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Revista de la

SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

Comité DE REDACCIÓN (BOARD OF EDITORS)

EDITOR DE PUBLICACIONES (EDITOR-IN-CHIEF)

Gonzalo Rodríguez Casero Apdo. 156, Mieres del Comino, Asturias, España

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Eugenia M* Martínez Cueto-Felgueroso Apdo. 156, Mieres del Camino, Asturias, España

EDITORES ADJUNTOS (ASSOCIATE EDITORS)

Benjamín Gómez Moliner Universidad del País Vasco, Vitoria, España

Ángel Antonio Luque del Villar Universidad Autónoma de Madrid, Madrid, España

Emilio Rolán Mosquera Universidad de Vigo, Vigo, España

José Templado González Museo Nacional de Ciencias Naturales, CSIC, Madrid, España

Jesús S. Troncoso Universidad de Vigo, Vigo, España

Comité EDITORIAL (BOARD OF REVIEWERS)

Kepa Altonaga Sustacha Universidad del Poís Vasco, Bilbao, España

Eduardo Angulo Pinedo Universidad del País Vasco, Bilbao, España

Rofael Araujo Armero Museo Nacional de Ciencias Naturales, Madrid, España

Thierry Bockeljau Institut Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica

Rúdiger Bieler The Field Museum, Chicago, Estados Unidos

Sigurd v. Boletzky Loboratoire Arago, Banyuls-sur-Mer, Francia

Jose Castillejo Murillo Universidad de Santiago de Compostela, Santiago de Compostela, España

Karl Edlinger Naturhistorisches Museum Wien, Viena, Austria

Antonio M. de Frias Martins Universidade dos Acores, Acores, Portugal

José Carlos García Gómez Universidad de Sevilla, Sevilla, España

Gonzalo Giribet de Sebastián Harvard University, EE.UU.

Edmund Gittenberger Notional Natuurhistorisch Museum, Leiden, Holanda

Serge Gofas Universidad de Málaga, España

Angel Guerra Sierra Instituto de Investigaciones Marinas, CSIC, Vigo, España

Gerhard Haszprunar Zoologische Staatssammlung Múnchen, Múnchen, Alemania

Yuri |. Kantor AN. Severtzov Institute of Ecology and Evolution, Moscú, Rusia

María Yolanda Manga González Estación Agrícola Experimental, CSIC, León, España

Jordi Martinell Callico Universidad de Barcelona, Barcelona, España

Ron K. 0'Dor Dalhousie University, Halifax, Canada

Tokoshi Okutani Nihon University, Fujisawo City, Japón

Marco Oliverio Universitá di Roma “La Sapienza”, Roma, Italia

Pablo E. Penchaszadeh Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina

Winston F. Ponder Australian Museum, Sydney, Australia

Carlos Enrique Prieto Sierra Universidad del Poís Vasco, Bilbao, España

Me de los Ángeles Ramos Sánchez Museo Nacional de Ciencias Naturales, CSIC, Madrid, España

Francisco Javier Rocha Valdés Instituto de Investigaciones Marinas, CSIC, Vigo, España

Paul 6. Rodhouse British Antarctic Survey, Cambridge, Reino Unido

Joundoménec Ros ¡ Aragones Universidad de Barcelona, Barcelona, España

María Carmen Solas Casanovas Universidad de Málaga, Málaga, España

Gerhard Steiner Institut fir Zoologie der Universitát Wien, Viena, Austria

Victoriano Urgorri Carrasco Universidad de Santiago de Compostela, Santiago de Compostela, España

Anders Worén Swedish Museum of Natural History, Estocolmo, Suecia

PORTADA DE /berus

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nombre a la revista. Dibujo realizado por José Luis González Rebollar “Toza”.

Iberus LIBRARY

OCT 19 2009

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REVISTA DE LA

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Vol. 21 (2) Oviedo, diciembre 2003

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Revista de la

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Iberus publica trabajos que traten sobre cualquier aspecto relacionado con la Malacología. Se

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ISSN 0212-3010

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S.E. mM >

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deal

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O Sociedad Española de Malacología ——____——T— Iberus, 21 (2): 1-17, 2003

Characterization and multivariate analysis of Patella inter-

media, Patella ulyssiponensis and Patella vulgata from Póvoa

de Varzim (Northwest Portugal)

Caracterización y análisis multivariante de Patella intermedia, Patella

ulyssiponensis and Patella vulgata de Póvoa de Varzim (noroeste de

Portugal)

Joáo Paulo CABRAL*

Recibido el 9-1X-2002. Aceptado el 13-1-2003

ABSTRACT

Samples of Patella intermedia, Patella ulyssiponensis and Patella vulgata were collected from

the lowest to the highest level of A Ver-o-Mar and Agucadoura beaches [Póvoa de Varzim,

Portugal). Identification of the specimens was based on the morphology of the radula pluri-

cuspid teeth and of the shell. Regressions of shell width or shell apexpos vs. shell length

yielded slopes not statistically different, but slopes of log shell height vs. shell length regres-

sion lines were significantly different, indicating that these limpets have different shell grow-

ing patterns. However, canonical discriminant analyses using only variables describing shell

form yielded poor discrimination between species. Analysis using only variables describing

radula relative size improved discrimination, was very satisfactory for P. ulyssiponensis spec-

imens, but resulted in a low identification of P. intermedia. The best results were achieved us-

ing both shell form and radula relative size variables, and the most discriminating variable

was radula length/shell height ratio, instead of radula length/shell length as reported in the

literature. However, whereas correct identification of P. ulyssiponensis specimens was very

good [higher than 95 %), for P. intermedia and P. vulgata, this value ranged from 70 —- 80

%. In light of data reported in the literature, morphometric characteristics such as those uti-

lized in this work are probably useful for characterization of P. ulyssiponensis as a species,

but are of limited value for P. intermedia and P. vulgata. Qualitative characters are thus still

indispensable for the discrimination between these two Patella species. Very good qualita-

tive discriminators were, for P. intermedia, the tall and broad cusp 2 and the dark marginal

rays in the shell interior surface, and for P. vulgata, the pointed protuberance on cusp 3, and

the silvery head scar.

RESUMEN

Muestras de Patella intermedia, Patella ulyssiponensis y Patella vulgata fueron colectadas en

una franja abarcando desde el nivel mas bajo al mas alto de las playas de A Ver-o-Mar y

Agucadoura (Póvoa de Varzim, Portugal). La identificación de los especimenes se basó en

la morfología de la concha y de los dientes pluricúspides de la rádula. Las pendientes de

las regresiones entre la anchura o el apexpos (distancia entre la proyección del ápice y la

parte posterior de la concha) de la concha y la longitud de la concha no difirieron estadís-

* University of Oporto, Center of Marine and Environmental Research (CIIMAR) and Faculty of Sciences, Rua

do Campo Alegre, 1191, 4150-181 Porto, Portugal. jspcabralGhotmail.com

Iberus, 21 (2), 2003

ticamente, mientras que las pendientes de las regresiones entre el logaritmo de la altura y

la longitud de la concha fueron significativamente diferentes, indicando que estas lapas pre-

sentan patrones de crecimiento distintos. Sin embargo, el análisis canónico discriminatorio

utilizando solo las variables que describen la forma de la concha tuvo un bajo poder de dis-

criminación entre especies. Al utilizar solo las variables que describen el tamaño relativo de

la rádula se mejoró el poder discriminatorio del análisis, y aunque permitió una identifica-

ción satisfactoria de P. ulyssiponensis, pero resultó tener bajo poder para la identificación

de P. intermedia. Los mejores resultados se obtuvieron utilizando variables relacionadas tanto

con la forma de la concha como con el tamaño relativo de la rádula, y la variable con mayor

poder discriminatorio fue la razón longitud de la rádula/altura de la concha, en lugar de

longitud de la rádula/longitud de la concha, como se ha reportado en la bibliografía. Mien-

tras que la correcta identificación de especimenes de P. ulyssiponensis fue muy alta (mayor

que 95 %), para P. intermedia y P. vulgata, este valor varió entre 70 - 80 %. A la luz de los

datos publicados en la bibliografía, características morfométricas tales como las utilizadas

en el presente trabajo, pueden ser de gran utilidad para la caracterización específica de P.

ulyssiponensis, pero tienen un valor limitado para la caracterización de P. intermedia y P.

vulgata. Las características cualitativas, por tanto, siguen siendo indispensables para la dis-

criminación ente estas dos especies de Patella. Para P. intermedia, la segunda cúspide alta

y ancha, así como los radios marginales oscuros en la superficie interna de la concha, cons-

tituyeron muy buenos discriminadores cualitativos, mientras que para P. vulgata, lo fueron

la protuberancia puntiaguda en la tercera cúspide, junto con la cicatriz de la cabeza de

color plateada.

KEY WORDS: Patella, radula, canonical discriminant analysis, morphometry.

PALABRAS CLAVE: Patella, rádula, anñalisis canónico discriminante, morfometría.

INTRODUCTION

The characterization and delimita-

tion of Patella intermedia Murray in

Knapp 1857, Patella ulyssiponensis

Gmelin 1791, and Patella vulgata Linné

1758, has been a source of debate and

controversy since they were proposed as

species (FISHER-PIETTE, 1935; RAMPAL,

1965; IBAÑEZ, 1982). This was due to the

high shell variability commonly exhib-

ited by these species. In regions of low

variability, P. intermedia, P. ulyssiponensis

and P. vulgata shells are usually distinct

from each other. In regions of high vari-

ability, however, some shells display

intermediate characteristics, and reliable

identification of these species based on

shell morphology alone is difficult

(FISHER-PIETTE, 1934, 1948, 1966; EVANS,

1947, 1958; FISHER-PIETTE AND GAIL-

LARD, 1959). Shell variability of P.

vulgata and P. intermedia is highest in the

Basque coast of France and Spain, and

in the south limit of their distribution,

the Algarve (Portugal) for P. vulgata, and

northern Africa for P. intermedia. P.

ulyssiponensis is the species with the

least variable shell (FISCHER-PIETTE AND

GAILLARD, 1959; FISHER-PIETTE, 1966).

The difficulties in the identification of

P. intermedia, P. ulyssiponensis and P.

vulgata specimens based on shell mor-

phology alone led to the search for new

characters. Dautzenberg, in the end of

the 18th century, pointed out the impor-

tance of radula length and pigmentation

of mantle tentacle to discriminate several

Patella species, being therefore the first

malacologist to propose alternative char-

acteristics to shell morphology (FISCHER-

PIETTE AND GAILLARD 1959; RAMPAL,

1965). The work of FISHER-PIETTE (1934,

1935) showed, for the first time, that the

morphology of the radula pluricuspid

teeth could be used to characterize

several Patella species, since they usually

display low intraspecific variability, and

CABRAL: Characterization and multivariate analysis of Patella spp.in Northwest Portugal

A VER-O-MAR

4127 N

417 26

41? 25'

41 24'

41 23'

B” 44” 843" W

Figure 1. Location of sampling sites (A Ver-o-Mar and Agucadoura beaches) near Póvoa de Varzim

Inset: Localization of the town of Póvoa de Varzim and Oporto district in Portugal.

Figura 1. Ubicación de la zona de muestreo (playas de A Ver-0-Mar y Agugadoura) en las cercanías de

Póvoa de Varzim. Recuadro: Ubicación del pueblo de Póvoa de Varzim y el distrito de Oporto en Portugal.

marked interspecific differences. Fisher-

Piette initial observations were con-

firmed by EsLick (1940) and EvANs

(1947). In a latter paper, FISHER-PIETTE

AND GAILLARD (1959) showed that radula

unicuspid teeth could also be used to

Characterize several Patella species.

Other characteristics have been pro-

posed for specific discrimination in the

genus Patella: the foot colour, the breed-

ing seasons, and the ratio of the radula

length / cubic root of the shell volume

(FISHER-PIETTE, 1935, 1941, 1948; EVANS,

1947, 1953, 1958; FISHER-PIETTE AND

GAILLARD, 1959; RAMPAL, 1965; POWELL,

1973).

In this study, samples of P. intermedia,

P. ulyssiponensis and P. vulgata from

Póvoa de Varzim (northwest Portugal)

were examined and compared with

respect to several morphological and

morphometric characters describing the

radula, shell, and soft parts. In particular

the following questions were raised:

- How variable are these species in

Póvoa de Varzim?

- Is shell form similar in these

limpets?

- Are radula and shell characteristics

equally important for the separation of

these species?

- Can these species be discriminated

by morphometric characters alone,

rather than by morphological character-

istics?

MATERIAL AND METHODS

Collections were made at A Ver-o-

Mar and Agucadoura beaches, two very

similar and exposed shores situated

Iberus, 21 (2), 2003

A Ae

[E Height ao)

Length

Figure 2. Shell measurements used in canonical discriminant analysis.

Figura 2. Mediciones de la concha utilizadas en el análisis canónico discriminatorio.

near the town of Póvoa de Varzim (Fig.

1). The upper shore is composed of dis-

persed and heavily eroded granite boul-

ders. Barnacles (Chthamalus) cover most

of the surfaces and P. vulgata is the dom-

inant limpet. Some mussels and gas-

tropods live in the crevices. The middle

shore is composed of horizontal granite

platforms, very eroded, with small

sandy beaches in between. P. intermedia

and P. vulgata are common in middle

shore. P. ulyssiponensis is restricted to

lower shore levels, where P. intermedia

and P. vulgata also occur. Most of the P.

ulyssiponensis shells are covered with

abundant Gelidium pulchelum (Turner)

Kútzing. Common macroalgae in the

middle and lower shore include Bifur-

caria bifurcata R. Ross, Chondrus crispus

Stackhouse, Codium tomentosum (Hud-

son) Stackhouse, coralline rhodophytes,

Cystoseira sp., Enteromorpha spp., Gi-

gartina pistillata (Gmelin) Stackhouse,

Gracilaria verrucosa (Hudson) Papenfuss,

Lythophyllum spp., Mastocarpus stellatus

(Stackhouse in Withering) Guiry and

Sargassum muticum (Yendo) Feusholt.

Sampling took place in January and No-

vember 2000, at low tide.

Specimens were collected from the

lowest to the highest level of the shores.

Squared areas with ca. 30 x 30 cm were

marked at random in each level, and all

the animals in each area removed from

the rock. The total number of collected

animals was 608.

Specimens were analysed for foot,

radula and shell characteristics. In the

laboratory, the animals were observed for

the foot colour, and then immersed for a

few minutes in boiling water to separate

the shell from the soft part. The radula

was removed from the visceral mass by

dissection, immersed in household

bleach to remove mucilaginous sub-

stances, washed in distilled water, and

measured to the nearest Imm using a

ruler. After air-drying, pluricuspid teeth

were first observed using a binocular mi-

croscope, with 80 x final magnification.

Structural details of the teeth were ob-

served by scanning electron microscopy,

in selected specimens. The samples were

gold coated (in a JEOL JFC1100 model;

film thickness less then 20 nm), and ob-

served in a JEOL JSM-35C model scan-

ning electron microscope, working at 15

keV, with 39 mm working distance.

The external and internal shell sur-

faces were examined and their charac-

teristics were recorded. Shell length,

width, height, apexante and apexpos

(Fig. 2 and Table I for the definition of

these measures) were then determined

to the nearest 0.01mm using a digital

calliper (Mitutoyo, model CD-15DC).

CABRAL: Characterization and multivariate analysis of Patella spp.in Northwest Portugal

Table I. Characters used in the canonical discriminant analysis.

Tabla [. Caracteres usados en el análisis canónico discriminante.

Acronym Description

SL Shell length: greatest distance between anterior and posterior ends

Sw Shell width: greatest distance perpendicular to the anterior-posterior axis

SH Shell height: greatest vertical distance from apex of the shell to the plane of aperture

SM Shell apexante: greatest distance between apex and anterior end

SAP Shell apexpos: greatest distance between apex and posterior end

sv Shell volume = (1/3) x (5W/2) x (SL/2) x SH

RL Rodula length

Morphometric analysis of the shell

and radula was carried out using the

variables commonly utilized in studies

of patellogastropoda (IBAÑEZ, 1982;

IBAÑEZ AND FELIU, 1983; FELIU AND

IBAÑEZ, 1984; HERNAÁNDEZ-DORTA, 1992;

SIMISON AND LINDBERG, 1999). The mean

and coefficient of variation of the mean

was calculated for each variable and

species. Linear regression analysis using

the least squares method was applied to

pairs of sheil or radula variables. These

calculations were carried out using

Microsoft Excel 2000 program. Com-

parison of the slopes was carried out by

analysis of covariance using the statisti-

cal procedure described by ZAR (1984).

Canonical discriminant analysis on

shell and radular morphometric vari-

ables was used to reveal differences

among the populations, and to identify

the variables that were responsible for

the majority of separation between

species. Discriminant analyses were car-

ried out in several steps. Firstly, using a

posteriori probabilities and no selection

of variables, four discriminant analyses

were carried out, two using only shell

Characters, one with only radula charac-

ters, and one using simultaneously shell

and radula characters. Secondly, using a

posteriori probabilities, and simultane-

ously shell and radula characters, dis-

criminant analysis was carried out with

stepwise selection of variables. Signifi-

cance level “to enter” and “to stay” the

variables was set at 0.15. Thirdly, cross-

validation analysis (using a priori proba-

bilities) was undertaken to assess the re-

liability of previous findings (using a

posteriori probabilities). For each species,

specimens were re-ordered randomly

using Excel 2000 program RAND func-

tion. Each group (species) was then di-

vided in two subgroups. For P. interme-

dia, these contained 152 specimens each,

for P. ulyssiponensis, 33 and 32, each, and

for P. vulgata, 119 and 120. The first

group (the analysis sample containing

the training or calibration data) was

used to derive the discriminant func-

tions, coefficients, and loadings. The

second group (the houldout sample con-

taining the test data) was used to test

the discriminant functions, by classify-

ing each specimen into one of the Patella

species. This process was repeated ten

times. At each run, a percent correct

identification to species was evaluated.

The means of these values were com-

pared with those obtained by using a

posteriori probabilities. Discriminant

analyses were performed using the

NCSS (NCSS Statistical Software, Utah,

USA) and XLSTAT 5.1 (Addinsoft, Paris,

France) software packages.

RESULTS

I. Delimitation of species

Based on radular and shell morphol-

ogy, the collected specimens were

divided into three groups. Within each

group, radula pluricuspid teeth dis-

played low variability, and shells exhib-

ited limited morphological variation.

Iberus, 21 (2), 2003

Figure 3. Morphology of pluricuspid teeth of the radula. Numbering of the cusps begins from left to

the right (corresponding to the outer and inner sides of the radula). Top: Patella intermedia. Cusp 2 is

taller and broader than cusps 1 and 3. Middle: Patella ulyssiponensis. Cusp 2 is directed to cusp 3.

Cusp 3 is wither than cusp 2. Cusp 3 has a protuberance on its outer side. Bottom: Patella vulgata.

Cusps 2 and 3 subequal. Cusp 3 has a pointed projection on its outer side border. Scale bars 100 um.

Figura 3. Morfología de los dientes pluricúspides de la rádula. La numeración de las cúspides comienza desde

la izquierda hacia la derecha (correspondiendo a los lados externos e internos de la rádula). Superior:

Patella intermedia. La segunda cúspide es más alta y ancha que la primera y tercera. Medio: Patella ulys-

siponensis. La segunda cúspide está dirigida hacia la tercera. Esta última es más ancha que la segunda. La

tercera cúspide tiene una protuberancia sobre el lado externo. Inferior: Patella vulgata. Las segunda y ter-

cera cúspides son subiguales. La tercera tiene una proyección puntiaguda en el borde externo. Escalas 100 ym.

The three groups displayed the fol- (left) and 3 (right). Margin of the shell

lowing features: rimose, with pointed extensions con-

Group I. Three unequal pluricuspid nected to the rays (Figure 4). External

teeth (Figure 3, top). Cusp 2 (centre) surface with few and prominent ribs.

much taller and broader than cusps 1 Interior with alternating dark and light

CABRAL: Characterization and multivariate analysis of Patella spp.in Northwest Portugal

Figure 4. Patella intermedia shells from A Ver-o-Mar and Agucadoura beaches.

Figura 4. Conchas de Patella intermedia provenientes de las playas de A Ver-o-Mar y Agucadoura.

rays, in the lower part. Head scar yellow-

orange, but creamy in a few specimens.

Group II. Three unequal pluricuspid

teeth (Figure 3, middle). Cusp 1 (left)

very small. Cusp 2 (centre) bent to cusp

3 (right). Base of cusp 3 wider than cusp

2. Cusp 3 with a protuberance on its

outer side — a vestigial fourth cusp.

Margin of the shell finely crenulate

(Figure 5). External surface crowded by

numerous, closely spaced, well-marked

ribs, of unequal size. Interior homoge-

neous, porcelanous white. Head scar

white, creamy or pale orange.

Group HH. Three unequal pluricuspid

teeth (Figure 3, bottom). Cusp 1 (left)

small. Cusps 2 (centre) and 3 (right)

subequal. Cusp 3 with a pointed protu-

Iberus, 21 (2), 2003

berance on its outer side border. Margin

of the shell entire or slightly indented

(Figure 6). External surface smooth at

the apex, with flat and spaced ribs

below. Interior with transparent nacre,

often with a green or blue iridescence.

Silvery head scar.

Based on shell and radula morpho-

logical characters reported in the litera-

ture for Patella spp. (FISHER-PIETTE, 1934,

1935; FISHER-PIETTE AND GAILLARD, 1959;

EVANS, 1947, 1953; ROLÁN, 1993; ROLÁN

AND OTERO-SCHMITT, 1996), groups 1, HI

and [Il were unequivocally identified as

belonging to P. intermedia, P. ulyssiponen-

sis and P. vulgata, respectively.

Radula pluricuspid teeth of P. inter-

media from Póvoa de Varzim is similar

to P. intermedia type B of EVANS (1953),

and to those reported by FISHER-PIETTE

(1934, figure 2), FISHER-PIETTE (1935,

figure 9, figure 17-2) and FISHER-PIETTE

AND GAILLARD (1959, figure 11 A and B).

Radula pluricuspid teeth of P.

ulyssiponensis from Póvoa de Varzim is

similar to P. ulyssiponensis type A of

EVANS (1953), and to those reported by

FISHER-PIETTE (1934, figure 2), FISHER-

PIETTE (1935 — figure 10, figure 17-9) and

FISHER-PIETTE AND GAILLARD (1959 —

figure 14 B).

Radula pluricuspid teeth of P.

vulgata from Póvoa de Varzim is similar

to P. vulgata types A and B of Evans

(1953), and to those reported by FISHER-

PIETTE (1934, figure 2), FISHER-PIETTE

(1935, figure 8, figure 17-1) and FISHER-

PIETTE AND GAILLARD (1959, figure 9A).

P. intermedia, P. ulyssiponensis and P.

vulgata shells from Póvoa de Varzim are

morphologically similar to those found

in Galiza (northwest Spain) (ROLÁN,

1993; ROLÁN AND OTERO-SCHMITIT, 1996)

and at Cardigan Bay (Wales, Great

Britain; EVANS, 1947).

The foot showed no constant

colouration within each group, and

therefore was not used in the identifica-

tion. P. intermedia foot was dark, from

grey or yellow to black. P. ulyssiponensis

foot was light, yellow or, most often,

orange. The colour of P. vulgata foot was

very variable, from light yellow to

black.

II. Morphometric delineation of

species

For all three Patella species, SW and

SA was a linear function of SL (Table ID.

However, SH was a curvilinear function

of SL, indicating that as these limpets

increase in size, the relative height of the

shell increases by a power function.

Similar relationship between SH and SL

has been reported for several Notoacmea

(LINDBERG, 1982) and Patella species

(DAVIES, 1969; MUÑOZ AND ACUÑA,

1994). Correlation between SH and SL

increased after log transformation of

SH, indicating that an exponential fit

was a better approach than a linear

function.

Regressions of SW or SAA vs. SL for

P. intermedia, P. ulyssiponensis and P.

vulgata specimens yielded slopes not

statistically different (Table ID.

However, slopes of log SH vs. SL regres-

sion lines were significantly different

(Table IM), indicating that the relative

increase in height as the shell grows was

not uniform in these Patella species.

Slopes increased in the order P. ulyssipo-

nensis, P. vulgata, and P. intermedia.

