
deberá etiquetarse con letras mayúsculas, el resto de las letras deberán ser minúsculas. Las escalas de dibujos y
fotografías deberán ser gráficas, utilizando unidades del sistema métrico decimal; no deberán hacerse referen¬
cias a los aumentos de una determinada ilustración, ya que éstos cambian con la reducción. En su caso, se
recomienda la utilización de mapas con proyección UTM. Cada figura, gráfica o ilustración deberá presentarse
en hojas separadas y con numeración arábiga (1, 2, 3,...), sin separar "Figuras" y "Láminas". Los pies de
figura, en una hoja aparte, deberán acompañarse de su traducción al inglés (o español si el inglés es la lengua
del trabajo). Utilícese el esquema siguiente:
Figura 1. Neodoris carvi. A: animal desplazándose; B: detalle de un rinóforo; C: branquia.
Las abreviaturas empleadas en las ilustraciones deberán incluirse en los pies de figura.
Los autores interesados en incluir láminas en color deberán consultar con el editor y sufragar el sobrecoste con
una contribución de 30 €por página. Por lo demás, deberán ajustarse a los mismos requisitos indicados para
las figuras.
Si se pretende enviar gráficas o ilustraciones en impresión de papel es imprescindible presentar originales de
buena calidad. Las imágenes en semitonos deben estar bien contrastadas y ajustarse al tamaño definitivo de
impresión; al componer fotografías sobre una hoja, procúrese que los espacios entre ellas sean regulares y que
estén debidamente alineadas. Téngase en cuenta que incluir fotografías de distinto contraste en una misma
página conlleva una pobre reproducción final. Las gráficas de ordenador deberán imprimirse con impresora
láser sobre papel de buena calidad.
• Las Tablas se presentarán en hojas separadas, siempre con numeración romana (I, II, III,...). Las leyendas se
incluirán en una hoja aparte acompañándose de una traducción al inglés. Deberán evitarse las tablas particu¬
larmente complejas. Se recomienda reducir el número y extensión de ilustraciones, láminas o tablas al mínimo
necesario.
• Los artículos que no se ajusten a las normas de publicación serán devueltos al autor con las indicaciones de
los cambios necesarios.
• El Comité Editorial comunicará al autor responsable del trabajo la fecha de recepción del trabajo y la fecha
de envío a revisión. Cada original recibido será sometido a revisión por al menos dos investigadores. El
Comité Editorial, a la vista de los informes de los revisores decidirá sobre la aceptación o no de cada manus¬
crito. El autor recibirá en cada caso copia de los comentarios de los revisores sobre su artículo. En caso de
aceptación, el mismo Comité Editorial, si lo considera conveniente, podrá solicitar a los autores otras modifica¬
ciones que considere oportunas. Si el trabajo es aceptado, el autor deberá enviar una copia impresa del mismo
corregida, acompañada por una versión en un CD, utilizando procesadores de texto en sus versiones corrien¬
tes de Windows (pero no el formato .docx generado por Word 2007, el habitual de Windows Vista) o Macin¬
tosh. La fecha de aceptación figurará en el artículo publicado.
• Las pruebas de imprenta serán enviadas por correo electrónico al autor responsable, exclusivamente para la
corrección de erratas, y deberán ser devueltas en un plazo máximo de una semana. Se recomienda prestar
especial atención en la corrección de las pruebas.
• De cada trabajo se entregarán gratuitamente 50 separatas, además de una versión electrónica en formato pdf.
Aquellos autores que deseen un número mayor, deberán hacerlo constar al devolver las pruebas de imprenta,
y nunca posteriormente. El coste de las separatas adicionales será cargado al autor. Los autores podrán hacer
uso de sus separatas en formato electrónico en una web personal o institucional, siempre que se trate de un
sitio sin ánimo de lucro.
INSTRUCTIONS TO AUTHORS
• Iberus publishes research papers, notes and monographs devoted to the various aspects of Malacology.