For all three Patella species, RL was a

linear function of SL and of “V(SV)

(Table ID). Similar relationship between

RE and SL has been reported for several

Patella species (BRIAN AND OWEN, 1952;

DAavIÉS, 1969; SELLA, 1976). Slopes of the

regression lines were significantly dif-

ferent, and increased in the order P.

ulyssiponensis, P. intermedia, and P.

vulgata (Table ID.

In P. intermedia, P. ulyssiponensis and

P. vulgata, radula relative size displayed

higher variability than shell form.

Whilst the coefficient of determination

for the regressions describing shell form

(SW vs. SL, log SH vs. SL, and SAP os.

SL) ranged between 0.597 and 0.952,

those describing radula relative size (RL

vs. SL, and RL vs. “W(SV)) varied

between 0.472 and 0.767 (Table ID.

Whilst the coefficient of variation of the

means describing shell form (SW/SL,

SH/SL, SAA/SAP, and SAP/SL) varied

between 5.4 and 22 %, those describing

radula relative size (RL/SL, RL/SH,

CABRAL: Characterization and multivariate analysis of Patella spp.1n Northwest Portugal

y pp 8

E E RS

Figures 5, 6. Patella shells from A Ver-o-Mar and Agucadoura beaches. 5: 2 ulyssiponensis; 6: P

vulgata.

Figuras 5, 6. Conchas de Patella provenientes de las playas de A Ver-o-Mar y Agucadoura. 5: P. ulyssipo-

nensis; 6: P. vulgata.

Iberus, 21 (2), 2003

Table II. Slopes of regression lines for morphometric characterization of the species, and their sta-

tistical comparison.

Tabla 11. Pendientes de las rectas de regresión de la caracterización morfomética de las especies, y su com-

paración estadística.

Comparison of slopes

Regression P. intermedia P. ulyssiponensis P. vulgata (DF=2, 404)

Slope E Slope p Slope y? F

Shell

SW vs. SL 0.859 0.952 0.791 0.918 0.821 0.929 JS

Logro SH vs. SL 0.0241 0.744 0.0149 0.597 0.0182 0.754 IAS

SAP vs. SL 0.608 0.859 0.570 0.834 0.627 0.939 ES

Rodula

RL ys. SL 1.29 0.472 DIN2AO625 1.40 0.501 OS

RL us. 1 (SV) 2.89 0.485 1.89 0.767 3.15 0.556 00%

5 Not significant at 0.02 level

"> Significant at 0.005 level

RL/SAA, and RL/%W(SV)) varied 9.6 and

28 % (Table III). Concerning the radula

relative size, P. ulyssiponensis was the

least variable species (Table II).

III. Discrimination between species

In order to determine which vari-

ables provided the best discrimination

between species, several canonical dis-

criminant analyses were carried out

using different techniques and different

sets of radula and/or shell characteris-

tics. Results are displayed in Tables IV -

VIL

Analysis I and Il, using only vari-

ables describing shell form, resulted in a

very ¡poor discrimination between

species, with only 65 % of the specimens

a posteriori correctly identified to species

(Table IV). The first canonical variable

accounted for 91 % of the variation

between species, and the variable with

the highest loading was SH/SL (data

not shown). Analysis HI, using only

variables describing radula relative size,

was better than analyses l and Il, and

was very satisfactory for P. ulyssiponen-

sis, but resulted in a low a posteriori

correct identification of P. intermedia

specimens (Table IV). The first canonical

variable accounted for 76 % of the varia-

tion between species, and the variables

with the highest loadings were RL/SH

followed by RL/*V(SV) (data not

shown) The simultaneous use of vari-

ables describing shell form and vari-

ables describing radula relative size

(Analysis IV) was better than the use of

one of these groups of variables alone

(Table IV). However, the improvement

of Analysis IV over Analysis II was

limited. A posteriori correct identification

of P. ulyssiponensis specimens was

unchanged (and very good), but identi-

fication of P. intermedia and P. vulgata

specimens was lower than 80 % (Table

IV). The first canonical variable

accounted for the great majority of the

variation between species, and the

second canonical discriminant variable

accounted for only 23.7 % of the vari-

ance (Table VI). The variables that pro-

vided the highest contrast between the

three Patella species were RL/SH fol-

lowed by RL/3V(SV), and then RL/SL

(Table VID. Individuals were plotted

along the two canonical variables

(Figure 7). There was a considerable

overlap between P. intermedia and P.

vulgata specimens, but a reasonable dis-

crimination of P. ulyssiponensis individu-

als. Stepwise selection of variables

describing shell form and radula rela-

CABRAL: Characterization and multivariate analysis of Patella spp.in Northwest Portugal

Table IM. Morphometric characterization of the species.

Tabla 11. Caracterización morfométrica de las especies.

Vorioble P. intermedia (n= 304) P. ulyssiponensis (n= 65) P. vulgata (n= 239)

Mean Y 0) Mean (00) Mean (V(%)

Shell

SW/SL 0.822 5.8 0.764 6.3 0.793 5.6

SH/SL 0.279 19 0.318 18 0.346 17

SAA/SAP 0.479 22 0.509 16 0.523 15

SAP/SL 0.682 7.0 0.665 10 0.658 5.4

Radula

RL/SL 1.68 19 0.934 12.4 1.55 20

RL/SH 6.24 26 2.99 14 4.54 20

RL/SAA 5.44 28 2.84 22 4.59 25

RL (SV) 4.32 19 2.35 9.6 3/3 19

Table IV. Number and percent correct identification of the specimens, based on canonical analyses

using different sets of shell and/or radula characters.

Tabla IV. Número y porcentaje de identificación correcta de los especímenes, basado en el análisis canó-

nico usando diferentes conjuntos de caracteres de concha y/o rádula.

P. intermedia (n= 304) P. ulyssiponensis (n= 65) P. vulgata (n= 239) Overall Mean

N % N % N % %

| SW/SL Sa 34 523 138 577 65.0

SH/SL

SMA/SAP

SW/SL IO 0 523 139 582 65.0

SH/SL

SAP/SL

Il RL/SL 202. 66.4 64 985 194 812 75.7

RL/SH

RL/SAA

RL/2V (SV)

V SH/SL O IA 64 985 187 782 17,5

SMA/SAP

RL/SL

RL/SH

RL/SAA

RL/2V (SV)

v SH/SL 3 64 985 187 782 77.8

RL/SL

RL/SH

RL/SM

REV (SV)

Analysis Variables

Iberus, 21 (2), 2003

Table V. Percent correct identification of the specimens based on canonical discriminant analysis

using shell and radula characters (SH/SL, SAA/SAB, RL/SL, RL/SH, RL/SAA, and RL/4V(SV).

Cross-validation analysis using half number of specimens for calibration and half for test.

Tabla V. Porcentaje de identificación correcta de los especímenes, basada en el análisis canónico discrimi-

nante usando caracteres de la rádula y concha (SH/SL, SAA/SAB RL/SL, RL/SH, RL/SAA, and

REAN(SV)). Análisis de validación cruzada usando la mitad de los especímenes para la calibración y la

otra mitad para el test.

P. intermedia P. ulyssiponensis P. vulgata Overall Mean

Mean 70.9 96.6 17.1 76.0

CV (0) 5.6 3.6 55) 38

Table VI. Eigenvalues and Wilks' A of canonical discriminant analysis using shell and radula cha-

racters (SH/SL, SAA/SAP, RL/SL, RL/SH, RL/SAA, RLPV(SV). Significance of Wilks' A was eva-

luated by Fisher's F values.

Tabla VI. Autovalores y A de Wilks del análisis canónico discriminante usando caracteres de la concha y

rádula (SH/SL, SAA/SAP RL/SL, RL/SH, RL/SAA, RLAN(SV)). La significatividad de la A de Wilks

se evaluó con los valores de la E de Eisher.

% of Variance

Eigenvalue A ela Canonical correlation Wilks A

First canonical variate 0.820 76.3 716.3 0.671 0.4395

Second canonical variate 0.255 2307) 100 0.450 DIN

2 Significant at 0.01 level

Table VII. Total canonical structure for discriminant analysis of Patella species. Variables with the

highest loadings are in bold.

Tabla VII. Estructura canónica total para el análisis discriminante de las especies de Patella. Las varia-

bles con mayores cargas están en negrita.

Voriable First canonical variable Second canonical variable

SH/SL 0.556 0.711

SAA/SAP 0.306 0.327

RL/SL 0.777 0.607

RL/SH 0.969 0,053

RL/SA 0.758 0.274

RL/AV(SV) 0.901 0.388

tive size (Analysis V, Table IV) resulted

in the removal of SAA/SAP, but the a

posteriori correct identification to species

was only very slightly improved. The

removal of SAA/SAP was expected

since it was the variable that provided

the lowest contrast between the three

limpet species (Table VID.

Analysis 1 — V were carried out using

a posteriori probabilities. In this tech-

nique, the same data set that is used to

derive the discriminant functions is also

CABRAL: Characterization and multivariate analysis of Patella spp.in Northwest Portugal

0 Patella intermedia

A Patella ulyssiponensis

OPatella vulgata

Second canonical variable

3 ES

2 Ú 12

First canonical variable

Figure 7. Scatter plot of the first and second canonical variables in the discriminant analysis using

shell and radula measurements (Analysis IV). The variables most closely associated with the dis-

crimination between species were RL/SH and RL/V(SV).

Fig. 7. Gráfica de la primera y segunda variables canónicas del análisis discriminatorio realizado uti-

lizando mediciones de la concha y la rádula (Análisis IV). Las variables que más fuertemente influen-

cian la discriminación entre especies fueron RL/SH y RLAN (SV).

used to test the accuracy of the predic-

tions using these functions. In order to

test the reliability of these results, an a

priori technique was used with a cross-

validation of the data. Mean percent-

ages of correct identification of the spec-

imens using this technique (Table V)

were very similar to those obtained with

a posteriori probabilities (Table IV, Analy-

sis IV), indicating high within-group

(species) homogeneity of specimens.

Only P. ulyssiponensis specimens were

very well correctly identified. Percent

correct identification of P. intermedia and

P. vulgata specimens were again lower

than 80 % (Table V).

DISCUSSION

I. Shell form

Results presented in this work

showed that P. intermedia, P. ulyssiponen-

sis and P. vulgata from Póvoa de Varzim

region have different shell growing pat-

terns. Although SW or SAA vs. SL regres-

sions showed slopes not significantly

different in these Patella species, log SH

vs. SL regressions displayed significantly

different slopes. The fastest growing

species in height was P. intermedia, and

the slowest P. ulyssiponensis, P. vulgata

displaying intermediate behaviour.

Iberus, 21 (2), 2003

TI. Radula relative size

Mean RL/SL ratios for P. intermedia,

P. ulyssiponensis and P. vulgata reported

in the literature for several coastal re-

gions of Spain, France, Great Britain and

Ireland display the following ranges:

1.60-2.20, 0.94-1.20, 1.29-2.29, respec-

tively (FISHER-PIETTE, 1934, 1935, 1941,

1948, Evans, 1947, 1953, 1958; BRIAN AND

OWEN, 1952; FISHER-PIETTE AND GAIL-

LARD, 1959; CHRISTIAENS, 1973; IBAÑEZ,

1982; FELIU AND IBANEZ, 1984). Mean

RL/2V(SV) ratios for P. intermedia, P.

ulyssiponensis and P. vulgata in several

coastal regions of Spain, France, and

Great Britain show the following varia-

tion: 4.66-4.71, 2.87-2.90, 3.90-4.38, re-

spectively (Fisher-Piette, 1941, 1948).

Our results for Póvoa de Varzim (north-

west Portugal) (Table III) were within, or

very near these ranges.

P. ulyssiponensis is usually the least

variable of these three limpets, if we

consider the inter-site and within-site

variability of the radula relative size

(FISHER-PIETTE, 1934, 1935, 1941, 1948,

EVANS, 1953, 1958; IBANEZ, 1982; FELIU

AND IBAÑEZ, 1984). This conclusion is

illustrated by data reported by IBAÑEZ

(1982), IBAÑEZ AND FELIU (1983) and

FELIU AND IBAÑEZ (1984) on several

Patella species in the Basque coast.

RL/SL ratios were determined at

several sites in this region, and on dif-

ferent months and different levels, at

each site. P. ulyssiponensis displayed

similar RL/SL ratios along the year and

between different levels in the beach.

On the contrary, RL/SL ratios for P.

intermedia and P. vulgata varied appre-

ciably along the year, between sites and

between different levels at each site.

Generally, RL/SL ratios increased from

winter to summer months. Our results

for these species in Póvoa de Varzim

agree with this conclusion. The coeffi-

cient of variation for the means of all

variables describing the radula relative

size were lowest in P. ulyssiponensis.

Data reported in the literature indi-

cate that in the genus Patella, the radula

relative size decreases, with lowering the

level of the occurrence of the animal in

the shore. This general trend has been

observed both within each species, and

between several species. BRIAN AND

OWEN (1952) studied the variability of

the RL/SL ratio for P. vulgata in the west

coast of Great Britain. Whilst animals

from high-water habitats, above high-

water neaps, displayed RL/SL ratios in

the range 1.83-1.96, animals from low-

water habitats, about or below low-wa-

ter neaps, showed RL/SL ratios between

1.38-1.61. DAviÉS (1969) observed that in

two sites in the Italian coast, intertidal

populations of Patella caerulea Linné 1758

displayed longer radula than submerged

populations. SELLA (1976) studied the

variability of the RL/SL and RL/SH ra-

tios in Patella aspera Róding 1798 (syn-

onym P. ulyssiponensis) and P. caerulea

from several Mediterranean stations in

the Tyrrhenian Sea. In both species, both

ratios decreased from animals living in

the mesolittoral zone (50 cm above to 50

cm below water level) to animals living

in the infralittoral region (6 to 26 m

depth). In P. ulyssiponensis, the RL/SL ra-

tio decreased, from the mesolittoral to

the infralittoral zones, from 0.80 to 0.66

at one station, and from 0.90 to 0.64 at

another site. The RL/SH ratio decreased

from 3.2 to 3.0 and from 3.4 to 3.0, at the

same sites and depths, respectively. Our

observations that radula relative size

was more variable than shell form, could

have been due to this dependence of the

radula relative size on the position of the

animal in relation to the water level,

since the animals were collected from the

low shore to high shore levels, and all

specimens of each species analysed as a

unique set. However, the coefficient of

variation of mean RL/SL and RL/“V(SV)

ratios was at maximum 20 %, a measure

of variability not uncommonly found in

biological populations. The exact extent

of the variability of the radula relative

size is expected to vary between sites in

the European coasts, considering the

polymorphism of the shorelines and the

variability of tides in this area.

IBAÑEZ (1982) and IBANEZ AND FELIU

(1983) found that in the Basque Coast,

Patella rustica Linné 1758, P. vulgata, Patella

depressa Pennant 1777 (synonym P. inter-

CABRAL: Characterization and multivariate analysis of Patella spp.in Northwest Portugal

media) and P. aspera (P. ulyssiponensis)

occupy successively lower positions in the

coast. Mean RL /SL ratios for these species

were 3.32, 1.76, 1.60-2.18, 0.986, respec-

tively. HERNÁNDEZ-DORTA (1992) reported

that in the Canary Islands, Patella candet

d'Orbigny 1840, Patella piperata Gould

1846, P. ulyssiponensis and Patella crenata

Gmelin 1791 (synonym P. ulyssiponensis)

habitats decreased in height in the shore-

line. Mean RL/SL ratios found for these

species in the region were 1.63, 2.29, 0.74

and 1.12, respectively. This general trend

of decreasing radula size from mesolit-

toral to the infralittoral species has also

been found at Póvoa de Varzim. P.

ulyssiponensis occupies the lowest posi-

tions in the shore, and displayed the

lowest RL/SL and RL/%V(SV) ratios.

TIT. Discrimination between species

Our results presented in this work

showed that, in the discrimination

between P. intermedia, P. ulyssiponensis

and P. vulgata from Póvoa de Varzim,

variables describing the radula relative

size were more important than variables

describing shell form. Whilst using only

variables related to shell form, the mean

a posteriori correct identification to

species was 65.0 %, using only variables

describing the radula relative size this

value rose to 75.7 %. Our results there-

fore confirm the observations reported

by several malacologists (see Introduc-

tion) on the importance of the character-

istics Of the radula to the discrimination

between P. intermedia, P. ulyssiponensis

and P. vulgata.

The RL/SL ratio is considered the

most important morphometric variable

for discriminating European Patella

species. HERNÁNDEZ-DORTA (1992) com-

paring P. candei, P. piperata and P.

ulyssiponensis from the Canary Islands

concluded that, from several shell and

radular variables, RL/SL was the most

discriminating variable, followed by

SH/SL. However, in the present work,

for the discrimination between P. inter-

media, P. ulyssiponensis and P. vulgata

from Póvoa de Varzim, the most impor-

tant variable was RL/SH, both in the

analysis using only radula relative size

variables and in the analysis using both

shell form and radula size variables.

RL/*V(SV) was the variable with the

second highest loading, and RL/SL the

third. This is a new finding, and can be

due to the use, in the present analysis, of

a wider range of variables describing

the radula relative size, four instead of

one or two commonly used. Alterna-

tively, our results can be related to a

specificity of these species at this site.

This observation that RL/SH was

the most discriminating variable

between P. intermedia, P. ulyssiponensis

and P. vulgata confirms other results also

found in this study on the importance of

the SH in the discrimination between

these species. Using only variables

describing shell form, SH/SL was the

most discriminating variable. Whilst

slopes of the regressions, SW or SAA vs.

SL were not statistically different, slopes

of log SH vs. SL regression lines were

significantly different.

The best discrimination between P.

intermedia, P. ulyssiponensis and P. vulgata

from Póvoa de Varzim was achieved by

using shell form and radula relative size

variables. However, the overall mean

correct identification to species (using a

priori or a posteriori probabilities) was be-

low 80 %, and only P. ulyssiponensis spec-

imens were correctly identified in high

percentage (higher than 95 %). It appears

therefore that quantitative characters re-

lated to shell form and radula size are

relatively poor discriminators between P.

intermedia and P. vulgata. Qualitative

characters are thus still indispensable for

the discrimination between these two

Patella species. Very good qualitative dis-

criminators were, for P. intermedia, the

tall and broad cusp 2 and the dark mar-

ginal rays in the shell interior surface,

and for P. vulgata, the pointed protuber-

ance on cusp 3, and the silvery head scar.

IV. The use of Canonical Discrimi-

nant Analysis in the discrimination

between Patella species

Canonical discriminant analysis has

been used in the literature to compare

Iberus, 21 (2), 2003

limpet species. These analyses have

used variables describing the shell and

the radula - shell size and radula ab-

solute size variables alone, or simultane-

ously, with shell form and radula rela-

tive size variables, which were the vari-

ables exclusively used in the present

work. The use of variables describing

shell and radula absolute sizes in the

comparison between limpet species has

however some limitations, because

limpets present extremely variable pop-

ulations structure. LEWIS AND BOWMAN

(1975) and THOMPSON (1980) reported

detailed studies on the biology and pop-

ulation dynamics of P. vulgata in several

sites Of the England and Ireland coasts.

Very marked differences in length-fre-

quency distributions were observed be-

tween populations occupying different

tidal levels and different habitats.

GUERRA AND GAUDENCIO (1986) studied

several populations of P. aspera (P.

ulyssiponensis), P. depressa (P. intermedia)

and P. vulgata from the Portuguese

coast, with monthly sampling. All these

three species exhibited marked varia-

tions in the population structure

throughout the year, and at a given

month, these limpets showed different

populations structures. These differ-

ences in populations structures have

been interpreted as the result of a com-

plex set of factors - gonad cycles,

spawning and settlement periods,

growth rates, survival, mortality and

longevity, or in brief, as the result of

population history (LEwIS AND

BOwMAN, 1975; THOMPSON, 1980;

GUERRA AND GAUDENCIO, 1986). There-

'm indebted to Dr. Joio Manuel

Martins for helpful discussions and com-

ments on the manuscript.

BIBLIOGRAPHY

BRIAN, M. V. AND OWEN, G., 1952. The relation

of the radula fraction to the environment in

Patella. Journal of Animal Ecology, 21: 241-249.

CHRISTIAENS, J., 1973. Révision du genre Patella

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National d'Histoire Naturelle, 182: 1305-1392.

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compared have similar structures,

namely percentage of juveniles and

adults, and sampling is exhaustive and

based on representatives of all age-

classes, comparisons between variables

describing shell and radula absolute sizes

might not represent real differences be-

tween species.

CONCLUSIONS

P. intermedia, P. ulyssiponensis and P.

vulgata from Póvoa de Varzim region

have different shell growing patterns.

The best discrimination between P.

intermedia, P. ulyssiponensis and P. vulgata

was achieved by using shell form and

radula relative size variables. However,

whereas correct identification of P.

ulyssiponensis specimens was higher

than 95 %, the identification of P. inter-

media and P. vulgata specimens ranged

from 70 to 80 %. Qualitative characters

describing the radula and the shell are

thus still indispensable for the discrimi-

nation between these two limpet

species.

Morphometric characteristics such

as those utilized in this work are proba-

bly useful for characterization of P.

ulyssiponensis as a species, but are of

limited value for P. intermedia and P.

vulgata.

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phological and molecular resolution of a

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Hall, New Jersey, 718 pp.

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O Sociedad Española de Malacología ——____——T— lIberus, 21 (2): 19-27, 2003

Toxic eftect of Thevetia peruviana and Alstonia scholaris lati-

ces on the freshwater snail Lymnaea acuminata

Toxicidad de los látex de Thevetia peruviana y Alstonia scholarís sobre

el molusco dulceacuícola Lymnaea acuminata

Sunil Kumar SINGH* and Ajay SINGH*!

Recibido el 4-X-2002. Aceptado el 21-1-2003

ABSTRACT

The aqueous and partially purified latex extracts of plants Thevetia peruviana and Alstonia

scholaris (Family Apocynaceae) have potent molluscicidal activity. Sub-lethal doses of

aqueous and partially purified latex extracts of both the plants also significantly alter the

levels of total protein, total free amino acid, nucleic acid (DNA and RNA) and the activity

of enzyme protease, acid and alkaline phosphatase in nervous tissue of the snail lymnaea

acuminata in time and dose dependent manner. The biologically active compounds pre-

sent in Thevetia peruviana plant are Apigenine (Flavonoid) and triterpenoid glycosides,

while a number of alkaloids (i.e. pseudo-akuammigine, Betulin, Ursolic acid and B-sitoste-

rol etc.), steroids and triterpenoids are present in Alstonia scholaris plant.

RESUMEN

Los extractos de latex acuosos y parcialmente purificados de las plantas Thevetia peru-

viana y Alstonia scholaris [Familia Apocynaceae) tienen potente actividad molusquicida.

Dosis subletales de dichos extractos alteran significativamente los niveles de proteínas

totales, aminoácidos libres, ácidos nucléicos y actividad de los enzimas proteasa, fosfa-

tasa ácida y alcalina en el tejido nervioso de Lymnaea acuminata de manera tiempo-

dependiente y dosis-dependiente. Los componentes activos de Thevetia peruviana son Api-

genina [Flavonoide) y glicósidos triterpenoides, mientras que en Alstonia scholaris

aparecen un cierto número de alcaloides (i.e. pseudo-akuammigina, betulina, ácido urso-

lico y B-sitosterol entre otros), esteroides y triterpenos.

KEY WORDS: Lymnaea acuminata, Thevetia peruviana, Alstonia scholarís, metabolism.

PALABRAS CLAVE: Lymnaea acuminata, Thevetia peruviana, Alstonia scholaris, metabolismo.

INTRODUCTION

The use of synthetic or petroleum REDINGER, 1976; SUSAN, VEERAIAH AND

based molluscicides for controlling TIiLAK, 1999). To overcome the problem

vector snails cause serious environmen- and to search for eco-friendly mollusci-

tal pollution (MIAN AND MULLA, 1992; cides, a number of extracts and essential

* Natural Products Laboratory, Department of Zoology, D.D.U. Gorakhpur University, Gorakhpur-273 009

(UP), INDIA E- mail: ajay_sCsancharnet.in

' Corresponding Author

Iberus, 21 (2), 2003

oils and their isolates have been evalua-

ted for use as molluscicides due to their

high toxicity, easy availability and easy

biodegradability (MARSTON AND HOS-

TETTMANN, 1985; KINGHORN AND EVANS,

1975; SINGH, SINGH, MISRA AND

AGARWAL, 1996). These materials have

shown encouraging results for vector

controlling properties with various snail

species.

Thevetia peruviana and Alstonia scho-

laris, are common medicinal plants of

family Apocynaceae. The latex of Theve-

tia peruviana is used in teeth cavities for

relief from toothache and the latex of

Alstonia scholaris is applied to ulcer,

sores, tumers and rheumatic pain and is

used for curing toothache (RAMA RAO,

1967). Both the plants have potent

molluscicidal and anti-cholinesterase

activity against harmful snails and slugs

(PANIGRAHI AND RAUT, 1994; SINGH,

YADAV AND SINGH, 2000; SINGH AND

SINGH, 2002). The mechanism by which

the active compounds present in both

the plants causes snail death is not

known.

The toxicological actions of Thevetia

peruviana, may be due to the presence of

apigenin-5-methyl ether (flavonoid) and

triterpenoid glycosides (VOIGTLANDER

AND BALSAM, 1970) while a number of

alkaloids (pseudo-akuammigine in

addition to betulin, ursolic acid and Í3-

sitosterol), steroids and triterpenoids are

present in Alstonia scholaris (BANERJI

AND BANERJL, 1977).

We are interested in knowing the

mode of action and long-term effect of

these plant products on snails, because

these substances cannot be put to com-

mercial use without a study of these

aspects as well. In the present study, the

effect of sub-lethal doses of aqueous

and partially purified latex extracts of

Thevetia peruviana and Alstonia scholaris

is examined on nitrogenous metabolism

of freshwater snail Lymnaea acuminata.

This snail is intermediate host of liver

fluke Fasciola hepatica and Fasciola gigan-

tica, which causes endemic fascioliasis

in cattle and live- stock in northern

parts of India (SINGH AND AGARWAL,

198):

MATERIALS AND METHODS

Latex of Thevetia peruviana and Alsto-

nia scholaris were collected from Botani-

cal garden of D.D.U. Gorakhpur Univer-

sity, Gorakhpur, India. White latex pro-

duced by these plants was drained into

glass tubes fallowing cutting of the stem

apices and lyophilised at 40"C and the

lyophilised dry powder was used for

further study. The wet weight of one ml

of latex of Thevetia peruviana and Alsto-

nia scholaris was 820 mg and 1000 mg

and dry weight was 340 mg and 400 mg,

respectively.

For aqueous extracts: The freeze-

dried powder was mixed with appro-

priate volume of distilled water to

obtain the desired concentrations.

For partial purification: The lyophi-

lised latex powder was extracted

sequentially with 5 mL each of chloro-

form -> carbon tetrachloride > acetone

> diethyl ether > ethyl alcohol. Centri-

fugation for 20 min at 2000 g. was

carried out in a refrigerated centrifuge,

at -4'C, after each extraction. The

solvent fraction was decanted and the

solvent allowed to evaporates. The

dried up soluble fraction was re-dissol-

ved in water for further experiments.

Adult snail Lymnaea acuminata

(2.6+0.3 cm in shell height) was collec-

ted locally and used as test animals. The

snails were maintained and treated with

aqueous and partially purified extracts

of latices of the test plants according to

SINGH AND AGARWAL (1990). Adult

Lymnaea acuminata were kept in glass

aquaria containing 3 litres of de-chlori-

nated tap water. Each aquarium contai-

ned 20 experimental animals.

Treatment protocol for dose- res-

ponse relation

Snails were exposed for 24h or 96h

to 40% and 80% of LCso doses of Thevetia

peruviana and Alstonia scholaris latices.

LCso doses were determined in an

earlier study by SINGH (2000). The 24h

or 96h doses for Thevetia peruviana were,

respectively, 0.43 mg/L and 0.17 mg/L

and for Alstonia scholaris were 4.76

mg/L and 1.76 mg/L.

SINGH AND SINGH: Toxic effect of Apocynaceae latices on Lymnaea acuminata

After completion of treatment the

test animals were removed from the

aquaria and washed with water. The

nervous tissue of Lymnaea acuminata was

excised and used for biochemical analy-

sis. Control animals were held in similar

conditions without any treatment. Each

experiment was replicated at least six

times and the values have been expres-

sed as mean +SE of six replicates. Stu-

dent's “t' test and analysis of variance

were applied to locate significant

changes (SOKAL AND ROHLEE, 1973).

Biochemical estimation

Protein: Protein levels were estima-

ted according to the method of Lowry,

ROSEBROUGH, FARR AND RANDALL (1951)

using bovine serum albumin as stan-

dard. Homogenates (5 mg/mL, w/v)

were prepared in 10% Trichloroacetic

acid (ICA).

Total free amino acids: Estimation

of total free amino acid was made accor-

ding to the method of SPICES (1957).

Homogenates (10 mg/mL, w/v) were

prepared in 95% ethanol, centrifuged at

6000 g and used for amino acid estima-

tion.

Nucleic acids: Estimation of nucleic

acid (DNA and RNA) was performed,

by the methods of SCHNEIDER (1957)

using diphenylamine and orcinol rea-

gents, respectively. Homogenates (1

mg/mL, w/v) were prepared in 5%

TCA at 90*C, centrifuged at 5000 g for

20 min and supernatant was used for

estimation of nucleic acids.

Protease: Protease activity was esti-

mated by the method of MOORE AND

STEIN (1954). Homogenate (50 mg/mL,

w/v) was prepared in cold distilled

water. Optical density was measured at

570 nm. The enzyme activity was

expressed in m moles of tyrosine equi-

valent/mg protein/h.

Acid and alkaline phosphatase: The

activity of acid and alkaline phospha-

tase in the nervous tissue was determi-

ned, according to the method of

ANDERSCH AND SZCYPINSKI (1947) as

modified by BERGMEYER (1967) and

SINGH AND AGARWAL (1983). Tissue

homogenates (2% w/v) were prepared

in ice-cold 0.9% sodium chloride solu-

tion and centrifuged at 5000 g at (0%C)

for 15 min. Optical density was measu-

red at 420 nm against a blank, prepared

simultaneously. The enzyme activity has

been expressed as amount of p-nitrop-

henol formed/30min/mg protein in

supernatant.

RESULTS

Data of sub-lethal (40% and 80% of

LC50o) exposure of freshwater snail

Lymnaea acuminata against aqueous and

serially extracted latex of Thevetia peru-

viana and Alstonia scholaris for 24h and

96h on nitrogenous metabolism in

nervous tissue are given in Table I-IV.

Thevetia peruviana

Exposure of snails to 40% and 80%

of LCso of aqueous latex extracts of The-

vetia peruviana for 24h or 96h caused sig-

nificant alterations in nitrogenous meta-

bolism in nervous tissue of the freshwa-

ter snail Lymnaea acuminata (Table I-II.

Total protein levels were reduced to 54%

and 38% of controls, respectively after

exposure to 40% and 80% of LCso (24h)

of aqueous latex extract. The maximum

decrease in protein level (30 % of con-

trol) was observed in snails treated with

80% of LCso (96h) of aqueous latex ex-

tract. DNA level was reduced to 81%

and 71% of controls after treatment with

40% and 80% of LC5o (24h), respectively.

The maximum decrease in DNA (35 %

of control) was observed in snails trea-

ted with 80% of LCso (96h) of aqueous

latex extract. RNA level was reduced to

88% and 70% of controls after treatment

with 40% and 80% of LCs5o (24h) of aque-

ous latex extracts respectively in ner-

vous tissue of Lymnaea acuminata. The

maximum decrease in RNA (42 % of

control) was observed in snails treated

with 80% of LCso (96h) of aqueous latex

extract. Total free amino acid levels were

induced to 149% and 166% of controls

after treatment with 40% and 80% of

LCso (96h) of aqueous latex extracts res-

pectively in nervous tissue of snail Lym-

naea acuminata (Table 1-ID.

Zal

Iberus, 21 (2), 2003

Table I. Changes in total protein, total free amino acids, nucleic acid (DNA and RNA) (ug/mg),

protease (ug moles of tyrosine equivalents/mg protein/h) and acid and alkaline phosphatase (p

mole substrate hydrolysed/30 min/mg protein) level in nervous tissue of Lymnaea acuminata after

exposure to 40% and 80% of LCso of aqueous and partially purified latex extracts of Thevetia

peruviana for 24 h.

Tabla 1. Cambios en los niveles de proteína total, aminoácidos libres, ácidos nucléicos (DNA y RNA)

(ug/mg), proteasa (ug moles de equivalentes de tirosinalmg proteínalh) y fosfatasas ácida y alcalina (y

moles sustrato hidrolizado/30 min/mg proteína) en el tejido nervioso de Lymnaea acuminata tras 24 h

de exposición a extractos acuosos y parcialmente purificados de latex de Thevetia peruviana al 40% y

80% de LC5o.

Nature of latex Control 40% of LC50 80% of LC50

Protein A 65.00+0.28 (100) 34.80+0.36+ (54) 24.50+0.38+ (38)

B 65.30+0.78 (100) 35.50+0.68+ (54) 27.50+0.84+ (42)

Amino acid A 34.60+0.36 (100) 48.50+0.68+ (140) 45.60+0.76+ (152)

B 34.40+0.65 (100) 51.20+0.77+ (137) 47.50+1.02+ (146)

DNA A 75.40+1.02 (100) 60.90+0.88+ (81) 53.70+0.70+ (71)

B 75.60+1.12 (100) 62.80+1.02+ (83) 55.50+1.05+ (73)

RNA A 60.10+0.52 (100) 52.80+0.60+ (88) 42.06+0.52+ (70)

B 61.20+0.82 (100) 54.60+0.70+ (88) 44.05+0.96+ (72)

Protease A 0.325+0.067 (100) 0.378+0.052+ (116) 0.420+0.048+ (129)

B 0.325+0.046 (100) 0.375+0.062+ (115) 0.415+0.062+ (128)

Acid phosphatase A 0.193+0.0005 (100) 0.171+0.0004+ (89) 0.148+0.0003+ (77)

B 0.193+0.0010 (100) 0.178+0.0003+ (92) 0.152+0.0006+ (79)

Alkaline phosphatase A 0.381+0.0015 (100) 0.338+0.0006+ (89) 0.251+0.0005+ (66)

B 0.382+0.0012 (100) 0.344+0.0005+ (90) 0.254+0.0003+ (66)

Values are mean +SE of six replicates

Values in parenthesis are % change with control taken as 100%

+, Significant (P<0.05) student's '1' test was applied between control and treated groups

A — Supernatant of aqueous solution of latex

B — Latex serially extracted through chloroform, carbon tetrachloride, acetone, diethyl ether and ethyl alcohol

Activity of acid phosphatase was

inhibited to 89% and 77% of controls

after treatment with 40% and 80% of

LCso (24h) of aqueous latex extracts res-

pectively in nervous tissue of snail. Acti-

vity of alkaline phosphatase was

reduced to 89% and 66% of controls

after treatment with 40% and 80% of

LC50 (24h) of aqueous latex extracts res-

pectively in nervous tissue of snail. The

maximum decrease in acid and alkaline

phosphatase 37 % and 45% of controls,

respectively, was observed in snails

treated with 80% of LCs5o (96h) of

aqueous latex extract. Protease activity

was increased to 116% and 129% of con-

trols after treatment with 40% and 80%

of LCso (24h) of aqueous latex extracts

respectively in the nervous tissue of

snail Lymnaea acuminata. The maximum

increase in protease activity (141% of

control) was observed in snails treated

with 80% of LCso (96h) of aqueous latex

extract (Table 1-ID.

Latex was sequentially extracted

with organic solvents also caused a

similar alteration in nitrogenous meta-

bolism of snail. Alterations caused by

SINGH AND SINGH: Toxic effect of Apocynaceae latices on Lymnaea acuminata

Table IL. Changes in total protein, total free amino acids, nucleic acid (DNA and RNA) (ug/mg),

protease (ug moles of tyrosine equivalents/mg protein/h) and acid and alkaline phosphatase (ul

mole substrate hydrolysed/30 min/mg protein) level in nervous tissue of Lymnaea acuminata after

exposure to 40% and 80% of LCso of aqueous and partially purified latex extracts of Thevetía

peruviana for 96 h.

Tabla 11. Cambios en los niveles de proteína total, aminoácidos libres, ácidos nucléicos (DNA y RNA)

(ug/mg), proteasa (ug moles de equivalentes de tirosinalmg proteína/h) y fosfatasas ácida y alcalina (p

moles sustrato hidrolizado/30 min/mg proteína) en el tejido nervioso de Lymnaea acuminata tras 96 h

de exposición a extractos acuosos y parcialmente purificados de latex de Thevetia peruviana al 40% y

80% de LC5o.

Nature of latex Control 40% of LC50 80% of LC50

Protein A 68.50+0.48 (100) 30.50+0.44* (44) 20.80+0.44* (30)

B 68.40+0.76 (100) 31.60+0.72* (46) 21.60+0.48* (32)

Amino acid A 35.30+0.98 (100) 52.50+0.26* (149) 58.60+0.36* (166)

B 35.30+0.98 (100) 40.50+0.46* (143) 56.70+0.78* (161)

DNA A 72.16+0.93 (100) 39.50+0.46* (55) 25.20+0.39* (35)

B 72.20+0.05 (100) 40.40+0.78* (56) 28.30+0.98* (39)

RNA A 62.50+1.02 (100) 36.40+0.45* (58) 26.45+0.78* (42)

B 62.80+1.12 (100) 38.50+0.78* (61) 28.35+0.98* (45)

Protease Á 0.345+0.058 (100) 0.420+0.072* (122) 0.186+0.025* (141)

B 0.346+0.068 (100) 0.418+0.075* (121) 0.128+0.032* (138)

Acid phosphatase A 0.192+0.0003 (100) 0.096+0.0005* (50) 0.071+0.0006* (37)

B 0.191+0.0004 (100) 0.101-0.0007* (53) 0.074+0.0011*(39)

Alkaline phosphatase A 0.400+0.0007 (100) 0.187+0.0010*(47) 0.181+0.0007*(45)

B 0.390+0.0015 (100) 0.194+0.0004* (50) 0.184+0.0004*(47)

Details are as given in Table |

sequentially extracted latex, though sta-

tistically in significant, appeared lower

than the water extracted latex at all the

doses and exposure periods (Table I-ID.

Alstonia scholaris

Exposure of snails to 40% and 80% of

LCso of aqueous latex extracts of Alstonia

scholaris for 24h or 96h caused significant

alterations in nitrogenous metabolism in

nervous tissue of the freshwater snail

Lymnaea acuminata (Table IM-IV). Total

protein levels were reduced to 55% and

40% of controls, respectively after expo-

sure to 40% and 80% of LCso (24h) of

aqueous latex extract. The maximum de-

crease in protein level (35 % of control)

was observed in snails treated with 80%

of LC5o (96h) of aqueous latex extract.

DNA level was reduced to 85% and 75%

of controls after treatment with 40% and

80% of LCso (24h), respectively. The ma-

ximum decrease in DNA (39 % of con-

trol) was observed in snails treated with

80% of LCso (96h) of aqueous latex ex-

tract. RNA level was reduced to 90% and

75% of controls after treatment with 40%

and 80% of LCso (24h) of aqueous latex

extracts respectively in nervous tissue of

Lymnaea acuminata. The maximum decre-

ase in RNA (45 % of control) was obser-

ved in snails treated with 80% of LCso

(96h) of aqueous latex extract. Total free

amino acid levels were induced to 145%

and 160% of controls after treatment

with 40% and 80% of LCso (96h) of aque-

ous latex extracts respectively in nervous

tissue of snail (Table II-IV).

Iberus, 21 (2), 2003

Table III. Changes in total protein, total free amino acids, nucleic acid (DNA and RNA) (1g/mg),

protease (ug moles of tyrosine equivalents/mg protein/h) and acid and alkaline phosphatase (p

mole substrate hydrolysed/30 min/mg protein) level in nervous tissue of Lymnaea acuminata after

exposure to 40% and 80% of LCso of aqueous and partially purified latex extracts of Alstonia schol-

aris for 24 h.

Tabla III. Cambios en los niveles de proteína total, aminoácidos libres, ácidos nucléicos (DNA y RNA)

(ug/mg), proteasa (ug moles de equivalentes de tirosinalmg proteínalh) y fosfatasas ácida y alcalina (y

moles sustrato hidrolizado/30 min/mg proteína) en el tejido nervioso de Lymnaea acuminata tras 24 h

de exposición a extractos acuosos y parcialmente purificados de latex de Alstonia scholaris al 40% y

80% de LC5o.

Nature of latex Control 40% of LC50 80% of LC50

Protein A 66.20+0.56 (100) 36.50+0.76* (55) 26.60+0.98* (40)

B 66.50+0.78 (100) 38.20+0.56* (57) 29.90+0.78* (45)

Amino acid A 35.50+0.57 (100) 47.90+0.78* (135) 52.50+0.68* (148)

B 36.20+0.56 (100) 47.80+1.05* (132) 52.50+0.68* (145)

DNA A 74.80+1.05 (100) 63.60+0.54* (85) 56.10+0.58* (75)

B 75.20+1.06 (100) 65.40+0.96* (87) 57.20+0.46* (76)

RNA A 61.20+1.06 (100) 55.10+0.58* (90) 45.90+0.78*(75)

B 60.10+0.54 (100) 54.70+0.82* (91) 46.30+1.02* (77)

Protease A 0.335+0.072 (100) 0.382+0.061*(114) 0.422+0.065* (126)

B 0.328+0.062 (100) 0.367+0.052* (112) 0.407+0.082*(124)

Acid phosphatase A 0.192+0.0003 (100) 0.172+0.0004* (89) 0.148+0.0007* (77)

B 0.192+0.0002 (100) 0.175+0.0004* (91) 0.150+0.0006* (78)

Alkaline phosphatase A 0.400+0.0010 (100) 0.366+0.0005* (92) 0.302+0.0005*(76)

B 0.400+0.0005 (100) 0.372+0.0012*(93) 0.308+0.0003*(77)

Details are as given in Table |

Activity of acid phosphatase was in-

hibited to 89% and 77% of controls after

treatment with 40% and 80% of LCs5o

(24h) of aqueous latex extracts respecti-

vely in nervous tissue of snail. Activity

of alkaline phosphatase was reduced to

92% and 76% of controls after treatment

with 40% and 80% of LCso (24h) of aque-

ous latex extracts respectively in ner-

vous tissue. Protease activity was incre-

ased to 114% and 126% of controls after

treatment with 40% and 80% of LCso

(24h) of aqueous latex extracts respecti-

vely in the nervous tissue of snail Lym-

naea acuminata. The maximum increase

in protease activity (140% of control)

was observed in snails treated with 80%

of LCso (96h) of aqueous latex extract,

respectively (Table HI-IV).

Latex of Alstonia scholaris was

sequentially extracted with organic sol-

vents also caused a similar alteration in

nitrogenous metabolism of snail. Altera-

tions caused by sequentially extracted

latex, though statistically in significant,

appeared lower than the water extracted

latex at all the doses and exposure

periods (Table HL-IV).

DISCUSSION

It is clear from the result described

above that the treatment with sub-lethal

doses of aqueous and partially purified

latex extracts of Thevetia peruviana and

Alstonia scholaris, significantly alter the

level of total protein, total free amino

SINGH AND SINGH: Toxic effect of Apocynaceae latices on Lymnaea acuminata

Table IV. Changes in total protein, total free amino acids, nucleic acid (DNA and RNA) (1g/mg),

protease (ug moles of tyrosine equivalents/mg protein/h) and acid and alkaline phosphatase (p

mole substrate hydrolysed/30 min/mg protein) level in nervous tissue of Lymnmaea acuminata after

exposure to 40% and 80% of LCso of aqueous and partially purified latex extracts of Alstonia schol-

aris for 96 h.

Tabla IV. Cambios en los niveles de proteína total, aminoácidos libres, ácidos nucléicos (DNA y RNA)

(ug/mg), proteasa (ng moles de equivalentes de tirosinalmg proteínalh) y fosfatasas ácida y alcalina (y

moles sustrato hidrolizado/30 min/mg proteína) en el tejido nervioso de Lymnaea acuminata tras 96 h

de exposición a extractos acuosos y parcialmente purificados de latex de Alstonia scholaris al 40% y

80% de LC5o.

Nature of latex Control 40% of LC50 80% of LC50

Protein A 65.60+0.48 (100) 30.20+0.84* (46) 22.90+0.78* (35)

B 66.20+0.58 (100) 31.80+0.96* (48) 23.80+0.48* (36)

Amino acid A 35.60+0.76 (100) 51.60+0.88* (145) 56.90+0.78* (160)

B 36.50+0.54 (100) 51.80+0.46* (142) 57.30+0.66* (157)

DNA A 74.80+1.05 (100) 44.90+0.78* (60) 29.20+0.48* (39)

B 75.60+1.08 (100) 46.90+0.78* (62) 31.00+1.02* (41)

RNA A 60.10+0.74 (100) 36.10+0.74* (60) 27.00+0.48* (45)

B 61.50+0.78 (100) 38.71+1.06* (63) 29.00+0.48* (47)

Protease A 0.340+0.076 (100) 0.408+0.035* (120) 0.476+0.076*(140)

B 0.338+0.066 (100) 0.399+0.067* (118) 0.463+0.064* (137)

Acid phosphatase A 0.193+0.0005 (100) 0.094+0.0003* (49) 0.074+0.1133* (38)

B 0.192+0.0008 (100) 0.099+0.0007* (52) 0.072+0.0006* (38)

Alkaline phosphatase A 0.380+0.0007 (100) 0.188+0.0010*(49) 0.145+0.00038* (38)

B 0.390+0.0007 (100) 0.195+0.0009*(50) 0.148+0.0004* (39)

Details are as given in Table |

acid and nucleic acid and activity of

enzyme protease, acid and alkaline

phosphatase. The rate of alteration in all

the cases was significantly (P<0.05) time

and dose dependent.

Proteins are mainly involved in

architecture of the cell. During chronic

periods of stress they are also a source

of energy (UMMINGER, 1977). The decre-

ase in protein level observed in this

study may be due to their degradation

and also to their possible utilization for

metabolic purposes. BRADBURY,

SYMONIC, COATS AND ATCHISON (1987)

pointed out that the decrease protein

content might also be attributed to the

destruction or necrosis of cells and con-

sequent impairment in protein synthesis

machinery.

The quantity of protein depends on

the rate of protein synthesis or its degra-

dation. It also affected due to impaired

in corporation of amino acids in to poly-

peptide chains (SINGH ET AL., 1996). The

synthesis of RNA plays an important

role in protein synthesis. The inhibition

of RNA synthesis at transcription level,

thus may affect the protein level. In this

study, a significant decline in RNA level

in exposed snail was observed. The

decrease in RNA level may also have

been a cause of protein depletion. On

the other hand, increase in protease acti-

vity may be the cause of increased

protein degradation.

Enhanced protease activity and

decreased protein level have resulted in

marked elevation of free amino acids

Iberus, 21 (2), 2003

that in the snail tissue. The accumula-

tion of free amino acids can also be attri-

buted to the lesser use of amino acids

(SESHAGIRI RAO, SRINIVAS, MOORTHY,

SWAMY AND CHETHY, 1987) and their

involvement in the maintenance of an

acid-base balance (MOORTHY, KASHI

REDDY, SWwAMY AND CHETHY, 1984).

NATARAJAN (1985) reported that stress

condition in general induces elevation

in the trans-amination pathway.

Extracts of both the plants also

decreased the level of nucleic acids sig-

nificantly in the nervous tissue of the

snail. Several reports are available on

the reduction in DNA and RNA level on

exposure to different pesticides (TARIG,

HAQUI AND ADHAMI, 1977; NORDENSK-

JOLD, SODERHALL AND MOLDEUS, 1979).

Data attained in this study made it clear

that these plant extracts are potential

inhibitor of DNA synthesis, which resul-

ted in the reduction in the RNA level.

MAHENDRU (1981) suggested that the

anti-AChE compounds attack many

enzymes responsible for normal meta-

bolism pathway. Thus it is possible that

lattices of both the plants might have

inhibited the enzymes necessary for

DNA synthesis, because the lattices of

both the plant tested in the present

study have potent anti-ACHE activity

(SINGH AND SINGH, 2002).

BIBLIOGRAPHY

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BANERJI, A. AND BANERJI J., 1977. Isolation of

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1972) and also involved in the synthesis

of certain enzymes (SUMNER, 1965).