Papers are manuscripts of more than 5 typed pages, including figures and tables. Notes are shorter papers.
Monographs should exceed 50 pages of the final periodical, and may be published as Supplements. Authors
wishing to publish monographs should contact the Editor. Manuscripts are considered on the understanding
that their contents have not been published or simultaneously submitted for publication elsewhere.
• Manuscripts and correspondence regarding editorial matters must be sent to: Serge Gofas, Editor de Publica¬
ciones, Departamento de Biología Animal, Universidad de Málaga, Campus de Teatinos, s/n, 29071, Málaga,
Spain and/or to the e-mail <sgofas@uma.es>.
• Manuscripts may be written in Spanish, English, Italian, French or Portuguese.
• Manuscripts must be typed double spaced (including the references, figure captions and tables) on one side
on A-4 (297x210 mm) with margins of at least 3 cm. An original and two copies must be submitted, together
with a CD or e-mail remittance containing the article written with a current Windows (but not .docx format
generated by Word 2007, mainly used with Windows Vista) or Macintosh word processor. When a paper has
joint authorship, one author must accept responsibility for all correspondence.
• The authors must inelude a list of at least 4 possible referees; the Editor can choose any others if appropriate.
• Papers should conform the following layout:
First page. This must inelude a concise but informative title, with mention of family of higher taxon when
appropriate, and its Spanish translation. It will be followed by all authors' ñames and surnames, their full
address(es), an abstract (and its Spanish translation) not exceeding 200 words which summarizes not only con¬
tents but results and conclusions.
Folloiving pages. These should content the rest of the paper, divided into sections under short headings. When-
ever possible the text should be arranged as follows: Introduction, Material and methods, Results, Discussion,
Conclusions, Acknowledgements and References. Unusual abbreviations used in the text must be grouped in
one alphabetic sequence after the Material and methods section.
• Notes should follow the same layout, without the abstract.
• Footnotes and cross-references must be avoided. The International Codes of Zoological and Botanical
Nomenclature must be strictly followed. The first mention in the text of any taxon must be followed by its
authority including the year. In systematic papers, when synonyms of a taxon are given, they must be cited IN
FULL, including the periodical, in an abbreviate form, where they were described, and the type localities in
square brackets when known. Follow this example (please note the punctuation):
Dendrodoris limbata (Cuvier, 1804)
Synonyms
Doris í imbata Cuvier, 1804, Ann. Mus. Hist. Nat. París , 4 (24): 468-469 [Type locality: Marseille].
Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop.-Car., 10: 275.
These references must not be included in the Bibliography list, except if referred to elsewhere in the text. If a
full list of references of the taxon is to be given immediately below it, the same layout should be followed (also
excluding from the Bibliography list those which are not cited elsewhere).
Only Latin words and ñames of genera and species should be underlined once or be given in italics. No word
must be written in UPPER CASE LETTERS. SI units are to be used, together with their appropriate symbols. In
Spanish manuscripts, decimal numbers must be separated with a comma (,), NEVER with a point (.) or upper
comma (').
• References in the text should be written in small letters or Small capitals: Fretter and Graham (1962) or
Fretter and Graham (1962). The first mention in the text of a paper with more than two authors must
inelude all of them [Smith, Jones and Brown (1970)], thereafter use et al. [Smith et al. (1970)]. If an author has
published more than one paper per year, refer to them with letters: (Davis, 1989a; Davis, 1989b). Avoid op. cit.
The references in the reference list should be in alphabetical order and inelude all the publications cited in the
text but only these. ALL the authors of a paper must be included. These should be written in small letters or
Small capitals. The references need not be cited when the author and date are given only as authority for a tax-
onomic ñame. Titles of periodicals must be given IN FULL, not abbreviated. For books, give the title, ñame of
publisher, place of publication, indication of edition if not the first and total number of pages. Keep references
to doctoral theses or any other unpublished documents to an absolute mínimum. See the following examples
(please note the punctuation):
Fretter V. and Graham A. 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp.
Ponder W.F. 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. In Ponder W.F.
(Ed.): Prosobranch Phylogeny. Malacological Review, suppl. 4: 129-166.
Ros J. 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Mis¬
celánea Zoológica, 3 (5): 21-51.
• Figures must be original and provided preferably in electronic format and adjusted to page format and
column size. These should be one column (57 mm) or two columns (121 mm) wide and up 196 mm high, or be
proportional to these sizes. Two columns format is recommended. It is essential that all figures be supplied in
their original format (e.g. photographs as high-grade .jpg or as .tif files, graphs as Excel spreadsheets or Corel-
Draw files), as the files inserted into WORD documents cannot be used for printing. Digital images must be
given their final printing size with a resolution at least 300 dpi for colour and halftones, and at least 600 dpi for
black/ white.
Take into account possible reduction in lettering drawings; final lettering must be at least 2 mm high. In com-
posite drawings, each figure should be given a capital letter; additional lettering should be in lower-case
letters. A scale line, labelled with SI units, must be used to indicate size; magnification ratio must be avoided
as it may be changed during printing. UTM maps are to be used if necessary. Figures must be submitted on
sepárate sheets, and numbered with consecutive Arabic numbers (1, 2, 3,...), without separating 'Plates' and
'Figures'. Legends for Figures must be typed in numerical order on a sepárate sheet, and a Spanish translation
must be included. Follow this example (please note the punctuation):
Figure 1. Neodoris carvi. A: animal crawling; B: rhinophore; C: gills.
If abbreviations are to be used in illustrations, they should be included in the figure captions.
Authors wishing to publish illustrations in colour should consult with the editor and will be charged a contri-
bution of 30 €per page. They should otherwise follow the same standards as black and white prints.
If the authors want to send Figures in printed format, it is essential to supply good quality origináis. Half-tone
images must be of good contrast, and should be submitted in the final printing size. When mounting pho¬
tographs in a block, ensure spacers are of uniform width. Remember that grouping photographs of varied con¬
trast results in poor reproduction. Computer graphics must be printed on high quality white paper with a
láser printer.
• Tables must be numbered with Román numbers (I, II, III,...) and each typed on a sepárate sheet. Headings
should be typed on a sepárate sheet, together with their English translation. Complex tables should be
avoided. As a general rule, keep the number and extensión of illustrations and tables as reduced as possible.
• Manuscripts that do not conform to these instructions will be returned for correction before reviewing.
• Authors submitting manuscripts will receive an acknowledgement of receipt, including receipt date, and the
date the manuscript was sent for reviewing. Each manuscript will be critically evaluated by at least two refer¬
ees. Based of these evaluations, the Editorial Board will decide on acceptance or rejection. Anyway, authors
will receive a copy of the referees' comments. If a manuscript is accepted, the Editorial Board may indicate
additional changes if desirable. Acceptable manuscripts will be returned to the author for consideration of
comments and criticism; a finalized manuscript must then be returned to the Editor, together with a CD con-
taining the article written with current Windows (but not a .docx format genera ted by Word 2007, mainly used
under Windows Vista) or Macintosh word processor. Dates of reception and acceptance of the manuscript will
appear in all published articles.
• Proofs will be e-mailed to the author for correcting errors and must be returned corrected within one week.
At this stage no stylistic changes will be accepted. Pay special attention to references and their dates in the text
and the Bibliography section, and also to numbers of Figures and Tables appearing in the text.
• Fifty reprints per article and a pdf file will be supplied free of charge. Additional reprints must be ordered
when the page proofs are returned, and will be charged at cost. NO LATER orders will be accepted. Authors
may post a pdf file of their papers on a personal or institutional webpage, with the condition that it should be
a non-profit website.