CONCLUSIONS

We therefore believe that these plant

extracts may eventually be of great

value for the control of aquatic target

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mosquito larvae.

ACKNOWLEDGEMENTS

One of the authors (Sunil Kumar

Singh) is thankful to Department of

Environment, (Ministry of Environment

and Forest) Govt. of India (Sanction No.

F-14/35/96 MAB-RE Dated: 09-11-1999),

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Gorakhpur University, Gorakhpur (U.P.),

India.

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plants Thevetia peruviana and Alstonia schola-

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S., 1999. A study on the bioaccumulation of

fenvalerate a synthetic pyrethroid, in the

whole body tissue of Labeo rohita, Catla catla,

Cirrhinus mrigala (Ham.) by gas-liquid chro-

matography. Pollution Research, 18 (1): 57 - 59.

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1977. Effect of thioptera on RNA and total

protein synthesis content in the testis of al-

bino rats. Indian Journal of Experimental Bio-

logy, 15: 804-805.

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sugar concentration in vertebrate to stan-

dard metabolic rate. Comparative Biochemistry

Physiology, 55: 457-460.

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Apigenin 5-methyl ether, ein neues Flavon

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intracellular metabolism. Postepy Higieny 1

Medycyny Doswiadczalnej, 13: 200-206.

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O Sociedad Española de Malacología —__——T— lberus, 21 (2): 29-36, 2003

A sibling species of Gibberula cordorae (de Jong and

Coomans, 1988) in the Leeward Antilles

Una especie gemela de Gibberula cordorae (de Jong and Coomans,

1988) en las Antillas de Sotavento

Franck BOYER*

Recibido el 13-XI-2002. Aceptado el 14-11-2003

ABSTRACT

The species Gibberula cordorae (de Jong and Coomans, 1988) is revised. lts distribution

ranges from the type locality of Curacao up to Saint-Vincent and Saint-lucia. The morpho-

logic variability of its shell is observed as being very low in Saint-Vincent and noticeably

higher in Curacao.

A sibling species, Gibberula colombiana sp. nov., is described from Martinique. lts distrib-

ution ranges up to eastern Guadeloupe. The morphologic variability of its shell is low, and

shows more affinities with the population from Curacao than with the SaintVincent's one.

The present distribution of these different populations is interpreted as resulting from dis-

tinct dispersion stages rather than from a continuous geographic cline.

RESUMEN

Se revisa la especie Gibberula cordorae (de Jong and Coomans, 1988). Su área de distri-

bución incluye desde la localidad típica de Curacao hasta Saint-Vincent y Saint-Lucia. La

variabilidad morfológica de su concha ha sido observada siendo muy baja en Saint-Vin-

cent y evidentemente alta en Curacao.

Una especie gemela, Gibberula colombiana sp. nov., es descrita de Martinique. Su área

de distribución llega hasta el este de Guadaloupe. La variabilidad morfológica de su con-

cha es baja, y muestra más afinidad con la población de Curacao que con la de Saint-

Vincent.

La presente distribución de estas differentes poblaciones es interpretada como el resultado

de distintos estados de dispersión más que como procedente de una variación clinal.

KEY WORDS: Gibberula, Lesser Antilles, sibling species, geographic dispersion, allopatric speciation, variability.

PALABRAS CLAVE: Gibberula, Antillas menores, especies gemelas, dispersión geografica, especiación alopátri-

ca, variabilidad.

INTRODUCTION

The species Gibberula cordorae (de m depth off Curacao. Except for few

Jong and Coomans, 1988) was not additional material sampled in Curacao

recorded in the literature since its by R. Moolenbeek (Zoologisch Museum

description on 19 shells collected at 30 Amsterdam) in the beginning of the

* 110, Chemin du Marais du Souci, 93270 Sevran, France.

Iberus, 21 (2), 2003

heighties, the species remained perfectly

elusive for many years in public and

private collections.

During the second half part of the

nineties, some few additional samples

have been yielded from Curacao by

dutch collectors, and similar morphs

have been discovered from Saint-

Vincent to Guadeloupe by french collec-

tors.

On the ground of these last findings,

J. Colomb (pers. comm.) observed the

constant occurrence of a brown patch on

the columella of the shells from Saint-

Vincent and Saint-Lucia, and the

absence of such a patch on the shells

from Martinique. By the fact of these

divergent features and of contrasted

ranges of length between both sets of

shells, J. Colomb suggested that two

allopatric species might be represented

under the form G. cordorae, the popula-

tion from Martinique constituting a

sibling species new to the science.

Stimulated by such a suggestion, the

revision of G. cordorae is conducted here-

RESULTS

under, and its variability is studied.The

specific identity of the population

ranging in Martinique is verified to be

distinct and to deserve a description as a

new species.

Gibberula cordorae "was initially

described as belonging to the genus Per-

sicula. In fact, both genera are very close.

They have the same kind of soft parts

external anatomy with bifurcated dead,

and their shells principally differ by the

presence of a thick external labial

margin bordered by a noticeable groove

in the species attributed to Persicula

(BOYER, NEEFS AND WAKEFIELD, 1998).

Abbreviations

ad= adult, juv= juvenile, spm= live col-

lected specimen, sh= dead collected

. Shell, splg= sampling.

MNHN Muséum national d'Histoire

naturelle, Paris.

ZMA Zoologisch Museum Amsterdam

FBC F. Boyer collection

JCC J. Colomb collection.

Family CYSTISCIDAE Coovert and Coovert, 1995

Genus Gibberula Swainson, 1840

Type species: Gibberula zonata Swainson, 1840 by monotypy (F=Volvarina oryza Lamark, 1822).

Gibberula cordorae (De Jong and Coomans, 1988) (Figs. 5-9, 10 A-C).

Persicula cordorae De Jong and Coomans, 1999, p. 99, pl. 41, figs. 543 A, B.

Type material: Holotype in ZMA (n” 3.87.093). Non studied.

Other material studied: 14 ad sh (L=5.05 to 6.10 mm), 27 juv and fragments, Saint Kruys Bay, Curacao,

20-30 m, FBC, leg. J. Neefs (Breda, Netherland); 4 ad sh (L=5.70 to 6.30 mm),1 juv, Curacao, 30-60

m, FBC; 1 ad spm (L=5 mm), 3 ad sh (L=5 to 5.30 mm), 3 juv, Curacao, 6-10 m, FBC; 118 ad sh (L=3.85

to 5.15 mm), 14 juv and fragments, Saint-Vincent, 30-40 m, J. Colomb splg 03-01, FBC; 21 ad sh (L=4.08

to 5.41 mm), Saint-Vincent, 30-40 m, J. Colomb splg 03-01, JCC (Figs. 5-8); 8 ad sh (L=4.51 to

5.22mm), Pigeon Island, Roadney Bay, Saint-Lucia, 2 m, J. Colomb splg 03-01, JCC (Fig. 9).

Type locality: Curacao, Santa Martha, at a depth of 30 m.

Original description: “Shell small with

a flat apex. Outer lip thickened and ser-

rated on the inside. Last whorl smooth

and shiny. The colour pattern consists of

alternately 2 prominent and 3 much less

prominent pattern-rows in spiral form.

The 2 prominent rows each consist of 5

to 6 well separated figures. Each figure

is built up of a large brown rectangle

and next to this on both sides a much

narrower brown rectangle. The figures

in the three less prominent rows each

BOYER: A sibling species of Gibberula cordorae in the Leeward Antilles

Figures 1-4. Gibberula colombiana sp. nov. 1, 2: holotype (UNHN), 6.0 x 3.8 mm; 3, 4: paratype 1

(CJC), 6.1 x 4.0 mm. Figures 5-9. Gibberula cordorae (De Jong and Coomans). 5, 6: Saint-Vincent,

40 m, 5.2 x 3.3 mm; 7, 8: Saint-Vincent, 40 m, 5.2 x 3.1mm; 9: Saint-Lucia, 10 m, 5.1 x 3.5 mm.

Figuras 1-4. Gibberula colombiana sp. nov. 1, 2: holotype (MNAN), 6,0 x 3,8 mm; 3, 4: paratype 1

(C]JC), 6,1 x 40 mm. Figuras 5-9. Gibberula cordorae (De Jong and Coomans). 5, 6: Saínt-Vincent,

40 m, 5,2 x 3,3 mm,;7, 8: Saint-Vincent, 40 m, 5,2 x 3,1Imm; 9: Saint-Lucía, 10m, 5,1 x 3,5 mm.

Iberus, 21 (2), 2003

count about 10 white squares and many

brown squares, giving a pattern differ-

ent from that of the prominent rows.

The background is greyish”.

Complementary description: On the

ground of the specimens from Curacao

sampled in 20-30 m and 30-60 m, con-

sidered as topotypes and showing the

same outline than the figured holotype

(DE JONG AND COOMANS, 1988: figs. 543

AandB), further details can be added.

The apex can be more or less flat and

wide, excavated or bulging, but gener-

ally it is submerged by enamel callus. In

some cases, the suture of the last whorl

makes a sharp elevation around the flat-

tened spire and the top of the shell

forms like a wide and shallow crater.

Rarely, the coiling of the first whorls is

suggested or partially visible. The place

of the protoconch can be suggested

where a more or less pronounced pit

occurs on the flattened apex (Fig. 10 A).

A thin varix-shaped fold borders the

edge of the labrum, enlarging progres-

sively towards the base, and making a

distinct oblique callus at the base of the

ventral side, after bypassing the deep

siphonal notch. About 30 very small

denticles are visible on the inner lip. The

4 anterior columellar plaits are strong

and arched (the first one being the

larger), and followed by about 5 col-

umellar lirations. The left basal keel is

generally shorter than the right one,

rarely reaching the same level.

The ground decoration consists of

golden to chestnut-orange oval ocelles

on a creamy-beige background (better

than greyish as reported for the holo-

type) and crossed by 2 prominent and 3

fainter spiral rows of large chestnut-

orange squares or rectangles with dark

borders and separated by white inter-

vals. The ground decoration of ocelles is

often disintegrated in irregular dashes

or spots, like it is visible in the holotype.

Among the 18 adult shells consid-

ered as topotypes, all but one show a

large dark brown patch covering most

of the external columella, from the level

of the second plait to the third posterior

quarter of the aperture. The develop-

ment of this patch occurs in the

SI2

subadult stage. This specific character is

not represented in the holotype.

The shell outline is oval subpyri-

form. Animal unknown.

Measurements: Holotype: 6.5 x 3.7

mm (ZMA). Topotypes: L=5.05 to 6.30

mm (FBC).

Distribution: The species is known

from Curacao, Saint-Vincent and Saint-

Lucia, mostly from 20 to 40 m.

J. Colomb remarked however (pers.

comm.) that the material from Saint-

Lucia consists of few dead shells from

one sampling at only 2 m deep, within

an anchorage where an artificial dyke

was strengthened by the supply of

sands brought from another place. Due

to the facts that he did not find the

species in other places along the

western coast of Saint-Lucia, and that

other “southern antillean species” (like

Conus aurantius Hwass, 1792) were also

found in dead conditions in the sur-

rounding of the same artificial dyke, J.

Colomb suggested that the lot of G. cor-

dorae from Saint-Lucia may have been

transported here by human industry.

The gap in the geographic distribu-

tion between Curacao and Saint-Vincent

is probably due to the lack of adequate

sampling efforts at appropriate depths

in this area.

Remarks: The morphology of the

shell and its decoration do not show any

remarkable variation along the distribu-

tion range of the species.

The population from Saint-Vincent is

very homogeneous and shows faint dif-

ferences with the topotypes from

Curacao: the shells from Saint-Vincent

present a smaller average size, a more

rounded shell, the apex being never

fully crater-excavated with a central pit

(Fig. 10 C), nor really bulging. The left

basal keel is always shorter than the

right one. The ground colour is honey to

beige, decorated of large oval ocelles at

both tips, which tend to disintegrate in a

flecked pattern at the median part of the

shell. The dark ventral patch is always

present in adult and subadult from

Saint-Vincent, even in very worn state.

The shells found in Saint-Lucia do

not show any difference with the shells

BOYER: A sibling species of Gzbberula cordorae in the Leeward Antilles

from Saint-Vincent, except for the pat-

tern of oval ocelles which do not tend to

disintegrate in the small lot at hand.

It must be underlined that several of

the features discussed here are very

subtile, and their comparative study

requires homogeneous conditions of

observation and preservation. For

instance, when ventral views of the

shells are not perfectly taken in the plan,

and when the aperture is slightly turned

towards the left whereas the base is

slightly carried towards the backside (as

in Fig. 9), so the labrum seems to be

thicker, the top wider and the left keel

longer than in reality.

The topotypes from Curacao show a

higher variability in morphology, pro-

portions and decoration, but the small

lot coming from shallow water (6-10 m,

4 ad + 3 juv) show the most divergent

features. The four adult shells have a

slender subcylindric outline, the top is

narrow, the top of the outer lip shows a

clear tendency to encompass the apex

(Fig. 10 B), the basal keels tend to reach

the same level, and only one of the 4

adult shells shows the ventral brown

patch. Their ground bottom is greyish

rather than creamy-beige, and the

pattern of small ground ocelles is very

ill defined.

Gibberula colombiana sp. nov. (Figs. 1-4, 10 D).

Type material: Holotype (Figs. 1, 2) in MNHN; 6 paratypes (Figs. 3, 4) (ad spm) in JCC; 4 paratypes

(1 ad spm, 1 subad spm and 2 juv spm) in FBC; all from the type locality.

Other material studied: 6 ad sh (L=5 to 5.6 mm), Anse d'Arlet, Martinique, 30 m, F. Boyer and R.

Delannoye splg 05-97, FBC; 1 ad sh (L=5.8 mm), 1 juv and 1 fragment, Cap Salomon, Martinique,

22-25 m, P. Clovel splg 28-06-97, FBC; 6 ad sh (L=5.4 to 6.2 mm), 6 juv, Cap Salomon, Martinique,

22-25 m, P. Clovel splg 12-08-97, FBC; 4 ad sh (L=4.9 to 5.7 mm), 5 juv, Cap Salomon, Martinique,

27 m, P. Clovel splg 29-08-97, FBC; 3 ad sh (L=5.4 to 5.9 mm), 1 juv, Cap Salomon, 30 m, P. Clovel

splg 4-09-97, FBC; 1 ad spm (5.9 mm), 4 ad sh (L=5.15 to 5.40 mm), 43 juv, Cap Salomon, Martinique,

22-30 m, P. Clovel splg 6-09-97, FBC; 1 ad spm (L=5.6 mm), 4 juv, Anse d'Arlet, Martinique, 30 m,

P. Clovel splg 10-98, FBC; 47 ad sh (L=5.42 to 6.27 mm), Grande Anse, Martinique, 20-25 m, J.

Colomb splg 2001, JCC; 3 ad sh (L=5.35 to 5.70 mm),1 juv, Grande Anse, Martinique, 20-25 m, Je

Colomb splg 2001, FBC; 1 ad sh, Guadeloupe, east coast, 15m, G. Paulmier splg and coll.

Type locality: Grande Anse, southwestern coast of Martinique, 20-25 m.

Etymology: From the name of Jacques Colomb (Marseilles, France), who first attracted the author's

attention on the specificity of the population of G. cf. cordorae represented in Martinique.

Description: Shell medium-sized for

the genus, outline oval subcylindric. The

top is flat, wide and excavated, the su-

ture of the last whorl making a sharp ele-

vation around the spire, which partially

reveals the coiling of the first whorls,

slightly merged by enamel callus. The

protoconch is suggested by a central pit.

The shoulder of the labrum is sharp

and slightly elevated behind the top, the

labrum is arched and thickened, and

bears numerous tiny denticles almost

obsolete on the upper part of the inner

lip. A thin varix-shaped fold is border-

ing the edge of the labrum, enlarging

progressively towards the base, encom-

passing the siphonal notch and making

an oblique callus at the base of the col-

umella. The 4 anterior columellar plaits

are strong and arched, and followed by

5 columellar lirations. The left basal keel

is much longer than the right one.

The ground decoration is made of

orange-grey oval ocelles on a creamy-

grey back, crossed by 2 prominent spiral

rows of large honey-brown rectangles

with dark borders and separated by

white intervals. 3 fainter spiral rows of

creamy-grey rectangles separated by

white intervals are distributed on either

sides of the 2 prominent spiral rows.

Some of the ground ocelles are partially

obsolete or divided in small spots and

flecks. Any colour patch is not present

on the columella.

Animal (from photos and drawings

made by R Delannoye and J. Colomb):

the wide foot is hyalinous covered of

Iberus, 21 (2), 2003

milky white large lateral patches (some-

times creamy white or light yellowish

towards the posterior part) and by

many smaller irregular spots more

densely grouped on the front of the

propodium and on the back axis of the

metapodium. Few orange dots are scat-

tered on the hyalinous ground and more

densely grouped on the back axis of the

metapodium. The bifurcate head and

the siphon are milky white, with some

orange dots around the eyes. The long

hyalinous tentacles are decorated by

whitish dashes and dots, and by scat-

tered orange dots. The eyes are black.

Measurements: Holotype: 6 x 3.8 mm

(MNHN). Paratypes 1 to 6, L=5.86 to

6.23 mm (JCC). Paratype 7 (ad.sh), L=5.9

mm, paratypes 8 to 10 (subad and

juv.sh), L=4.9 to 5.5 mm (FBC).

Distribution: The species is known

from Martinique by many individuals

and from Guadeloupe by only one shell.

This last one is absolutely similar to the

material from southwest Martinique,

and the datum given by G. Paulmier is

confirmed as right. So the species must

be ranging at least from Martinique to

Guadeloupe, the extension of the distri-

bution towards northern Lesser Antilles

remaining possible on the ground of

new discoveries.

POSTS ,

”

4 y

( .

( e

SSA

Figures 10 A-C. Gibberula cordorae (De Jong and Coomans). A: Curacao, 30-60 m; B: Curacao, 6-

10 m; C: Saint-Vincent, 40 m. Figure 10 D. Gibberula colombiana sp.nov., Martinique, 20-25 m.

Figuras 10 A-C. Gibberula cordorae (De Jong and Coomans). A: Curacao, 30-60 m; B: Curacao, 6-

10 m; C: Saint-Vincent, 40 m. Figure 10 D. Gibberula colombiana sp.n0v., Martinique, 20-25 m.

Remarks: G. colombiana sp. nov shows

very homogeneous shell morphology

and decoration, and presents close simi-

larity with G. cordorae. Most of the fea-

tures represented in G. colombiana belong

to the range of variability represented in

the populations of G. cordorae from

Curacao. The large size and the exca-

vated top of the shell of G. colombiana

(Fig. 10 D) match the features occurring

in several topotypes of G. cordorae (Fig.

10 A). The subcylindric outline and the

greyish ground of decoration found in G.

colombiana match the features of the

shells of G. cordorae studied from shallow

water in Curacao (6-10 m).

However, two special characters can

be considered as distinctive features of

G. colombiana: the longer left basal keel

and the total lacking of a ventral brown

patch. The first character is especially

original, as such a produced keel is very

uncommon within the genus. The

second character is significant from a

statistic point of view, as the ventral

patch is represented as highly dominant

in G. cordorae, lacking only in few shells

from Curacao and never lack in the

shells from Saint-Vincent and Saint-

Lucia. The presence of a ventral patch

can be considered as a specific feature of

G. cordorae, accidentally absent (or much

BOYER: A sibling species of Gibberula cordorae in the Leeward Antilles

Hispaniola

Aruba, Curacao

Figure 11. Map of the Eastern Caribbean Sea.

Figura 11. Mapa del Este del Mar Caribe.

obsolete) in some shells, whereas the

total lack of ventral patch in G. colom-

biana can be interpreted as a proper spe-

cific feature.

On the other hand, it must be under-

lined that the very homogeneous shell

features of G. colombiana show many

similarities with some of the forms of G.

cordorae ranging in Curacao, but much

DISCUSSION

On the ground of the phenotypic

variability patterns observed within the

populations of G. cordorae and of G.

colombiana, the adjacent distribution of

both species cannot be interpreted as

resulting from a step by step settling

leading to a continuous geographic

cline, which is the most current situation

within the non-planktotrophic groups

such as cystiscids.

It seems more likely, in the present

case, that the “G. cordorae group” has

been subject to several periods of dis-

persion associated to geologic and

hydroclimatic events along the per-

Puerto yin:

Rico Virgin Islands )

Bonaire

Anguilla

St. Martin

Y eo mo e _ St. Barthelemy

St. Thomas $7 se, Barbuda

| e Antigua

Sab / rr

. 2 Saba

St. Croix St. Eustatius l/ Ñ ge edo:

St. Kitts E

Nevis Y

Y Martinique

St. Lucia Q Barbados

St. Vicent 9

Grenada Y

Margarita e Tobago

= iio

Tortuga

less with the populations from Saint-

Vincent and Saint-Lucia which range in

a closer vicinity. In these conditions, the

neighbouring populations from Mar-

tinique on one hand and from Saint-

Vincent and Saint-Lucia on the other

hand show as well-contrasted morphs,

here interpreted as representing two

allopatric sibling species.

turbed history of the eastern antillean

region (FAIRBRIDGE, 1966). According to

this view, G. cordorae may represent a

“southern stock” belonging to the

Venezuelan area, and G. colombiana may

represent an “eastern stock” coming

from the old inner arc of the Limestone

Caribbes.

The G. cordorae group has apparently

not any other representant in the

Caribbean Province, but a close relative

is represented in the Panamic Province

as G. phrygia (Sowerby, 1846), which

presents a very constant pattern from

Western Mexico to Galapagos Islands.

Iberus, 21 (2), 2003

ACKNOWLEDGEMENTS

Il am indebted to Regis Delannoye

(Trinité, Martinique), to Pierre Clovel (Fort-

de-France, Martinique) and to Jacques

Colomb (Marseilles, France) for their ac-

tive contribution to this study, for their

sampling efforts and precious observa-

BIBLIOGRAPHY

BOYER, F., NEEFS, J]. AND WAKEFIELD, A., 1998.

A new species of Persicula from the Venezue-

lan region (Gastropoda: Cystiscidae). Journal

of Conchology, 36 (4): 15-20.

COOVERT, G. A. AND COOVERT, H. K., 1995. Re-

vision of the Supraspecific Classification of

Marginelliform Gastropods. The Nautilus,

109 (2-3): 43-110.

DE JONG, K. M. AND COOMANS, H. E., 1988. Ma-

rine Gastropods from Curacao, Aruba and

Bonaire. E. J. Brill, Leiden, 261 pp.

tions as well as for the gift or the loan of

material. Gérard Paulmier (Monteaux,

France) also loaned useful material.

A special thanks is due to Tiziano

Cossignani (Ancona, Italy) who per-

formed the photos and the digital plates,

and to Rafael La Perna (Catania, Italy)

for the helpful correction of the text.

FAIRBRIDGE, R. W., 1966. The Encyclopedia of

Oceanography. Reinhold Publishing Corp.,

New York, 779 pp.

SOWERBY, G. B., IL, 1846. Monograph of the

genus Marginella, in Thesaurus Conchyliorum,

or figures and descriptions of Recent Shells, 1 (7):

239-406, pls. 68-78. Sowerby, London.

O Sociedad Española de Malacología Iberus, 21 (2): 37-60, 2003

Contributions to West-Mediterranean Solenogastres

(Mollusca) with three new species

Contribuciones al conocimiento de los Solenogastres (Mollusca) del

Mediterráneo occidental, con tres nuevas especies

Luitfried v. SALVINIEPLAWEN*

Recibido el 1-X-2002. Aceptado el 13-111-2003

ABSTRACT

Six species of West-Mediterranean Solenogastres are treated, three of which are

described as new to science: Micromenia subrubra n.sp. from off Malta, Macellomenia

adenota n.sp. from off Ceuta, Urgorria monoplicata n.sp. from off the Costa Brava.

Records of other species (Nematomenia banyulensis, Tegulaherpia sp. and Eleutheromenia

sierra) contribute to our knowledge of respective geographical distribution. The refinding

of E. sierra also results in the definition of a new genus Scheltemaia n.g. for two South-

Australian, former Eleutheromenia species.

RESUMEN

Se estudian seis especies de Solenogastres del Mediterráneo occidental, incluyendo tres

nuevas especies: Micromenia subrubra n.sp. de Malta, Macellomenia adenota n.sp. de

Ceuta, Urgorria monoplicata n.sp. de la Costa Brava. Se aportan informaciones sobre

otras especies, Nematomenia banyulensis, Tegulaherpia sp. y Eleutheromenia sierra, que

contribuyen a un mejor conocimiento de sus distribuciones. La redescripción de E. sierra

justifica la definición de un nuevo género Scheltemaia n.g. para dos especies descritas

del Estrecho de Bass, Australia meridional.

KEY WORDS: Mollusca, Solenogastres, geographical distribution, new species.

PALABRAS CLAVE: Mollusca, Solenogastres, distribución geográfica, nuevas especies.

INTRODUCTION

The Solenogastres are a small class of

aplacophoran Mollusca, characterised by

the mantle cover of cuticle and unicellu-

larily formed, calcareous mantle bodies

. Or sclerites (scales, spicules, a.o.) and by

the laterally rounded body with the foot

narrowed to a longitudinal pedal groove.

With respect to the mantle cover, to char-

acters of the alimentary tract (no true

radula ribbon, midgut without separate

digestive gland) and to the lack of partic-

ular excretory organs (no emunctoria),

they appear to represent a very conserva-

tive off-shoot of Mollusca. Apart from the

plesiomorphic, aplacophoran mantle and

a few other features, they are by diphyly

also clearly different from the Caudo-

foveata (cf. SALVINI-PLAWEN, 2003).

Solenogastres are generally small

(average 3 mm - 3 cm, rarely longer, up

* Institut fir Zoologie, Universitat Wien, A-1090 Wien (Vienna/Austria), Althanstrasse 14

Iberus, 21 (2), 2003

to 30 cm), and about 230 species are

nominally known at present. Their

purely marine habitats (mostly below 50

m depth), their organisation based

mainly on internal systematic charac-

ters, together with the great effort and

expense required to collect them have

clearly prevented broader assessments

and a greater familiarity with this

group. Much material from more recent

collections (expeditions and individual

samplings) awaits elaboration and will

no doubt contribute to a still fairly frag-

mentary knowledge. This is also valid

for basic faunistics (biodiversity, bio-

geography, etc.). Due to such incom-

pleteness, all information - even if frag-

SYSTEMATICS

mentary - is of interest. In regard to the

revised classification (SALVINI-PLAWEN

1978), systematic problems remain;

these also concern several descriptions

that are fragmentary compared to the

required standard and await supple-

mentation or revision.

The present contribution compiles

Solenogastres from different samplings

of the western Mediterranean Sea. Apart

from enlarging our information on

organisational diversity and providing

geographical insight, this study also

improves our understanding of the bio-

diversity of Solenogastres even within a

fairly well investigated biogeographical

region (cf. SALVINI-PLAWEN 1986, 1990).

Ordo PHOLIDOSKEPIA Salvini-Plawen, 1978

Solenogastres with a single layer

of adpressed and overlapping solid,

scaly mantle sclerites, sometimes

supplemented by other solid scle-

rites; cuticle thin, no epidermal papi-

llae.

Family DONDERSIDAE Simroth, 1893

The family is characterised by two

different types of mantle scales, by a

monoserial radula provided with

paired, long denticles, by the foregut

glandular organs as subepithelial folli-

cles (so-called type A in SALVINI-

PLAWEN, 1972, 1978), as well as by the

lack of respiratory organs. It includes

seven genera (cf. SALVINI-PLAWEN

1978):

Genus Nematomenia Simroth, 1893

Type species: Dondersia flavens Pruvot, 1890.

Dondersia Hubrecht, 1888, partim; Myzomenia Simroth, 1893; Herpomenia Heath, 1911.

Definition: Solenogastres-Dondersi-

idae with common atrio-buccal

opening; radula rudimentary: either

monoserial radula with two pairs of

distally touching denticles (bifid-mono-

stichous type), or radula sheath without

radula, in part forming the unpaired

outlet for the paired foregut glandular

organs; midgut without serial constric-

tions; terminal sense organ present; sec-

ondary genital opening single, no copu-

latory stylets.

Nematomenia banyulensis (Pruvot, 1890)

Dondersia banyulensis Pruvot; Myzomenia banyulensis (Pruvot) in Simroth 1893;

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

Diagnosis: Body distinctly red, elon-

gate and slender, up to 30 mm, middor-

sal scales forming a longitudianl crest;

main type of scales short and wide, dis-

tally pointed, proximally in part

notched; mid-foregut forming a protrusi-

ble cone, without radula, radula sheath

forming the single outlet of paired glan-

dular organs and, by elongation, open-

ing at the tip of the cone, posterior

foregut curving anteriorly to open pre-

cerebrally into frontal midgut; no papil-

lated dorsal foregut gland; musculus

longitudinalis distinctly separated.

Spawning ducts fused throughout, axi-

ally with one pair of receptacula seminis.

Remarks: This well-known species

(up to 30 mm in length) is recorded

along the European coasts with wide

gaps from off Dalmatia/Croatia to the

Trondheimsfjord/Norway (PRUVOT

1891, NIERSTRASZ AND STORK 1940,

SALVIN-PLAWEN 1997, HANDL AND

SALVINI-PLAWEN 2001). There is a recent

finding of several 3-5 mm long, red spec-

imens by C. Mifsud (26.8.2002) from off

Ras il-Wahx/NW-Malta (ca.35%57N,

1419E) at 120-140 m, and another

record (25.6.1994) of two small individu-

als comes from the campaign FAUNA

IBERICA MI (PB 923-0121), station 186 A,

from off Cabo Cros in front of Punta na

Foradada /NE-Mallorca (39 47.64'-39"

49.66' N, 2' 40.78'-2" 38.71” E) at 59-61 m

from a bottom also characterised by the

presence of Hydrozoa-Sertulariidae. This

latter record interbridges the known oc-

currence from the Gulf of Naples and the

Cóte Vermeille (Banyuls).

One of the Malta animals (3.6 mm)

and the larger individual of the Mal-

lorca specimens (preserved only 3.2 mm

long, 0.5 mm high and 0.4 mm wide)

were serially cross sectioned (semithin

ribbons of 2 um with glassknife); they

show all typical characters of N. banyu-

lensis (particularly curved foregut with

cone, etc.). Although the Mallorca

animal (deposited in the Mus. Nacional

Cienc. Nat., Madrid, mol. no. 15.02/15)

contained eggs of (“W 80-100 ym and

sperm, the receptacula seminis were not

yet differentiated. As PRUVOT (1891: 716)

indicates that young individuals are

pale reddish or orange, the distinct red

colour of both the small Mallorca

animals (maximum size of only 3.5 mm)

is Of interest.

Genus Micromenia Leloup, 1948

Rupertomenia Schwabl, 1955

Type species: Micromenia simplex Leloup, 1848.

Definition: Solenogastres-Dondersi-

idae with atrial sense organ (vestibu-

lum) and mouth separate; monoserial

radula with one pair of lateral, curved

denticles touching each other distally

(bifid-monostichous type); midgut

without serial constrictions; terminal

sense organ (s) present; secondary

genital opening single, no copulatory

stylets.

Micromenia subrubra spec. nov.

Holotype: NW-Malta, off Ras il-Wahx, 35%57'N, 1419E, yellow mud and sand at 140 m (Sept.

1996); spicules and section series (cs 2 qm) on slides: Naturhist. Mus. Wien (NHMW), Moll. IVN

1002446.

Paratype 1: NW-Malta, off Ras il-Pellegrin, about 35%55'N, 14*19'E, 120 m (April 1994); spicules and

section series (cs 2 jm) on slides: Nat. Hist. Mus. London, Rg.no. 20020531.

Paratype 2: NW-Malta, off Delli, grey mud at 140 m (June 1999); whole animal, 5 mm long (Fig.

10), preserved in ethanol: Naturhist. Mus. Wien (NHMW), Moll. IVN 102447.

Derivatio nominis: Latin subrubrus = not very bright red, reddish.

Iberus, 21 (2), 2003

Diagnosis: Body reddish, to 5 mm x

0.5 mm in size (stretched in life up to 7

mm), dorso-posterior body end gener-

ally elongated. Calcareous mantle scle-

rites of two types: slender shovel-like

scales and slightly curved solid needles.

Anteriorly concentrated subepithelial

glands opening into dorsal pharyngeal

pouch; paired ventral foregut glandular

organs with single opening; midgut

with short rostral caecum. With one pair

of receptacula seminis and vesiculae

seminales. Mantle cavity extending as a

ventrally open gutter onto the elongated

dorso-posterior body portion, ventro-

rostrally with pouch; 2-4 dorsoterminal

sense organs (Figs 1-4, 10, 12).

Material and methods: Several speci-

mens had been collected by Constantine

Mifsud (Malta) in April June /July 1994,

June 1995, August, Sept. and Nov. 1996,

June 1999 and May 2001 from muddy

bottoms off Western Malta in the area

off Ras il-Pellegrin (between the Gnejna

and Fomm ir-Rih bays), about 35%5'N,

14919E, at 120-160 m (see MIFSUD, 1996),

and off cape Ras il-Wahx, about

35%57'N, 1419, at 140 m.

Ribbons of semithin serial sections

(cs 2 qm) of several specimens have

been made with glassknifes and stained

with Richardson's solution.

External appearance: The slender

animals are generally 2.5-5 mm in length

(Fig. 10), but when stretched in life may

even reach a length of up to 7 mm. They

are fairly susceptible to mechanical effects

and easily damaged through sampling

treatment. The animals exhibit in life a dis-

tinct, not very intensive reddish colour

throughout, ranging from pink to red (see

Fig. 4 in MIrsuD 1996). The hind end in

adults shows a beak-like extension of the

dorso-posterior body (Fig. 10); in smaller

(juvenile or sub-mature) individuals (up

to 3 mm in length) the posterior body in

life may be swollen and appear to be still

devoid of the dorso-posterior body elon-

gation. The mantle sclerites are generally

adjoining and give the body surface a

somewhat shiny appearance; the mid-

dorsal scales may form a low crest.

Mantle: Cuticle 7-15 qm. Calcareous

mantle sclerites of two types (Fig. 1): a)

very slender shovel- to trowel-like

scales 55-110 pm long (handle 12-20 qm

long) and 7-15 ym wide; b) slightly

curved solid needles with indistinct

handle, 80-110 qm long x 4-6 qm wide;

both types may extend beyond the

general body contour.

Foot and mantle cavity: Pedal ciliary

pit flat and wide (125 um), densely cili-

ated, without folds or ridges. Foot

formed by posterior narrowing of the cil-

iated epithelium, changing along its

course between a flattened ridge (25 qm

wide) with centrally high cells and a real

fold. It does not enter the mantle cavity.

Pedal gland voluminous with large

follicles at both sides in the anterior

body. Sole glands along pedal groove

small, scattered, of same histological

quality as pedal glands.

Mantle cavity as an elongated, ven-

trally open gutter extending along entire

dorso-posterior beak-like body portion,

densely ciliated without respiratory for-

mations. At the beginning of this beak-

like portion the ventroterminal body

wall including “abdominal” spicules is

curved dorso-anteriorly to form a wide

dorsally open gutter facing the gutter of

the mantle “cavity”. This entire internal

space is rostrally three-partitioned (Fig.

4): dorsal epithelium closing up as rectal

opening (anus); central portion closing

up to represent the short outlet of the vo-

luminous spawning duct; the ventral

portion becoming a rostral, wide and

flattened pouch which - at least dorsally

- appears to be lined by mantle epithe-

lium (with cuticle-like substance and

embedded small sclerites). More

frontally this pouch either (holotype)

continues ventrally in a fairly circular

blind sack (Figs. 4, 12), or shows a mid-

ventral ridge, implying distinct symme-

try of the wide pouch; the latter condi-

tion (paratype 1) extends anteriorly until

just below the end of the pericardium.

Musculature: Body wall musculature

distinct but not very compact, its fibres

showing a loose arrangement enclosing

lacunae. No lateroventral reinforcement

of longitudinal fibres. The regular

dorsoventral bundles are weak and run

peripherically, only occasionally causing

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

30 um

100 um

Figures 1-4. Micromenia subrubra spec.nov. 1: mantle sclerites; 2: organisation of the anterior

body; 3: one radula tooth/plate; 4: organisation of the posterior body. Abbreviations, as: atrial

sense organ; ce: cerebral ganglion; co: suprarectal commissure; du: outlet of ventral foregut glandu-

lar organs (type A); fo: pedal fold (foot); gl: follicles of glands; go: gonad; hg: hindgut (rectum);

mc: mantle cavity; mg: midgut; mo: mouth opening; pc: pericardium; pd: pericardioduct; ph:

pharynx; pi: pedal pit; rs: radula sheath; sd: spawning duct; so: terminal sense organ; sr: receptacu-

lum seminis; vg: (first) ventral ganglion; vs: vesicula seminalis.

Figuras 1-4. Micromenia subrubra spec.nov. 1: Escleritas del manto; 2: organización de la parte ante-

rior del cuerpo; 3: diente radular; 4: organización de la parte posterior del cuerpo. Abreviaturas, as:

órgano sensorial atrial; ce: ganglio cerebral; co: comisura suprarrectal; du: abertura de los órganos diges-

tivos ventrales anteriores (tipo A); fo: pie; gl: grupos de glándulas; go: gónada; hg: recto; mc: cavidad del

manto; mg: tubo digestivo; mo: abertura bucal; pc: pericardio; pd: pericardioducto; ph: faringe; pi:

foseta pedal; rs: rádula; sd: conducto de la puesta; so: organo sensorial terminal; sr: receptáculo seminal;

vg: (primer) ganglio ventral; vs: vesícula seminal.

Iberus, 21 (2), 2003

constrictions of the midgut. At the begin-

ning of the pericardium, the inner/up-

per pair of dorsoventral bundles fuses

with the peri-pericardial musculature.

Sensory system: Cerebral ganglion

(110-125 qm wide, 75-80 qm high, 70 qm

long) with separate connectives, the

buccal ones first and very close to the

ventral connectives. Four laterofrontal

pairs of medullary roots of cerebral

nerves (holotype), a ventro-lateral, a

lateral and two subsequent dorsolateral

ones; the lateral and dorsolateral roots

connect at each side to an adjacent basal

ganglion; this latter gives rise to only

two (rather than three) nerves which

lead together with the separate ventro-

lateral cerebral nerve to the atrial region.

All these (six) nerves are medullary.

Lateral ganglia elongate (55 qm x Y

30 pm) and adjacent to the cerebral

ganglia. Ventral ganglia voluminous (Y

60 x 45 qm), interconnected by two com-

missures, followed by another pair of

ventral ganglia with their commissure,

all posterior to the ciliated pedal pit

(Fig. 2). Buccal ganglia (Y up to 45 x 25

pm) posterio-lateral of the radula sheath

between the follicles of the foregut glan-

dular organs, interconnected by an

exceptionally long commissure dor-

soterminally of the sheath (150 um

when relaxed in the here 300 um wide

body, 60 pm when contracted and

curved).

Suprarectal commissure overlying

rectum just below end of the peri-

cardium (Fig. 4). The lateral cords con-

tinue medullarly into the dorsoposterior

body extension and form two additional

thick commissures.

Atrial sense organ clearly separated

from mouth. A distinct pre-atrial, cili-

ated pit is formed which leads into the

voluminous atrium. Atrial ciliary tract

ruming at the periphery of the opening

only, frontally continuous with the pre-

atrial pit. No dorsal ciliation. Cavity

itself with several stoutish papillae,

most being elaborated as pairs.

There are two (holotype) to four

(paratype 1) subsequent dorso-terminal

sense organs at the posterior elongation

of the body (Fig. 4).

Alimentary tract: Mouth opening

wide, separate from the atrium, leading

into a voluminous (or but longitudinally

folded) foregut with a dorsal enlarge-

ment below the cerebral ganglion. This

pouch receives frontally subepithelial

follicles of glandular cells (Fig. 2)

without forming a papilla. Behind

pouch, foregut with some distinct circu-

lar musculature.

Radula (Fig. 3) consists of monoser-

ial teeth or plates (20 ym wide), each

with a curved denticle laterally (up to 20

pm long), both which are very close

medially and touch distally. Ventral

foregut glandular organs present as

wide ducts or ampullae (Fig. 2) with

large subepithelial follicles filling the

space to the midgut; wide ducts fuse to

open by a single outlet ventral of the

anterior radula.

Foregut in radular region with

dorsal and lateral coating of small

subepithelial foregut glands adjacent to

strengthened circular musculature.

Postradular foregut opening axially

without sphincter into the (surround-

ing) midgut. Frontal midgut caecum

short. Midgut, depending on the indi-

vidual, at a more or less distinct dis-

tance from the body wall. Regionally

showing irregular bulges, but no true

serial constrictions due to the dorsoven-

tral musculature and not forming

regular lateroventral pouches. Middor-

sal ciliary tract present. In two speci-

mens small nematocysts were embed-

ded in the midgut epithelium.

Circulatory system: Atrium of heart

forming a spacious invagination of the

pericardial roof, being free in its anteri-

ormost portion only and interconnected

with the ventricle by one distinct

opening. Ventricle for half or two-thirds

its entire extension as an invagination of

anterior pericardium.

Body spaces (pseudocoel) between

the organs in part filled by distinct mes-

enchyme as well as blood cells. The

latter consisting of round cells (U 2-5

pm) with in part densely granulated

contents and well visible nucleus, and of

more irregular, vacuolised cells (Y 3-7

pm).

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

Gonopericardial system: The two

gonads distincly separated throughout,

containing eggs of at most 40 ym in

diameter. Terminal portion of the gonads

in fully mature animals serves as a volu-

minous sperm bag, then being ventrally

continuous with the two short and wide

gonopericardioducts, which are ciliated

throughout. Ducts open dorsally into the

pericardium, ciliation continuing as a

paired, wide lateral tract throughout

pericardium and entering pericardiod-

ucts. Pericardioducts emerging termino-

laterally, each elaborating in its subse-

quent curve towards anterior an elongate

and wound, anteriorly directed vesicula

seminis (Fig. 4). Ciliated pericardioducts

open rostrally from laterodorsal into the

respective spawning duct.

Both spawning ducts connected ante-

riorly by a curved duct (stalk) with a

voluminous receptaculum seminis (Fig.

4). After a short length, both glandular

spawning ducts open anterio-laterally

into a voluminous continuation showing

a rostro-median pouch between the

openings and being provided with a dis-

tinctly lower, ciliated epithelium. This

subsequently single spawning duct soon

widens laterally and becomes dorsally as

well as ventrally lined by a high glandu-

lar epithelium and provided with some

musculature. It opens by means of a

short and simply-lined, central outlet,

surrounded by musculature, into the

“mantle cavity” (see above).

Discussion: It should be mentioned

that two sub-mature animals showed an

atrial sense organ without papillae, and

it cannot be evaluated whether this

refers to the sub-mature status or

reflects a teratological condition.

The characteristic radula classifies

the present animals within the Donder-

siidae in the closer relationship of Don-

dersia Hubrecht, 1888, itself (see SALVINI-

PLAWEN, 1978) and of Micromenia. As

already pointed out earlier (SALVINI-

PLAWEN, 1972: 218), the genus Microme-

nia differs from Dondersia by the absence

of serial midgut constrictions due to the

peripherical course of the dorsoventral

muscle bundles. Such a condition,

however, could perhaps be correlated

with body size and would then not rep-

resent a good generic character. Apart

from this, note the closer similarities

with respect to the shovel- to trowel-like

scales in the present species and in Don-

dersia festiva Hubrecht, 1888, as well as

in Heathia porosa Heath, 1911 (see SCHEL-

TEMA, 1998a). There is also a remarkable

multiplication of dorsoterminal sense

organs in some Dondersia species: two in

D. festiva and D. indica Stork, 1941, three

in D. annulata Nierstrasz, 1902, and

eleven in D. californica Heath, 1911.

With respect to the mentioned elabo-

ration of the midgut, the species in ques-

tion belongs to Micromenia. At present,

two Micromenia species are known, i.e.

M. simplex Leloup and M. fodiens

(SCHWABL, 1955). Besides the known

geographic occurrence off Spitzbergen,

M. simplex differs from the specimens at

hand by several dominant characters,

e.g. the lack of the dorsoposterior body

elongation, the differentiation of scat-

tered pharyngeal glands, the paired

opening of the ventral foregut glandular

organs and the unpaired spawning duct

(LELOUP, 1948; SALVINI-PLAWEN, 1972).

M. fodiens is known from SW Sweden

and from Norway in the area off Bergen

(SALVINI-PLAWEN, 1988); new evidence

comes from the Skagerrak (two localities,

70-80 m) and the Trondheimsfjord

(several localities, 185-530 m) (pers.

comm. C. Handl, Vienna). M. fodiens

differs from the present animals by the

whitish body colour and the lack of the

dorsoposterior body elongation as well

as Of a pedal fold, the shape of the scales,

the rudimentary atrial sense organ

without papillae, and also by details of

the gonopericardial system (SCHWABL

1955, SALVINI-PLAWEN 1972, 1988).

M. subrubra spec.nov. is thus well-

separated and its finding enlarges the

biodiversity of Micromenia by a Mediter-

ranean species. With respect to the body

colour, together with Nematomenia banyu-

lensis (above) and the enigmatic Ne-

matomentia (?) corallophila (KOWALEVSKY,

1881) recorded upon Corallium rubrum

(L.) off La Calle/ Algeria (cf. SALVINI-

PLAWEN 1997), M. subrubra is the third

“red Solenogastre” in the region.

Iberus, 21 (2), 2003

Family MACELLOMENIIDAE Salvini-Plawen, 1978

The family is characterised by the

nail-shaped type of mantle sclerites,

by a monoserial radula of serrate

plates, by the foregut glandular

organs as subepithelial follicles (so-

called type A in SALVINI-PLAWEN,

1972, 1978), as well as by the presence

of respiratory organs; at present it

includes one genus only (cf. SALVINI-

PLAWEN, 1978).

Genus Macellomenia Simroth, 1893

Type species: Paramenia palifera Pruvot, 1890.

Paramenia Pruvot, 1890, partim [non Brauer and Bergenstam, 1889]

Definition: — Solenogastres-Macel-

lomeniidae with solid acicular sclerites

with enlarged basal plate; paired

ventral foregut glandular organs (type

A) with distally single outleading duct;

terminal sense organ present; second-

ary genital opening unpaired; no copu-

latory stylets.

Macellomenia adenota spec. nov.

Material: A single specimen 1.65 mm long and 0.75 mm across was collected in May 1986 from the

Strait of Gibraltar, off Punta Almina (North of Ceuta: 35%54.1'N, 0%16.5'W; 25-40 m). After exami-

nation of the mantle cover, ribbons of semithin serial sections (somewhat oblique cs 2 jm) were

made with a glassknife and stained with haematoxyline-eosine.

Holotype: Mus.Nat.Hist.Nat. (Malacologie), Paris; section series

Derivatio nominis: Greek adén = gland, Latin -otus = provided with; referring to the paired pre-

pallial “abdominal gland” in addition to the foregut glands.

Diagnosis: Body less than 2 mm long;

mantle sclerites with spine slightly

curved and up to 125 qm long, not form-

ing a crest; atrial sense organ (vestibu-

lum) and mouth opening separate;

radula plates curved with seven denti-

cles, the median one most promenent;

midgut with paired rostral caecum, with-

out regular lateral constrictions; with

paired “abdominal gland” and paired ab-

dominal spicules, all opening into a pre-

pallial mantle invagination (Figs 5, 6, 16).

Description: Characteristic solid man-

tle sclerites (Figs. 5a, b) with a 70-100

um long, slightly curved spine and an

oblique basal plate (20-25 qm x 12-15

pm) reinforced at the free proximal rim.

Some sclerites embedded in epidermis

which includes several larger gland

cells. Mantle cuticle thin, about 15 um

(13-20 qm).

Rear of body with mantle cavity not

retained. With pre-pallial ventral mantle

invagination with opening of a paired,

slender pouch, each including a bundle

of at least 12 slender spicules or spines

(abdominal spicula). In addition, a

paired short tube also opens aside the

end of the foot lateral in the groove of

the mantle invagination; it is elaborated

just anterio-dorsal of each spicule

pouch, provided with an epithelium of

well elaborated glandular cells and sur-

rounded by musculature (“abdominal

gland”, Fig. 16).

Pedal pit of foot simple, receiving

the outlets of the pedal gland follicles.