La Sociedad Española de Malacología
Junta Directiva desde el 11 de julio de 201 1
Presidente
Vicepresidente
Secretario
Tesorero
Editor de Publicaciones
Bibliotecario
Vocales
Jesús S. Troncoso
Gonzalo Giribet
Ramón Álvarez Halcón
Luis Murillo Guillén
Serge Gofas
Rafael Araujo Armero
José Templado González
Carmen Salas Casanova
Alberto Martínez Ortí
José Ramón Arrébola Burgos
Benjamín Gómez Moliner
La Sociedad Española de Malacología se fundó el 21 de agosto de 1980. La sociedad se registró como una aso¬
ciación sin ánimo de lucro en Madrid (Registro N° 4053) con unos estatutos que fueron aprobados el 12 de
diciembre de 1980. Esta sociedad se constituye con el fin de fomentar y difundir los estudios malacológicos
mediante reuniones y publicaciones. A esta sociedad puede pertenecer cualquier persona o institución interesada
en el estudio de los moluscos.
Sede social: Museo Nacional de Ciencias Naturales, d José Gutiérrez Abascal 2, 28006 Madrid, España.
Cuotas para 2014:
DESDE ABRIL
Inscripción: 6 euros, además de la cuota correspondiente.
A los socios residentes en España se les aconseja domiciliar su cuota. Todos los abonos deberán enviarse al
Tesorero (dirección reseñada anteriormente) el 1 de enero de cada año. Los abonos se harán sin recargos para la
sociedad y en favor de la Sociedad Española de Malacología y no de ninguna persona de la junta directiva. Aque¬
llos socios que no abonen su cuota anual dejarán de recibir las publicaciones de la Sociedad. Los bonos de ins¬
cripción se enviarán junto con el abono de una cuota anual al Tesorero.
A los residentes en el extranjero se les ruega que abonen su cuota mediante giro postal en euros (internatio-
nal postal money orders in euros sent to the Treasurer). Members living in foreing countries can deduce 6 euros
if paid before 15 April.
Cada socio tiene derecho a recibir anualmente los números de Iberus, Reseñas Malacológicas y Noticiarios que
se publiquen.
SMITHSONÍAN INSTITUTION LIBRARES
3 9088 01751 3839
Indice
Iberus
32 (1) 2014
RoláN E. AND Gori S. New information on the marginellids of Sao Tomé and Principe, with
. new records and the description of four new species
Nueva información sobre los marginelidos de Sao Tomé y Principe, con nuevas citas y descrip¬
ción de cuatro especies nuevas . 1-25
Holyoak D.T. AND Holyoak G.A. An undescribed species of Cryptosaccus (Gastropoda: Hygro-
kmiidae) from the south-west of the province of León, NW Spain
Una nueva especie de Cryptosaccus ( Gastropoda: Hygromiidae) del sur de Province León, NO
España . 27-34
BARROS A. Estado de conservación de la náyade Margaritifera margaritifera (Linnaeus 1758) en el
curso bajo del río Mandeo (A Coruña)
Status ofthe freshwater pearl mussel Margaritifera margaritifera (Linnaeus 1758) in the lower
course of the river Mandeo (A Coruña) . 35-43
GOFAS S., Kantor Y. AND LUQUE A.A. A new Aforia (Gastropoda: Conoidea: Cochlespiridae)
from Galicia Bank (NW Iberian Peninsula)
Una nueva Aforia (Gastropoda: Conoidea: Cochlespiridae) del banco de Galicia (NO Penín¬
sula Ibérica) . 45-5 1
Vanhaelen A., Massin C., Martin J. and Laffargue P. Kaloplocamus ramosas (Cantraine,
1835) (Gastropoda: Polyceridae): new records in the Bay of Biscay, with notes on distribu-
tion and food
Kaloplocamus ramosus (Cantraine, 1835) (Gastropoda: Polyceridae): nuevos sitios en el Golfo
de Vizcaya, con datos sobre su distribución y su dieta . 53-64
COSEL R.V., GOFAS S. and Poutiers J. Nomenclatural notes on some European marine bivalve
species
Apuntes nomenclaturales sobre algunas especies de bivalvos de Europa . 65-85
ISSN 0212-3010