The single pedal fold soon flattens to a

wide ciliated ridge ending with mantle

invagination. Sole glands along groove

present.

Body wall musculature without par-

ticular ventral reinforcement of the lon-

gitudinal layer. Serial dorsoventral

bundles weak, running fairly close to

body wall, not constricting midgut.

Preoral atrial sense organ ending as

blind sack (Fig. 6), with few single,

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

100 um

Figures 5, 6. Macellomenia adenota spec.nov. 5: mantle sclerites from lateral (a) and frontal (b), and

one radula plate; 6: organisation of the anterior body. Abbreviations, as: atrial sense organ; bu:

buccal ganglion; bw: body wall (mantle and musculature); ca: midgut caecum; ce: cerebral gan-

elion; fo: pedal fold (foot); gl: ventral foregut glandular organ; go: gonad; mg: midgut.

Figuras 5, 6. Macellomenia adenota spec.nov. 5: escleritas del manto en vista lateral (a) y frontal (b), y

diente radular; 6: organización de la parte anterior del cuerpo. Abreviaturas, as: órganos sensorial

atrial; bu: ganglio bucal; bw: pared del cuerpo (manto y musculatura); ca: ciego digestivo; ce: ganglio

cerebral; fo: pie; gl: órgano glandular digestivo ventral; go: gónada; mg: tubo digestivo.

stoutish papillae. Alimentary canal with

mouth opening separate from atrium,

leading into a short foregut. Animal

with somewhat protruded radula appa-

ratus, foregut forming a posterior pouch

above radula sheath as well as a flat

blind sack extending rostrally over the

cerebral ganglion. Plates of monoserial

radula only 25 pm wide and curved;

with seven denticles, the median one

distinctly being strongest (Fig. 5). No

particular radula support.

Ventral foregut glandular organs,

each consisting of a densely packed

bunch of subepithelial follicles extending

lateroventrally of the midgut and empty-

ing at each side into short duct; both

ducts uniting from lateral to a median

outlet with circular musculature. Duct

running below radula sheath and then

below free radula to open beneath ante-

riormost plate of protruded radula.

Foregut opening into midgut above

beginning radula, midgut giving rise

from this area to a paired rostral caecum

with high epithelium (30-120 qm) equal

to that of midgut. Midgut with some

nematocysts.

Nervous system with single cerebral

ganglion (110 qm wide, 40 qm high, 35

pm long), connectives of body cords

emerging separately. Buccal ganglia (9

25 um) aside radula sheath (Fig. 6), their

commissure over sheath. Ventral ganglia

(9 65 x 35 pm) adjacent to pedal pit.

Animal mature, paired gonad with

relatively large eggs (Y up to 110 um).

Other organs of gonopericardial system

(pericardium, spawning ducts, etc.) and

mantle cavity not retained.

Iberus, 21 (2), 2003

Discussion: Within Macellomeniidae,

only two species of Macellomenia have

been described so far, viz. M. palifera

(Pruvot, 1890) and M. aciculata Schel-

tema, 1999, both provided with a similar,

remarkable mantle cover of solid acicu-

lar spicules with enlarged basal plate.

The present specimen fits well into that

frame, but exhibits specific differences.

M. aciculata possesses radula plates with

five equal denticles, the mouth opens at

the back of the atrial sense organ and

there is a lack of abdominal spicules

(SCHELTEMA 1999). The present specimen

coincides with the geographically closer

type species M. palifera by the radula

plates with seven denticles and by the

presence of invaginated pouches for ab-

dominal spicules (PrRUVOT, 1891: 790-

791). There are specific differences how-

ever: (a) Curved radula plates with a dis-

tinctly prominent median denticle; (b)

paired midgut caecum (not present in M.

palifera; PRUVOT 1891: 790); (c) the paired

“abdominal gland” (see below) empty-

ing into the mantle invagination close in

front of the abdominal spicules; (d) body

length (though mature, less than 2 mm,

M. palifera = 4-8 mm, Pruvot 1891: 727)

and lack of a middorsal crest of spicules

(Pruvor, 1891: 727 and Fig. 4). Most

probably (e), also in M. palifera the

mouth opening is located within the pre-

oral atrium.

Though the gonopericardial system,

the mantle cavity and the terminal sense

organ of the present specimen are not

known, this animal clearly represents a

proper species, Macellomenia adenota

spec. nov.. An association of abdominal

spicula with a paired gland has been

reported also in Squamatoherpia tricuspi-

data (BUCHINGER AND HANDL, 1996),

though in different mutual arrange-

ment. The histology of the present

“abdominal glands” looks similar to

that of spawning ducts; as no characters

of the pericardium and mantle cavity

are known, the glands could be the

anlagen of those organs. If so, however,

the spawning ducts would represent

fairly unusual, lateral organs with

medial opening into a pre-pallial mantle

invagination. The presence or absence of

a terminal sense organ, as well as the

common (M. aciculata; M. palifera?) or

separated atrial and buccal openings

(M. adenota) are judged to be generic

characters (SALVINI-PLAWEN, 1967b,

1978); due to the currently limited

insight in species organisation and

species diversity of Macellomeniidae,

however, no reclassification is proposed

herein.

No accurate evaluation is possible

with regard to the Macellomenia speci-

mens reported by Caudwell et al. (1995)

from the Irish Sea at 80 m and 120 m. In

view of the present species from the

Strait of Gibraltar and of M. aciculata

from 50% N, 13 W, the geographical dis-

tribution of each Macellomenia species

appears to be limited. This is in contrast

to an earlier estimation (Salvini-Plawen,

1997) and speaks against a conspeci-

ficity of the Irish specimens with the

Mediterranean M. palifera from the Cóte

Vermeille. However, as already sur-

mised (Salvini-Plawen 1997: 44), only an

anatomical investigation of the Irish

animals can clear up the conspecificity

issue.

Family LEPIDOMENIDAE Pruvot, 1902

The family is characterised by one

type of scaly mantle sclerites, by a disti-

chous radula, by the foregut glandular

organs as subepithelial follicles (so-

called type A in SALVINI-PLAWEN, 1972,

1978), as well as by the lack of respira-

tory organs; it includes at present three

genera (cf. SALVINI-PLAWEN, 1978, 1988).

Genus Tegulaherpia Salvini-Plawen, 1983

Type species: Tegulaherpia stimulosa Salvini-Plawen, 1983.

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

Definition: Solenogastres-Lepidome-

niidae with atrial sense organ (vestibu-

lum) and mouth opening separate; ter-

minal sense organ present; secondary

genital opening unpaired; with copula-

tory stylets; no respiratory organs.

Tegulaherpia (?) spec.

Material: A single specimen was collected by Diego Moreno (Cabo de Gata / Almería) from sand

below stones at Punta del Esparto nearby Los Escullos, Cabo de Gata (Southeast Spain, » 36%50'N,

02%03'W) at 1 m depth only. After examination of the mantle scales, ribbons of semithin serial sec-

tions (cs 2 qm) were made with a glassknife and stained by Richardson's solution. The slides (mantle

scales and series section) are deposited in the Mus. Nacional Cienc. Nat., Madrid, mol. no. 15.02/14.

Description (Fig. 7): Specimen of 1.25

mm length and 0.55 mm across with an

evenly rounded body (Fig. 7 A), not yet

mature, though developing eggs visible.

Of whitish colour and smooth surface

with small, densely adjoining and

strongly overlapping, delicate scales

measuring 23 x 18 qm (Fig. 7 C) tightly

adjacent. Atrial opening with anterior

stereocilia (“bristles”), mouth and foot

well visible in life (Fig. 7 A).

Unfortunately, the animal was very

poorly preserved (partly in histolytic

state) and only a few characters can be

verified. Mouth opening distinctly sepa-

rate from the atrial sense organ and

leading into a fairly straight foregut sur-

rounded by strong longitudinal muscu-

lature. Distichous radula consisting of

paired, erect teeth with symphysis, each

provided with a distal hook and four

median denticles (Fig. 7 B). Foregut

glandular organs not traceable. Single

cerebral ganglion above the mouth (90

pm wide, 60 um high, 45 qm long) gives

rise to the connectives separately. Apart

from the gut, no organ of the posterior

body (spawning ducts, etc.) were clearly

traceable. Gut containing several nema-

tocysts, among them stenoteles, of a

hydrozoan food.

Discussion: The mantle scales, the

distinctly separate atrial sense organ

and mouth, and the distichous radula fit

well within the characters of the genus

Tegulaherpia. Other characters, however,

particularly the foregut glands and the

copulatory stylets, could not be verified

to confirm the generic classification. The

shape and size of the scales (23 x 18 yum),

and the radula (with four median denti-

cles per tooth) do not coincide with the

two known European Tegulaherpia

species (cf. SALVINI-PLAWEN, 1988, 1997).

No specific relationship can therefore be

hypothesised. The Adriatic T. stimulosa

Salvini-Plawen, 1983 possesses 30-40 ym

x 20-25 ym sized scales and 4-5 radula

denticles; the geographically overlap-

ping, West-Mediterranean and NE-

Atlantic T. myodoryata Salvini-Plawen,

1988 (= T. celtica Caudwell et al., 1995)

shows somewhat larger scales, 30-45 ym

x 20-30 um, and 4-5 radula denticles

(SALVINI-PLAWEN, 1988; HANDL AND

SALVINI-PLAWEN, 2001). Fig. 7 D shows a

scale of T. myodoryata from the most

eastern record off Malta (see MIFSUD,

1996: Fig. 3) for comparison. Most spe-

cific differences between both known

species, however, concern the arrange-

ment of the copulatory stylet apparatus

and the outlet of the (fused) spawning

duct (SALVINI-PLAWEN, 1988); no com-

parison of the present animal can there-

fore be made. According to the mantle

scales, the specimen appears to repre-

sent a species proper.

The habitat (sand) could also point

to Meiomeniidae which have mesop-

sammic species; yet, these are charac-

terised by mantle sclerites of two or

three different kinds (SALVINI-PLAWEN,

1985; GARCIA-ÁLVAREZ, URGORRI AND

CRISTOBO, 2000b).

Apart from the possible presence of

an independent, third Tegulaherpia

species, this finding is likewise of inter-

est as the shallowest record of a

solenogastre (one meter depth only).

Due to the need of calmer waters for cil-

iary gliding, Solenogastres species are

Iberus, 21 (2), 2003

generally very rare on substrata shal-

lower than about 50 meters, including

Urgorria monoplicata (below) or the

mesopsammic representatives (cf. GAr-

CÍA-ALVAREZ ET AL., 2000b). However,

exceptions such as Epimenia arabica

Salvini-Plawen and Benayahu, 1991, a

large-sized species (13-21 cm) living hid-

den in Alcyonaria-colonies at 2-5 meters

depth, occur.

Ordo CAVIBELONIA Salvini-Plawen, 1978

Solenogastres with acicular, gener-

ally hollow mantle spicules, cuticle

mostly thick, with epidermal papillae.

Remarks: As some findings indicate

(SALVIN-PLAWEN, 1978; SCHELTEMA AND

KUZIRIAN, 1991; HANDL AND SALVINI-

PLAWEN, 2002), the main character of this

taxon - the hollow acicular spicules -

strictly taken may be polyphyletic (not

supported, however, by computerised

cladograms; cf. SALVINI-PLAWEN, 2003).

Investigations have also called attention

on differences in the developmental

arrangement of the spicules at the generic

level, e.g. in Pararrhopaliidae (cf. Table D)

or in Simrothiellidae: they are either (1) in

a radial or (2) in a tangential alignment

(HOFFMAN 1949). The latter may be pro-

duced (2a) in a single, obliquely disposed

layer, or (2b) they are arranged in two or

more fairly rectangularly intercrossing

layers of low angle and almost embed-

ded within the cuticle (and also termed

“skeletal”; SCHELTEMA, 1999, SCHELTEMA

AND SCHANDER, 2000). Another criterion

exists with respect to the enclosed cavity:

the spicules may be either thick-walled or

thin-walled. Spicules may be distally

hooked (or barbed), asymmetrically flat-

tened and serrate, or asymmetrically axe-

like enlarged (termed “captate” by

ARNOVSKY, 2000).

Family PARARRHOPALIIDAE Salvini-Plawen, 1972

(Parameniidae Simroth, 1893; Paramenidae Pruvot, 1902; Perimeniidae Nierstrasz, 1909*); Pruvo-

tiniidae Heath, 1911; Pruvotinidae Scheltema, 1998)

* As NIERSTRASZ (1909: 291-292) consciously introduced a synonymy (Perimenia instead of Pru-

votina), his family “Perimeniidae” cannot be accepted as valid.

The family is characterised by a dis-

tichous radula, by the foregut glandular

organs generally as subepithelial folli-

cles (so-called type A in SALVIN-PLAWEN

1972, 1978), as well as by the additional

presence of hooked mantle spicules

and/or of a middorsal papillous pha-

ryngeal gland and/or of respiratory

organs (SALVINI-PLAWEN, 1978, GARCÍA-

ÁLVAREZ, SALVINIPLAWEN AND

URGORRI, 2001); it includes 15 genera

(Table D.

Genus Eleutheromenia Salvini-Plawen, 1967

(Paramenia Pruvot, 1890, partim [non Brauer and Bergenstamm, 1889]; Pruvotina Cockerell, 1903,

partim; Perimenia Nierstrasz, 1909, partim)

Type species: Paramenia sierra Pruvot, 1890; Costa Brava.

Definition (amended): Solenogastres

with hollow spicules in more than one

layer, also including hooked ones; with

common atrio-buccal opening; radula

present, ventral foregut glandular

organs subepithelial with paired outlet

(type A); no dorsal papillous foregut

gland; midgut with serial ventro-lateral

constrictions; with dorsoterminal sense

organ, with respiratory organs; unpaired

secondary genital opening; no receptac-

ula seminis and no copulatory stylets.

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

20 um

Figure 7. A-C: Tegulaherpia (7) sp. A: living animal from ventrolateral (sketch by Diego Moreno,

Cabo de Gata); B: one pair of radula teeth with symphysis; C: one mantle scale. D: Mantle scale of

Tegulaherpia myodoryata from off Malta. Abbreviations, as: atrial sense organ; mo: mouth opening;

fo: pedal fold (foot).

Figure 7. A-C: Tegulaherpia (2) sp. A: animal vivl en vista ventrolateral (esquema de Diego Moreno, Cabo

de Gata); B: par de dientes radulares con sínfisis; C: escala del manto. D: Escala del manto de Tegulaherpia

myodoryata de Malta. Abreviaturas, as: órgano sensorial atrial; mo: abertura bucal; fo: pie.

Eleutheromenta sierra (Pruvot, 1890)

Paramenta sierra Pruvot, 1890

Material: One specimen (preserved 11,5 mm long, with lobes up to 1,7 mm high) of typical aspect

and shape (Fig. 11) comes from the Skjórn Fjord north of Trondheim /Norway at 218 m depth (col-

lected 2.3.1992 by G. Steiner, Vienna). Ribbons of semithin serial sections of this specimen (cs 2 qm)

were made with glassknives and stained with toluidine-blue.

Type material no longer existent. The present material (series sections on slides) is deposited in the

Naturhist. Mus. Wien (NHMW), Moll. IVN 102448.

Diagnosis: Body up to 12 mm x 2 mm

with middorsal carina extended to a se-

ries Of 15 prominent lobes; cuticle mod-

erately thick, no epidermal papillae;

spicules upright, tangential and inter-

crossed, as well as hooked, all hollow;

elongate scales along the pedal groove.

Abdominal spicules in a paired bundle,

each consisting of numerous hollow

straight elements. Radula teeth with dis-

tal hook, no median denticles; ventral

foregut glandular follicles with paired

duct, midgut with rostral caecum. Vesic-

ulae seminales present, no receptacula

seminis; up to 28 gill folds. Costa Brava,

Bretagne, Irish Sea, Trondheim area; 40-

128 m (Figs. 8, 11, 15).

Body wall: Body marked by a mid-

dorsal carina elaborated into three

smaller and twelve subsequent promi-

nent lobes (Fig. 11). In these bulges, cir-

cular musculature splits into a small

subepithelial fraction (entering the

bulge) and main fibres traversing the

base of bulge, thus as usual delimiting

body cavity. Space between this split

musculature filled by mesenchyme and

large vacuole (gland) cells. Regions

between the lobes normally structured.

Epidermis covered by a 25-30 ym

thick cuticle, locally reaching 50 qm.

Especially in the posterior body, big,

pluricellular epidermal glands, but no

true epidermal papillae; only a few epi-

Iberus, 21 (2), 2003

dermal protrusions scattered ventrally

at both sides of the posterior pedal

groove (mantle rims).

Mantle with three main types of

hollow spicules (Fig. 8) in part extend-

ing beyond cuticle and thus resulting in

a somewhat rough surface. Spicule

types are (a) more or less straight ones

(130-350 ym), (b) similar type with a

harpoon-like indentation at the solid

distal portion (up to 400 pum; see

“species incerta II” in SALVINI-PLAWEN,

1978: 118-119 and Unciherpia in GARCÍA-

ÁLVAREZ ET AL., 2001), and (c) somewhat

bent spicules (up to 125 pm) with solid

distal portion forming a hook with knob

at the turn. In addition, some very

slender acicular spicules and, sporadi-

cally, ones with distal serration (d)

occur; along the pedal groove (e) elon-

gate scales (up to 80 x 13 um).

Behind foot, a pair of ventromedially

directed invaginations of the body

epithelium, each embedded within

ventral body wall musculature and

housing a bundle of straight, 130 um (or

more) long, hollow abdominal spicules

(Fig. 15). Epidermis as usual underlain

by circular and longitudinal muscles;

only weak lateroventral reinforcement

of longitudinal musculature.

Foot and mantle cavity: The ciliated

pedal pit gives rise to a single longitudi-

nal fold which runs through the pedal

groove. The foot ends, however, in front

of the abdominal spicules, not entering

the mantle cavity. The pedal gland is

voluminous, and the sole glands open

all over into the pedal fold and groove.

The mantle or pallial cavity holds at

its posterior wall blunt gill folds (28 in

Pruvor, 1890, 1891). The hindgut opens

frontally and, even more anteriorly, the

outlet of the spawnings ducts opens

ventrally. Between the end of the foot

and the opening of the mantle cavity the

invaginated bundles of the abdominal

spicula are elaborated (Fig. 15). There

are no copulatory stylets.

Sensory system: Cerebral ganglion

unpaired (240 um wide, central Y 130

pm) giving rise to connectives sepa-

rately; two pairs of small ganglia imme-

diately adjacent to it, innervating the

atrial region. Lateral body cords loosely

provided, ventral ones more densely

provided with nuclei; in the posterior

body, cords are clearly medullary. In the

ventral system, ganglia with commis-

sures fairly densely elaborated, without

correspondance to lateroventral connec-

tives. The (first) ventral ganglia (9 80

pm) showing two commissures; without

commissural sack. Buccal ganglia like-

wise prominent (Y 60 ym).

Suprarectal commissure 150 qm long

(Y 40 um) and interconnecting the

ganglia posteriora superiora medullarly;

at least two pairs of posterior nerves

originate there.

Atrial sense organ with stoutish

single or bifurcated papillae and bor-

dered by a horseshoe-shaped ciliary

fold, the dorso-posterior incurvings

exclude the buccal groove continuous

with the mouth.

Region of the dorsoterminal sense

organ (close to the body end, according

to PRUVOT, 1891) in present animal

destroyed.

Alimentary tract: Mouth opening in

the dorsoposterior area of the common

atrio-buccal cavity, connected with the

sensory region by a groove. Buccal

space with high folds leading into the

pharyngeal foregut; foregut with dis-

tinct pre-radular circular musculature

and intercellularly opening, subepithe-

lial pharyngeal glands; without com-

pacted dorsal foregut gland (with

papilla). Radula typically distichous,

each tooth (25-30 um high) with a distal

hook; median denticles could not be dis-

cerned in the sections. Radula support

represented by an assemblage of muscu-

lar and connective tissue. Ventral

foregut glandular organs as follicles

emptying at each side into distinct

opening lateroventrally of radula (type

A in SALVINI-PLAWEN, 1972, 1978).

Midgut with rostral caecum and con-

stricted only ventro-laterally by serial

dorsoventral muscle bundles to form

pouches. No nematocysts were oOb-

served within the gut. Middorsal ciliary

tract widens to cover entire hindgut

opening into mantle cavity above and

behind unpaired spawning duct outlet.

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

ANA

(9)

(q)

50 um

Figure 8. Eleutheromenia sierra (Pruvot) from the Trondheim area, types of mantle sclerites.

Figura 8. Eleutheromenia sierra (Pruvot) del área de Trondheim, tipos de escleritas del manto.

Gonopericardial system: Paired gonad

showing developing germ cells in the

posterior region within lateral sacks, the

most posterior pair representing vesicu-

lae seminales. Paired gonopericardial

ducts opening dorsofrontally into peri-

cardium with a paired anterior begin-

ning. Pericardium voluminous with

paired lateral ciliary tract and containing

mature eggs (Y 140 qm or 150 x 130 qm);

probably due to such conditions, PRU-

voT (1891) cosistently termed the peri-

cardium as “sac oOvigere”. Heart

throughout as mediodorsal invagination,

anteriorly very wide, then hanging by a

double “mesenterium” into the pericar-

dial lumen. Atrium clearly paired and

fusing just before the axial transition into

ventricle. Two kinds of blood cells:

round, homogeneous ones (Y 10 ym)

and oval to round, vacuolated or vari-

ously granulated cells (10-20 um long).

Pericardioducts emerging posteri-

orly, forming in their curve towards

anterior short sacculations (vesiculae

seminales) and opening laterodorsally

into respective spawning duct. No

receptacula seminis present. Spawning

ducts volumirous, highly glandular,

and subdivided into two equal sections:

paired portion not axially continuous

with unpaired section, the latter extend-

ing somewhat below the former, with

paired dorsoventral interconnection.

Unpaired section opening by means of

simple outlet ventrally to exterior; this

opening representing the anterior-most

area of the mantle cavity opening.

Discussion: Though the present speci-

men comes from Skjórn Fjord (Trond-

heim region), the type locality of Parame-

nia sierra (Pruvot) is off the Costa Brava

(Catalunia /Spain); for comparison with

the description by Pruvot (1891), a full

descriptive presentation of the new

finding was given here. The specimen

from Norway exhibits an almost identi-

cal organisation to E. sierra which

includes, for example, also the dorso-

frontal opening of the gonopericar-

dioducts into the pericardium (PRUVOT,

1891: Fig. 17). Some characters are not

mentioned by Pruvot (such as, e.g. the

posterior-most gonad serving as seminal

vesicles, or the subdivision of the spawn-

ing ducts); there appear to be, however,

just three discrepancies. First, the mantle

spicules are not exactly identical.

Second, in the region of the posterior

body drawn as a cross section by

PruvoT (1891: Fig. 61), Pruvot claims a

“glande cloacale” (Fig. 61: y). Here, apart

Iberus, 21 (2), 2003

from muscle fibres, numerous sole

glands, the medullary ventral nerve

cords as well as two connected ganglia

innervating the spawning ducts are

present in the Norwegian specimen.

Some of these structures could represent

Pruvot's gland as no real glandular

organ is discernible; also the distance to

the mantle cavity (“cloaca”) is too long

in both Pruvot's animal and the present

specimen to represent a pallial gland.

A third and more essential point of

disagreement of the new specimen with

Paramenia sierra appears due to PRUVOT's

(1891: 790) somewhat misleading decrip-

tion of the “petits caecums” at the end of

the pedal groove, which he equalises with

“Y appareil spiculaire pénial de Proneome-

nia vagans” [= Pararrhopalia pruvoti

Simroth, 1893]. This equalisation led to the

(incorrect) definition of Paramenia sierra as

possessing copulatory stylets (THIELE,

1913b, SALVINEPLAWEN, 1967b). As Pruvot,

however, describes correctly in detail

(Pruvor, 1891: 790), P. sierra possesses

“une paire de petits caecums” (plural!)...

“plongés dans les muscles longitudineaux

ventraux, dans chacun desquels” he recog-

nised “un faisceau de spicules”. This is an

exact description of a paired bundle of

abdominal spicules, such as exists in the

present individual (Fig. 15).

The Norwegian specimen described

here is thus regarded as conspecific with

Eleutheromentia sierra (Pruvot).

Apart from the single, no longer

existent type specimen (11-12 mm x 2

mm) of the original description by PRU-

voT (1890, 1891) from off Portaló Island

(Cap Creus/Costa Brava) at 75-80 m,

and a finding from off Roscoff / Bretagne

at about 40 m (Pruvot, 1897), several

other records have been reported from

the Irish Sea (cf. SALVINI-PLAWEN, 1997).

Though without closer examination,

due to their typical appearance (lobu-

lated keel), however, these latter speci-

mens may likewise belong to E. sierra;

this is biogeographically supported by

Pruvot's record from off Roscoff and by

the present specimen (Trondheim area).

Paramenta sierra Pruvot was generically

transferred by SALVINI-PLAWEN (1967b) to

a new genus Eleutheromenia. A second

species, Pruvotina impexa THIELE (1913a),

after re-examination was generically sep-

arated later as Labidoherpia impexa (Salvini-

Plawen, 1978). Most recently, SCHELTEMA

(1998b) and SCHELTEMA AND SCHANDER

(2000) described two Eleutheromenia species

from the Bass Strait, South of Australia.

According to the descriptions and a pre-

liminary re-examination of specimens (E.

mimus from Slope Station 40, E. bassensis

from BSS-S 202; a full description of the

anatomies is in preparation by Cl. Handl,

Wien /Vienna, Austria), both these latter

species differ markedly in several charac-

ters from Eleutheromenia sierra (PRUVOT,

1891 and above):

(1) They do possess copulatory

stylets, in contrast to E. sierra (with

abdominal spicules, see above);

(2) The ventral foregut glandular

organs are tubular with epithelial gland

cells (type C in SALVINI-PLAWEN 1972,

1978);

(3) In both species a “ventral com-

missure sack” of unknown function is

present, similarily as in the gymnomeni-

ids Genitoconia, Wirenia [= Aesthoherpia]

and Gymnomenia (SALVINI-PLAWEN,

1967a, 1988, HASZPRUNAR, 1986, SCHEL-

TEMA, 1999);

(4) The glandular anterio-ventral

portion of the pericardium (not in open

communication with the mantle cavity)

- in connection with the pericardial

glands - speculatively might represent

part of some kind of excretory system.

With respect to the generic charac-

ters among the family Pararrhopaliidae

(Table D, at least the above characters (1)

and (2) concern the generic level. Apart

from the different type of the foregut

glandular organs in both Australian

species (see also Unciherpiinae in GAR-

CÍA-ÁLVAREZ ET AL., 2001), copulatory

stylets are only present in Labidoherpia

Salvini-Plawen (with papillous dorsal

pharyngeal gland) and in Pararrhopalia

Simroth (with papillous gland, mouth

separate). Thus, both these species, E.

mimus and E. bassensis, do not fit into an

existing genus and consequently are

transferred to a separate genus Schel-

temaia gen.nov. (see also the cladistic

analysis in SALVINI-PLAWEN, 20093).

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

Table I. Generic characters in Pararrhopaliidae (see SALVINI-PLAWEN 1967b, 1978; GARCÍA-

ÁLVAREZ ET AL., 2001, GARCÍA-ÁLVAREZ AND URGORRI, 2001). A: ventral foregut glandular

organs with subepithelial follicles (type A); C: ventral foregut glandular organs epithelial (type C);

dts: dorso-terminal sense organ; pphf: peri-pharynggeal ring of follicular glands; -: absent; +:

present.

Tabla 1. Caracteres genéricos en Pararrhopaliidae (ver SALVINI-PLAWEN 1967b, 1978; GARCÍA-

ÁLVAREZ ET AL., 2001, GARCIA-ÁLVAREZ Y URGORRI, 2001). A: órganos glandulares digestivos ven-

trales con folículos subepiteliales (tipo A); C: órganos glandulares digestivos ventrales epiteliales (tipo C);

dts: órgano sensorial dorso-terminal; pphf: anillo perifaríngeo de glándulas foliculares; -: ausente; +: pre-

sente.

hooked dorsal ventral respiratory mouth copulatory radula cutide dis

spicules gland glonds folds separate stylets

Pruvotina + + A + + thick +

Pararrhopalia + + A - + 4 + thick +

Labidoherpia + + A + + + thick +

Eleutheromenia + A + + thick +

Gephyroherpia + A + + + thick +

Luitfriedia + A + thick +

Lophomenia - + A ? + + thick +

Metameria A + + thick +

Hypomenia A + + thick ?

Halomenia A + + - + thick +

Forcepimenia A ? + ? + thin

Scheltemaia + C + + + thick +

Unciherpia + pphf + thin +

Uncimenia - (2) pphf + + - - thin +

Sialoherpia - - pphf ? ? - thick +

Scheltemaia gen. nov.

Type species (with respect to the more accurate description): Eleutheromenia mimus Scheltema

and Schander, 2000; Bass Strait (South Australia).

Definition: Solenogastres-Parar- 1972, 1978); no dorsal papillous foregut

rhopaliidae with hollow spicules includ- gland; with “commissure sack” between

ing hooked ones; with common atrio- (first) ventral ganglia; with dorsotermi-

buccal opening; radula distichous; nal sense organ, with respiratory or-

paired ventral foregut glandular organs gans; unpaired secondary genital open-

each as tubes or sacks with epithelial ing; with copulatory stylets; no recepta-

gland cells (type € in SALVINI-PLAWEN cula seminis. Two species known.

Family RHOPALOMENIIDAE Salvini-Plawen, 1978

The family is characterised by a thick glandular organs subepithelial (type A)

cuticle and hollow acicular spicules in and/or epithelial (type C; SALVINI-

several layers, without hooked spicules; PLAWEN, 1972, 1978); no respiratory

radula distichous or lacking; no papillate organs. It includes five genera (GARCÍA-

dorsal foregut gland, ventral foregut ÁLVAREZ AND SALVINI-PLAWEN, 2001).

DS)

Iberus, 21 (2), 2003

os |

100 um

Ear dad ca

Figure 9. Urgorria monoplicata spec. nov. Organisation of the anterior body. Abbreviations, as:

atrial sense organ; bw: body wall (mantle and musculature); ca: midgut caecum; ce: cerebral gan-

elion; fo: pedal fold (foot); gl: ventral foregut glandular organ; go: gonad; mu: muscle bundle; ph:

pharynx.

Figura 9. Urgorria monoplicata spec. nov. Organización del cuerpo anterior. Abreviaturas, as: órgano

sensorial atrial, bw: pared del cuerpo (manto y musculatura); ca: ciego digestivo; ce: ganglio cerebral; fo:

pie; gl: órgano glandular digestivo ventral; go: gónada; mu: unión muscular; ph: faringe.

Genus Urgorria García-Álvarez and Salvini-Plawen, 2001

Type species: Urgorria compostelana García-Álvarez and Salvini-Plawen, 2001; off northwestern

Spain.

Definition: Solenogastres-Rhopalo- foregut glandular organs epithelial

meniidae with epidermal papillae and (type C in SALVINI-PLAWEN 1972, 1978);

hollow acicular spicules in several inter- dorso-terminal sense organ present; sec-

crossing layers; mouth opening within ondary genital opening unpaired, no

the atrium; without radula; paired copulatory stylets.

Urgorria monoplicata spec. nov.

Material: A single specimen of 2.2 mm length (9 0.5 mm) with evenly rounded body end origi-

nates from off the Costa Brava (Girona /Spain), from the “Furrió de Tamariu”, sandy bottom in 35

m depth (collected 16.9.1990 by Luis Dantart, Barcelona).

After examination of the sclerites, ribbons of semithin serial sections (cs 2 qm) were made with a

glassknife and stained with RICHARDSONSs solution.

The specimen (series sections on slides) is deposited as holotype in the Mus. Nacional Cienc. Nat.,

Madrid, mol. no.15.02/13.

Derivatio nominis: Greek monos = one, Latin plica = fold; referring to the single pedal fold.

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

Figure 10. Micromenia subrubra spec.nov. (paratype 2), preserved 5 mm specimen; note beak-like ex-

tension of dorsal body end (below). Figure 11. Eleutheromenia sierra (Pruvot) from Trondheim area,

preserved animal (11.5 mm long). Figure 12. Micromenia subrubra spec.nov., cross section through

posterior body behind pericardium [hg: hindgut; po: anterior pouch of mantle cavity with separated

anterio-ventral portion; sd: spawning duct] scale bar: 50 um. Figure 13. Urgorría monoplicata spec.nov.,

cross section through mantle bridge (retracted) between atriobuccal opening and pedal pit with pre-

pedal scales (sc) [ca: midgut caecum (paired); ce: cerebral ganglion; ph: pharynx] scale bar: 100 um.

Figura 10. Micromenia subrubra spec.nov. (paratipo 2), especimen fijado, $ mm; nótese la extensión

con forma de pico en la parte final (abajo). Figura 11. Eleutheromenia sierra (Pruvot) del área de

Trondheim, animal fijado (11.5 mm). Figura 12. Micromenia subrubra spec.nov., sección del cuerpo

posterior tras el pericardio [hg: intestino posterior, po: saco anterior de la cavidad del manto con porción

separada antero-ventral, sd: conducto de puesta] escala: 50 qm. Figura 13. Urgorria monoplicata

spec.nov., sección entre la abertura atriobucal y la foseta pedal con escalas pre-pedeales (sc) [ca: ciego

digestivo, ce: ganglio cerebral, ph: faringe] escala: 100 jon.

Iberus, 21 (2), 2003

Diagnosis: Body 2.2 mm x 0.5 mm,

with evenly rounded ends; cuticle fairly

thin without keel formation; foot with

one single fold only until mantle cavity;

no radula or vestige of radula sheath,

ventral foregut glandular organs as very

small sacks opening ventrolaterally, ter-

minal foregut with two pairs of strong

muscle bundles to lateral body wall;

midgut with large rostral caecum, ros-

trally paired, and with regular constric-

tions. Spawning ducts in their distal half

fused, with simple opening; one pair of

non-stalked, dorsoposterior receptacula

seminis. Figs 9, 13, 14.

Mantle: Mantle producing a moder-

ately thick cuticle only (40-50 pm)

without marked thickenings; epidermal

papillae distally balloon-shaped.

Spicules (average 100 um long) acicular

and hollow, in tangential and intercross-

ing layers; no hooked or distally serrate

spicules; along foot elongate scales

(about 40 qm). Mantle bridge between

atrio-buccal opening and pedal pit like-

wise with distinct scales (Fig. 13).

Foot and mantle cavity: Pedal pit

receiving outlets of the voluminous

pedal gland follicles that extend at both

sides in the anterior body. Ciliated

epithelium of pit not forming folds. At

end of pit a single, distinct fold is differ-

entiated, continuing until the pallial

cavity. Sole glands parallel internally the

foot. Mantle cavity representing a

simple ciliated space receiving dorsome-

dially the rectum and medioventrally

the single outlet of the posteriorly fused

spawning ducts. There are neither respi-

ratory organs nor accessory genital

organs.

Musculature: Body wall musculature

not very prominent, longitudinal

portion showing an only moderately

elaborated ventral reinforcement. Simi-

larly, the dorsoventral bundles are typi-

cally differentiated, but not very strong.

Their fairly medial course serially in dis-

tances of 50-80 um causes deep constric-

tions of the midgut, forming respective

lateral pouches. Special musculature is

elaborated in connexion with the poste-

rior foregut.

Sensory system: Fused cerebral gan-

glion above the mouth exhibiting a tri-

angular outline in cross section (ven-

trally 125 qm wide, 60 qm high, 50 qm

long). Laterofrontal pairs of small gan-

glionic formations innervate the atrial

sense organ and the mouth opening.

The connectives emerge laterally, the

very short (20 um) ventral and lateral

ones in close contact. The (first) lateral

ganglion at each side is latero-posteri-

orly elongated. The first “ventral”

swelling, due to the short connective, is

(Right page) Figure 14. Urgorria monoplicata spec. nov., somewhat oblique cross section through

region of terminal foregut [bu: right buccal ganglion; dv: dorsoventral muscle bundle; fo: pedal

fold (foot); gl: left foregut gland; mg: midgut; mu: musculature forming bundles; nv: medullary

ventral nerve cord; pg: (portion of) pedal gland; ph: pharynx; sg: sole gland], scale bar: 30 um.

Figure 15. Eleutheromenia sierra (Pruvot) from Trondheim area, cross section through pre-pallial

(post-pedal) region with abdominal spicules (sp) [sd: spawning duct], scale bar: 50 um. Figure 16.

Macellomena adenota spec. nov., somewhat oblique cross section through pre-pallial (post-pedal)

mantle invagination (mi) with “abdominal gland” (ag) just before opening, and with opening of

pouch of abdominal spicules (sp) [mg: midgut], scale bar: 30 um.

(Página derecha) Figura 14. Urgorria monoplicata spec. nov. Sección oblicua a la altura de la región

terminal del digestivo anterior [bu: ganglio bucal derecho; dv: musculatura dorsoventral; fo: pie; gl:

glándula digestiva izquierda; mg: intestino medio; mu: unión muscular terminofaringeal; nu: cordón

nervioso ventral; pg: (parte de) glándula pedal; ph: faringe; sg: glándula de la suela), escala: 30 jom.

Figura 15. Eleutheromenia sierra (Pruvot) del área de Trondheim, sección de la región prepaleal (post-

pedal) con espículas abdominales (sp) [sd: conducto de la puesta), escala: 50 qm. Figura 16. Macellom-

ena adenota spec. nov., sección a la altura de la invaginación prepaleal (post-pedal) del manto (mi) con

“glándula abdominal” (ag) justo antes de la abertura y con la abertura de la bolsa de espículas abdomi-

nales (sp) [mg: intestino medio], escala: 30 ym.

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

Iberus, 21 (2), 2003

located lateroventrally of the cerebral

ganglion; the true ventral ganglia (9 60

pm x 30 um) are formed above the

beginning pedal fold. The remaining

nervous system exhibits no unusual

configuration. The 125 pm long

suprarectal commissure is medullary (Y

20 qm).

The atrial sense organ is bordered by

the horseshoe-shaped ciliary tract, the

dorsal limbs running parallel in distinct

distance. The papillae are single and

blunt (Y 12-15 qm). A single dorsotermi-

nal sense organ is elaborated middor-

sally above the posterior rectum.

Alimentary tract: Mouth opening dor-

sally at the rear end of the atrium,

behind the curved ciliary tracts. Foregut

representing an elongate tube, coated

throughout its course by a delicate cir-

cular and a distinct longitudinal pha-

ryngeal musculature. The epithelium

consists in the anterior third of cubical

cells (about 6 qm high), further on of

irregularly club-shaped cells (8-12 yum

high). It is intruded (partly in the ante-

rior third, all along posterior third) and

forms up to 12 distinct longitudinal

folds.

A short distance in front of the

ventral opening of the foregut into the

midgut, the dorsal and lateral longitudi-

nal pharyngeal muscles are concen-

trated to a pair of dorsolateral bundles

which run transversely to the lateral

body wall (Figs 9, 14). In that narrow

area, the ventral pharyngeal muscles

also become concentrated to a pair of

ventral bundles; both extend (in the

present specimen) asymmetrically at the

same body side far posteriorly to join

the musculature of the body wall (not

before the middle of the body).

In this terminal portion of the

foregut (formation of muscle bundles)

the buccal ganglia (Y 25 ym) are elabo-

rated laterally (Fig. 14). The ventral

glandular organs are separated by the

buccal commissure ventral to the

foregut; they open ventrolaterally into

the foregut (Fig. 14). The organs are very

small (40-50 qm long, Y 30 qm), some-

what irregular sacks with glandular

epithelium and a narrow outleading

lumen (type C in SALVIN-PLAWEN 1972,

1978).

Midgut possessing a voluminous

rostral caecum extending to body tip

(Fig. 9), splitting frontally in the region

of the mouth to a paired organ. Due to

the serial muscle bundles, there are

regular, deep lateral constrictions of the

midgut. The midgut and its caecum

include, among dissolved food material,

also nettle capsules of Cnidaria.

Gonopericardial system: Paired,

hermaphoditic gonad. Eggs at the con-

necting median walls measure maxi-

mally Y 50 ym, indicating that full

maturity has not been reached. The

single heart auricle is an invagination of

the pericardial roof, and the ventricle

represents a largely free organ. There

are two kinds of blood cells: a) small

and coarsely structured granulocytes (Y

5 ym); b) finely granulated, round

haemocytes (9 10 qm).

The pericardioducts emerge lat-

eroterminally. They open from dorsal

into the respective spawning duct, the

latter bending dorso-posteriorly into a

scarcely separated, wide tube: receptac-

ulum seminis. The spawning ducts are

paired in their anterior portion; single

opening into the central mantle cavity

represents a simple, narrowed and short

outlet (pore). There are neither vesiculae

seminales nor accessory genital organs

(copulatory stylets, abdominal spicula).

Discussion: Among the five genera of

the family Rhopalomeniidae (GARCÍA-

ÁLVAREZ AND SALVINI-PLAWEN, 2001),

the present specimen coincides on the

generic level with Urgorria based on the

mantle elaboration, the atrio-buccal

cavity, the lack of the radula and the

configuration of the genital apparatus.

The conformity with the hitherto sole

species U. compostelana even includes

such characters as the special scales at

the mantle bridge between the atrio-

buccal opening and the pedal pit, the

similar musculature of the foregut, the

unpaired to paired midgut caecum, and

the receptacula seminis. Only few char-

acters actually differ from those in Ul.

compostelana: the body cuticle forms no

middorsal reinforcement (in U. com-

SALVINI-PLAWEN: West-Mediterranean Solenogastres with three new species

postelana the 40-100 yum thick cuticle

with spicules locally forms a middorsal

crest up to 150 qm); only a single pedal

fold is elaborated, which continues to

the mantle cavity (in U. compostelana

there are two pedal folds which fuse

into a single one in the region of the

anterior spawning ducts and flatten to a

mere ridge close to the mantle cavity to

a mere ridge); the position of the dor-

soterminal sense organ differs (above

the anterior mantle cavity; in U. com-

postelana at the rear of the body); the

heart ventricle represents a free tube (in

U. compostelana it remains an invagina-

tion throughout); the different relation

of the foregut muculature (the muscle

layer around the foregut in U. composte-

lana is weaker than the bundles at the

terminal foregut, which are also much

stronger than those in the present speci-

men 1). The present specimen thus rep-

resents a proper species Urgorria mono-

plicata spec.nov.

The common atrio-buccal opening in

Urgorria separates this genus from all

other Rhopalomeniidae, and Urgorria is

thus well-defined (GARCÍA-ALVAREZ

AND SALVINI-PLAWEN, 2001). At the

same time, the condition of the ventral

foregut glandular organs (type C in

SALVINI-PLAWEN, 1972, 1978) required

somewhat altering the definition of the

BIBLIOGRAPHY

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PLAWEN, 1978). The type species

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O Sociedad Española de Malacología Iberus, 21 (2): 61-65, 2003

Poliquetos asociados a Isognomun alatus (Gmelin, 1791)

(Bivalvia: Isognomonidae) en la costa nororiental de

Venezuela.

Polychaete worms associated to /sognomun alatus (Gmelin, 1791)

(Bivalvia: Isognomonidae) from northeastern coast of Venezuela

Óscar DÍAZ DÍAZ e Ildefonso LINÑERO-ARANA*

Recibido el 9-1X-2002. Aceptado el 9-1V-2003

RESUMEN

Treinta y ocho especies de poliquetos fueron encontradas asociadas a 175 ejemplares de

la ostra boba Isognomun alatus (Gmelin, 1791) colectados en la costa nororiental de

Venezuela. Las familias Serpulidae y Eunicidae fueron las mejores representadas con trece

y siete especies, respectivamente. Entre las especies identificadas, Caulleriella alata Sout-

hern, 1914, Eunice cariboea Grube, 1856 y Nematonereis hebes Verrill, 1900, constitu-

yen primeros registros para Venezuela.

ABSTRACT

Thirty-eight polychaete species were found on valves of 175 specimens of the oyster Isog-

nomun alatus (Gmelin, 1791) collected in the northeastern coast of Venezuela. Serpulidae

and Eunicidae were the families best represented with thirteen and seven species, respecti-

vely. Caulleriella alata Southern, 1914, Eunice cariboea Grube, 1856 and Nematonereis

hebes Verrill, 1900, are new records for Venezuela.

PALABRAS CLAVE: Bivalvos, /sognomun alatus, Isognomonidae, Poliquetos, epifauna, Gran Caribe.

KEYS WORDS: Bivalves, lsognomun alatus, Isognomonidae, Polychaetes, Epifauna, Great Caribbean.

INTRODUCCIÓN

La ostra boba, Isognomun alatus

(Gmelin, 1791), es un bivalvo común de

las costas venezolanas (GÓMEZ, 1999).

Este molusco está ampliamente distri-

buido en el Mar Caribe desde Florida

hasta el sur de Brasil (ABBOTT, 1974;

SIUNG, 1980; Díaz Y PUYANA, 1994) se de-

sarrolla a poca profundidad y se encuen-

tra asociado principalmente a substratos

duros, especialmente a raíces de man-

gles, donde llega a formar densas colo-

nias (ABBOTT, 1974). A pesar de ser muy

abundante es la menos conocida entre

las especies del género, entre los pocos

estudios se citan los trabajos de TRUE-

MAN Y LOWE (1970) quienes estudiaron el

efecto de la temperatura y exposición li-

toral sobre la fisiología cardíaca, y SIUNG

(1980) sobre la biología de la especie.

Este bivalvo es desestimado comer-

cialmente por la escasa cantidad de

carne, sin embargo, en algunas comuni-

* Instituto Oceanográfico de Venezuela-Universidad de Oriente, Cerro Colorado, Edif. 1.O.V. Piso 2, Lab. 208.

Cumaná, Estado Sucre-Venezuela. e-mail: ecobentos1 24 hotmail.com, linerofPhotmail.com

Iberus, 21 (2), 2003

dades es consumido por sus pobladores

y es ofrecido engañosamente a los turis-

tas como ostras de mangle (Crassostrea

rhizophorae Guilding, 1828). SIUNG (1980)

señala que en Jamaica, el consumo de 1.

alatus se ha incrementado considerable-

mente debido a la reducción de las áreas

de C. rhizophorae, ocasionada principal-

mente por la sobre-explotación de la

misma y destrucción del manglar.

La superficie de las valvas de este

molusco es lisa-escamosa, proporcionando

un substrato adecuado para la fijación de

organismos epibiontes que completarán

allí su ciclo de vida; por otro lado, la dis-

posición del molusco en grupos con nume-

rosos individuos, muy próximos entre sí,

brinda microhábitats disponibles a ser

colonizados por otras especies, además de

servir refugio contra los depredadores. En

el Gran Caribe existen pocos estudios sobre

la fauna asociada a moluscos bivalvos de

interés comercial, excepto el de LIÑERO-

ARANA (1999) sobre poliquetos asociados

a de Perna viridis (Linné, 1758).

En el presente trabajo se analiza la

comunidad de poliquetos asociados al

bivalvo 1. alatus colectados de las raíces

de mangles en la costa nororiental de

Venezuela.

MATERIAL Y MÉTODOS

El estudio se realizó entre enero y abril

de 2002, los organismos fueron colectados

en la Bahía de Mochima (64? 25" 10” N, 11?

15" 20” O), los mismos fueron retirados de

las raíces del mangle Rhizophora mangle

(Linné, 1753), y colocados en bolsas plásti-

cas debidamente etiquetadas y éstas, a su

vez, dentro de cavas con hielo y agua de

mar para su traslado. Una vez en el labo-

ratorio, se procedió a colocar los bivalvos

en acuarios con aireación. Posteriormente,

se procedió a separar los poliquetos de la

superficie de las valvas y a fijarlos en una

solución de formaldehído al 8% en agua

de mar. Para la identificación de los mis-

mos se emplearon las claves regionales:

JONES (1962), BLAKE (1971), Day (1973),

SALAZAR-VALLEJO Y CARRERA-PARRA (1997),

UEBELACKER, JONHSON Y VITTOR (1984),

LINÑERO-ARANA, (1996), entre otras.

RESULTADOS Y DISCUSIÓN

Se analizaron 175 ejemplares del

bivalvo 1. alatus y 658 ejemplares de poli-

quetos identificándose 38 especies perte-

necientes a 12 familias (Tabla 1), entre

éstas, las mejores representadas fueron

Serpulidae con trece especies, lo que

representa el 34,21% del total de especies

identificadas, seguida por Eunicidae

(18,42%) con siete y Terebellidae (10,53%)

con cuatro. Mientras que, en cuanto a la

abundancia por familia, Serpulidae pre-

sentó el mayor número de individuos

(59,27%), seguida por Syllidae (10,49%) y

Terebellidae (8,21%). Entre las especies

más abundantes destacan Pileolaria milita-

ris Claparede, 1868 (34,65%), Salmacina

sp. (8,21%), Exogone dispar (Webster, 1879)

(6,84%) y Serpula sp. (6,23%).

I. alatus habita principalmente en la

zona submareal conformando densas

agrupaciones, aunque es capaz de sobre-

vivir en la zona intermareal, en zonas de

gran turbidez, en las que otros organismos

filtradores no podrían habitar, SIUNG (1980)

señala que puede soportar rangos de sali-

nidad entre 10,9 - 40,0%o demostrando

una gran adaptabilidad a condiciones

extremas e indica que la especie crece y se

reproduce aun en presencia de especies

incrustantes, sin embargo, no las men-

ciona. Estas agregaciones de individuos

proporcionan una mayor cantidad de

microhábitats para la macrofauna bentó-

nica, y específicamente a los poliquetos, así

como protección contra los depredado-

res, alimento y áreas de reproducción. Por

otro lado, la superficie escamosa del

molusco permite la colonización de aque-

llas especies sedentarias, principalmente

serpúlidos, que en algunos casos llegan a

cubrir casi por completo la superficie de

las valvas, y terebélidos.

Entre los poliquetos, varias especies

de Polydora Bosc, 1802, han sido identifi-

cadas como plagas de cultivos de bival-

vos, especialmente de pectínidos

(CIOCCO, 1990; BASILIO, CANETE Y ROZ-

BACZYLO, 1995; (CÁCERES-MARTÍNEZ,

2001), afectando principalmente el creci-

miento del bivalvo, debilitando la

concha y haciéndolo más susceptible a

infecciones O ruptura de las valvas. P.

Díaz Y LINERO-ARANA: Poliquetos asociados a lsognomun alatus en Venezuela

Tabla I. Poliquetos asociados a la ostra boba /. alatus en la Bahía de Mochima. MI: móvil intersti-

cial; Er: errante; E: epibionte; Sl: sedentario intersticial.

Table 1. Polychaete associated to 1. alatus in Bahía de Mochima. MI: mobile interstitial: Er: errant; E:

epibiont; SI: sedentary interstitial.

Especie Localidad Material Microhúbitat —— Distribución

tipo examinado

Spionidae

Polydora websteri Hartman, 1943 Nueva Inglaterra 39 SI Pacífico (Golfo de

California, Hawai, Chile),

Atlántico (Quebec, New

Foundland hasta Florida, Golfo

de México, Brasil, Argentina,

Venezuela)

Cirratulidae

Coulleriella alata Southern, 1914 Mar de Irlanda 16 SI Cosmopolita

Phyllodocidae

Phyllodoce (Anaitides) madeirensis Langerhans, 1880. Madeira l MI Cosmopolita

Hesionidae

Podarke obscura Verrill, 1873 Mar Caribe 11 MI Gran Caribe

Hesione splendida Savigny, 1818 Mar Rojo 2 MI Cosmopolita

Syllidae

Syllis (Typosyllis) lutea Hartmann-Schróder, 1960 Mar Rojo 21 MI Cosmopolita

Ehlersia cornuta Raihke, 1843 Noruega 3 MI Cosmopolita

Exogone dispar (Webster, 1879) Virginia 45 MI Cosmopolita

Nereididae

Nereis riisei Grube, 1857 Mar Caribe 6 MI Gran Caribe

Ñ. falsa Quatrefages, 1865 Mar Mediterráneo 3 MI Cosmopolita

Amphinomidae

Euryihóe complanata (Pallas, 1766) Mar Caribe 14 Er Circuntropical

Eunicidae .

Eunice vittata (delle Chiaje, 1828) Nápoles 7 Er Cosmopolita

Eunice cariboea Grube, 1856 Mar Caribe 10 Er Costas americanas del

Ailántico y del Pacífico

E. aphroditois (Pallas, 1788) Océano indico l Er Circuntropical

Lysidice ninefta Audouin y Milne-Edwards, 1833 — Chancey, Isla (Francia) 7 Er Cosmopolita

Lysidice collaris Grube, 1870 Mar Rojo l Er Circuntropical

Marphysa sanguinea (Montagu, 1815) Costas de Devon (Inglaterra) 4 Er Cosmopolita

Nematonereis hebes Verrill, 1900 Mar Adriático 4 MI Cosmopolita

Dorvilleidae

Dorvillea cerasina Ehlers, 1901 Chile 4 Er Costas americanas del

Atlántico y del Pacífico

Terebellidae

Terebella pterochaeta Schmarda, 1861 Cabo de la Buena 24 SI-E Cosmopolita

Esperanza

Terebella sp. - SI -

Streblosoma hartmanae Kritzler, 197 Mar Caribe 22 SHE Gran Caribe

Polycirrus denticulatus Saint-Joseph, 1894 Francia 4 SI Atlántico

Iberus, 21 (2), 2003

Tabla I. Continuación.

Table I. Continuation.

Especie Localidad Material Microhábitat — Distribución

tipo examinado

Sobellidae

Branchiomma nigromaculata (Baird, 1865) Antillas 8 SI-E Circuntropical

Hypsicomus phaeotaenia Schmarda, 1861 Mar Mediterráneo SI-E Circuntropical

Serpulidae

Spirobranchus giganteus giganteus (Pallas, 1766) Antillas 2 E Gran Caribe

S. dendropoma (Schmarda, 1861) Antillas l E Gran Caribe

S. tetraceros (Schmarda, 1861) Antillas 2 E Cosmopolita

Hydroides dirampha Mórch, 1863 Antillas 32 E Circuntropical

H. bispinosa Bush, 1910 Bermuda 4 E Gran Caribe

H. elegans (Haswell, 1883 Port Jackson (Australia) 1 E Cosmopolita

Hydroides sp. - l E :

Vermiliopsis annulata (Schmarda, 1861) Jamaica 10 E Costas americanas del

Atlántico y del Pacífico

Pseudovermilia occidentalis (McIntosh, 1885) Bermuda 12 E Costas americanas del

Ailántico y del Pacífico

Salmacina sp. 54 E -

Serpula sp. - 41 E -

Protula submedia Augener, 1906 Mar Caribe (Sur) 2 E Gran Caribe

Pileolaria militaris Claparede, 1868 Francia 228 E Cosmopolita

Total de individuos 658

websteri Hartman, 1943, único espiónido

registrado en este estudio, representa

menos del 6% del total de poliquetos

colectados, siendo su abundancia y fre-

cuencia relativamente baja, conside-

rando que se analizaron 175 ejemplares

del bivalvo. Los tubos mucosos del poli-

queto están principalmente adheridos a

los filamentos del biso, y no se observa-

ron ampollas ni tubos mucosos sobre las

valvas ni en el interior de ellas.

Al comparar los resultados obteni-

dos en este trabajo con otros similares,

se Observa que el número de especies

registrado es superior a la mayoría.

KEOUGH (1984) registró tres especies de

espirórbidos como epibionte de Pinna

bicolor Gmelin, 1791, señalando que el

número de individuos era considerable-

mente bajo y que constituye menos del

2% de los epibiontes; LINERO-ARANA

(1999) registró 11 asociadas a P. viridis; y

DE LEÓN, LENA-TRISTÁN Y SALAZAR-

VALLEJO (1993), registraron 28 especies

asociadas a Spondylus princeps unicolor

Sowerby, 1847, siendo Serpulidae la

familia mejor representada, tanto en

número de especies como en abundan-

cia, con siete especies, mientras que en

el presente estudio se registran 38 espe-

cies de las cuales 13 corresponden a ser-

púlidos, la dominancia de esta familia

en ambos es debido a que las conchas de

los bivalvos constituyen un substrato

apropiado para su colonización y esta-

blecimiento de tubos de carbonato de

calcio construidos por los miembros de

esta familia. La presencia de sabélidos y

terebélidos, especies que normalmente

construyen sus galerías en el sedimento,

sobre las valvas obedece a la acumula-

ción de sedimento entre las escamas lo

que permite la colonización de estos

microhabitats.

La distribución geográfica indica

un predominio de especies cosmopo-

DÍAZ Y LINERO-ARANA: Poliquetos asociados a lsognomun alatus en Venezuela

litas (15) lo que representa más del

44%, seguido por aquellas especies

cuya distribución está restringida al

Gran Caribe (20,59 %), las circuntropi-

cales (17,65%), las anfiamericanas, es

decir que se encuentran tanto en las

costas americanas del Pacífico y del

Atlántico (11,76%), una sola especie (P.

websteri) que se comporta como

disyunta (anfiamericana-transpacífica),

dado que ha sido registrada en Hawai,

y una restringida al Atlántico, ambas

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with a key to all species recorded from North

Carolina. NOAA Technical Report NMES Circ.

375: 1-140.

De León GONZALEZ, J. A., LEIJA-TRISTÁN, A.Y

SALAZAR-VALLEJO, S., 1993. Epifauna del os-

tión espinoso Spondylus princeps unicolor (Mo-

llusca: Bivalvia) de Puerto Escondido, Golfo

de California, México. Revista de Biología Tro-

pical, 41 (3): 877-881.

Díaz ,J. M. Y PUYANA. M,, 1994. Moluscos del

Caribe Colombiano. Un catálogo ilustrado. COL-

CIENCIAS, Fundación Natura e NVEMAR,

Bogotá. 367 pp.

GÓMEZ, A., 1999. Los recursos marinos renovables

del Estado Nueva Esparta, Venezuela. Gómez-

Gaspar Ed. Nva. Esparta Venezuela, 215 pp.

JONES, M. L., 1962. On some polychaetous anne-

lids from Jamaica, the West Indies. Bulletin

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con 2,94%. No se registraron especies

endémicas.

Éste constituye el primer estudio sobre

los poliquetos asociados de 1. alatus.

AGRADECIMIENTOS

Los investigadores agradecen a

FONACIT, por el financiamiento parcial

de esta investigación a través del Pro-

yecto S1-2000000946.

KEOUGH, M., 1984. Dynamics of the epifauna of

the bivalve Pinna bicolor: interaction among

recruitment, predation and competition. Eco-

logy, 63 (3): 677-688.

LINERO-ARANA, I., 1996. Aspectos bioecológi-

cos de los poliquetos y descripción de algu-

nas especies bénticas de la costa nororiental

de Venezuela. Trabajo de Ascenso a Profesor Ti-

tular. Universidad de Oriente, Cumaná Ve-

nezuela, 254 pp.

LINERO-ARANA, IL, 1999. Poliquetos (Annelida:

Polychaeta) asociados al mejillón verde Perna

viridis, en la Península de Araya, Venezuela.

Boletín del Instituto Oceanográfico de Venezuela.

Universidad de Oriente-Venezuela, 38 (2): 53-62.

SALAZAR- VALLEJO, S. Y CARRERA-PARRA, L.,

1997. Taxonomía de Poliquetos. CONACYT-

ECOSUR: 64 pp.

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nomun alatus Gmelin (Bivalvia: Isognomoni-

dae) with notes on its potencial as a com-

mercial species. Bulletin of Marine Science, 30

(1): 90: 101.

TRUEMAN, E. R. Y LOWE, G. A., 1970. The effects

of temperature and littoral exposure on the

heart rate of a bivalve mollusk, Isognomun ala-

tus, in tropical condition. Composition Bio-

chemical Physiology, 38A: 555-564.

UEBELACKER, J. M., JONHSON, P. G. Y VITTOR,

B., 1984. Taxonomic guide to the polychaetes of

the northern Gulf of Mexico. Final Report to the

Mineral Management Service, contract 14-12-

001-29091. J.M. Uebelacker, P.G. Jonhson y

B. Vittor (Eds.) Barry A. Vittor y Associates,

Inc., Mobile, Alabama. Vol. I-VIL.

YONGE, C. M., 1968. Form and habit of species

of Mulleus (incluiding the “Hammer Oys-

ters”) with comparative observations on [sog-

nomun isognomun. Biological Bulletin of Marine

Biology Laboratory of Woods Hole. 135: 378-

405.

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O Sociedad Española de Malacología —___——— lberus, 21 (2): 67-90, 2003

The family Elachisinidae (Mollusca, Rissooidea) in the tem-

perate and tropical Atlantic

La familia Elachisinidae (Mollusca, Rissooidea) en el Atlántico tem-

plado y tropical

Emilio ROLÁN* and Serge GOFAS**

Recibido el 29-1-2003. Aceptado el 18-1V-2003

RESUMEN

Se revisa el género Elachisina Dall, 1918 (Mollusca, Rissooidea) en el océano Atlántico, in-

cluyendo aquellas especies que previamente habían sido situadas en otros géneros. En total,

hay 4 especies ya conocidas (E. floridana (Rehder, 1943), del Caribe, E. eritima (Smith,

1891) de la Isla de Sta. Helena, E. canarica (Nordsieck y García-Talavera, 1979) (combina-

ción nueva) de Canarias y E. canaliculata Rolán y Rubio, 2001, de las Islas de Cabo Verde),

7 que se describen como nuevas para la ciencia (E. azoreana n. sp., de las Azores, E. te-

nuisculpta n. sp., E. pergrandis n. sp., E. pelorcei n. sp., E. senegalensis n. sp., E. gubbiolii

n. sp. y E. catenata n. sp., de la costa occidental africana) y una más que figuramos sin nom-

brarla por carecer de material suficiente. Las especies insulares son todas endemismos de sus

respectivos arquipiélagos y tienen una protoconcha pauciespiral, lo que sugiere un desarro-

llo no-planctotrófico. Cuatro de las seis especies de la costa africana tienen protoconcha mul-

tiespiral y probablemente un desarrollo planctotrófico, pero solo tres de ellas se han encon-

trado en un área geográfica amplia, desde Mauritania o Senegal hasta Angola. Teniendo

en cuenta las nuevas especies aquí descritas, la provincia Oeste-Africana se presenta como

un centro de mayor riqueza específica conocido para la familia.

ABSTRACT

The Atlantic species of the genus Elachisina Dall, 1918 (Mollusca, Rissooidea), including those

hitherto placed in other genera, are revised. In total there are four previously known species (E.

foridana (Rehder, 1943), from the Caribbean, E. eritima (Smith, 1891), from St. Helena Island,

E. canarica (Nordsieck and Garcia-Talavera, 1979) (comb. nov.), from the Canaries and E. cana-

liculata Rolán and Rubio, 2001, from the Cape Verde Islands), besides seven species which are

described as new to science (E. azoreana n. sp., from the Azores, E. tenuisculpta n. sp., E. per-

grandis n. sp., E. pelorcei n. sp., E. senegalensis n. sp., E. gubbiolii n. sp. and E. catenata n.

sp. from the West African coasts) and one more species which we figure without naming it

awaiting appropriate material. The insular species are endemic of their respective archipelagoes,

and have a paucispiral protoconch that suggests non-planktotrophic development. Four out of

the six West African species have a multispiral protoconch and a probable planktotrophic deve-

lopment, but among them only three have been collected in an extensive geographical range

from Mauritania or Senegal to Angola. Taking into account the new species described herein

and current knowledge, West Africa appears as a center of high species richness for the family.

KEY WORDS: Rissooidea, Elachisina, new species, Atlantic, West Africa.

PALABRAS CLAVE: Rissooidea, Elachisina, especies nuevas, Atlántico, África occidental.

* Investigador del Museo de Historia Natural, Campus Universitario Sur, 15782, Santiago de Compostela.

** Dept. de Biología Animal, Facultad de Ciencias, Universidad de Málaga, E-29071 Málaga.

Iberus, 21 (2), 2003

INTRODUCTION

The genus Elachisina was introduced

by DaLL (1918), and the taxon received

several different interpretations revie-

wed in PONDER (1985a). In this latter

work, the anatomy of the soft parts and

the taxonomy of the known species were

detailed and the new family Elachisini-

dae was proposed. This author presen-

ted the species which he considered wit-

hin this genus, from the Caribbean, wes-

tern North and Central America, and the

Indo-Pacific. No species were reported

from the Eastern Atlantic mainland co-

ast, but Rissoa eritima Smith, 1890 from St

Helena, was included in Elachisina. Two

more species have been described more

recently and will be considered here: E.

canarica (Nordsieck and García-Talavera,

1979) from the Canary Islands, and E. ca-

naliculata Rolán and Rubio, 2001, from

the Cape Verde Islands.

From the material collected in

several expeditions to West Africa by

both authors, as well as from sediments

sent to us by amateur collectors and

from museum material, we have found

several additional species of this genus.

The present work results from the study

of this material. The type material

collected by E.R. is deposited in Museo

Nacional de Ciencias Naturales,

Madrid, and paratypes in several collec-

tions; that collected by S.G. is in the

Muséum National d'Histoire Naturelle,

Paris.

The genus has a fossil record in the

West European Tertiary, going back to the

TAXONOMIC PART

Eocene with Elachisina minutissima (Des-

hayes, 1861), Elachisina loveni (Bayan, 1873),

(see Le Renard, on-line “fossils” database

http: / /www.somali.asso.fr/fossils). Se-

veral other genera (Entomope, Cirsope, Pseu-

docirsope, Lacunella, Dumasella) are therein

accepted as Elachisinidae. Representative

species in the Neogene are Elachisina mo-

ravica (Rzehak, 1893) and Pseudocirsope

burdigalica (Cossman and Peyrot, 1919)

from the Miocene, but there are several ot-

her undescribed species (pers. comm. P.

Lozouet, MNHN).

Abbreviations

AMNH American Museum of Natural

History, New York

BMNH The Natural History Museum,

London

MNCN Museo Nacional de Ciencias

Naturales, Madrid

MNHN Muséum National d' Histoire

Naturelle, Paris

CER private collection of Emilio Rolán

CFR private collection of Federico Rubio

CFG collection Franco Gubbioli

CJP private collection of Jacques Pelorce

CPR private collection of Peter Ryall

CWE private collection of Winfried Engl

spm specimen with soft parts

s empty shell

jjuvenile

f fragment

sta. station (of a sampling cruise)

coll. in the collection of

leg. “legit”, collected by

Family ELACHISINIDAE COMPLETA AUTOR Y AÑO

Genus Elachisina Dall, 1918

Type species: Elachisina grippi Dall, 1918; by original designation. Recent, western North

America.

Elachisina floridana (Rehder, 1943) (Figs. 1-6)

Microdochus floridanus Rehder, 1943a p. 193-194, pl. 20, fig. 6 [Type locality: originally stated as

Bonefish Key, Fla. corrected to Missouri Key in REHDER 1943b].

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

Figures 1-6. Elachisina floridana (Rehder, 1943). 1: paratype from Missouri Key, Florida, leg. B.R. Bales

(MNHN; height 2.35 mm); 2: detail of the microsculpture of another paratype; 3: protoconch (same

specimen as 2); 4: shell from Treasure Cove, Abaco, Bahamas, leg. C. Redfern (height 2.45 mm); 5: detail

of the microsculpture (same shell as 4); 6: protoconch, same locality as 4, 5. Scale bars 100 um.

Figuras 1-6. Elachisina floridana (Rehder, 1943). 1: paratipo de Missouri Key, Florida, leg. B.R. Bales

(altura 2.35 mm); 2: detalle de la microescultura de otro paratipo; 3: protoconcha (mismo ejemplar que

2); 4: concha de Treasure Cove, Abaco, Bahamas, leg. C. Redfern (altura 2.45 mm); 5: detalle de la micro-

escultura (misma concha que 4); 6: protoconcha, misma localidad que 4, 5. Escalas 100 yo.

Type material: Holotype in United States National Museum, not examined, paratypes in Los Angeles

County Museum, Los Angeles; 8 spm, leg. B.R. Bales, in coll. Staadt, MNHN.

Other material examined: Bahamas: 9 s, Treasure Cove, Abaco, in beach drift, leg. C. Redfern 1981;

5 s, Treasure Cove, Abaco, in beach drift, leg. C. Redfern 1977; 2 spm collected alive, Treasure Cay,

Bahamas, under stone in 1 m depth, leg. C. Redfern 1982.

Iberus, 21 (2), 2003

Description: Shell (Figs. 1, 4) small,

ovate-conical, solid, not transparent,

with convex whorls separated by a deep,

chamnelled suture. Protoconch (Figs. 3, 6)

of a little more than one whorl with a

diameter of 280-300 pm, depressed

dome-shaped, and smooth. Teleoconch

covered by numerous spiral grooves

(Figs. 2, 5) which are about 8-10 very

faint at the beginning and, in the last

whorl, are about 30 up to the base and

become stronger abapically. Spiral groo-

ves about as wide as the interspaces, and

rather smooth. Umbilicus narrow, borde-

red by a blunt keel and with small striae

inside. Aperture piriform, with an angu-

lation in the adapical part, and with the

abapical edge slightly protruding at the

point where the umbilical keel meets the

columella. Peristome continuous, with

simple outer lip. Colour whitish. Dimen-

sions up to 2.45 mm height.

Distribution: USA: Florida Keys, Loui-

siana; Mexico; Panama; Bermuda;

Bahamas: Abaco and Grand Bahama

Island; Brazil (see details in Malacolog

database http://erato.acnatsci.org

/wasp /search.php /2200)

Remarks: This taxon was described in

a new genus Microdochus of the Rissoi-

dae, and later recognized as a species of

Elachisina by PONDER (1985a).

Elachisina eritima (E. A. Smith, 1890) (Figs. 7-10)

Rissoa eritima E. A. Smith, 1890. Proc. Zool. Soc. London,18: 289, fig. 40. [Type locality: St. Helena

island].

Type material: 5 syntypes (Fig. 7) in BMNH.

Other material examined: St. Helena: 6 s, coll. Turton (MNHN); 1 j, off Jamestown 15? 54.5' S, 5?

42 W, leg. G. Kouyoumontzakis, Orstom (MNHN).

Description: Shell (Figs. 7, 8) small,

ovate to ovate-conic, solid, opaque, with

convex whorls. Protoconch (Fig. 10) of

about one whorl and 1/8 more, light

brown in colour, depressed dome-

shaped, smooth, about 330 qm in dia-

meter with a nucleus of 125 um. Teleo-

conch covered by numerous fine spiral

grooves (Fig. 9) which are about 11-13 at

the beginning, 25 on the last spire

whorl, and about 60 on the last whorl.

Spiral grooves of similar size as inters-

paces, frequently with a thread in the

interspaces. Umbilicus narrow, bordered

by a sharp ridge and with fine commar-

ginal striae inside. Aperture almost

semicircular, with a blunt angulation in

the adapical part, and with the abapical

edge slightly protruding at the point

where the umbilical keel meets the colu-

mella. Peristome continuous, with

simple outer lip. Colour whitish.

Dimensions: between 1.3 and 1.7 mm

height.

Distribution: Only known from St.

Helena. Supposedly endemic of this

island.

Remarks: E. eritima differs from other

Atlantic species by its characteristically

globose shape.

Elachisina canarica (Nordsieck and García-Talavera, 1979) (Figs. 11-17)

Cithna tenella canarica Nordsieck and García-Talavera, 1979. p. 289, pl. 11, fig. 1. [Type locality:

Arrecife, Lanzarote 1., Canary ls.].

Type material: In Museo de la Naturaleza y el Hombre, Santa Cruz de Tenerife, not examined.

Other material examined: Canary Islands: 3 spm, 3 s, La Restinga, El Hierro, 30 m (CWE).

Description: Shell (Figs. 11, 12) small,

ovate to ovate-conic, solid, not transpa-

rent, with convex whorls. Protoconch

(Fig. 15) of a little more than one whorl

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

Figures 7-10. Elachisina eritima (E.A. Smith, 1890). 7: syntype from St Helena (BMNH); 8: shell

from St Helena, coll. Turton (MNAHN; height 1.66 mm); 9: microsculpture of the same shell; 10:

protoconch, same locality. Scale bars 100 um.

Figuras 7-10. Elachisina eritima (E.A. Smith, 1890). 7: sintipo de Santa Helena (BMNA); 8: concha

de Santa Helena, coll. Turton (MNHN; altura 1.66 mm); 9: microescultura de la misma concha; 10:

protoconcha, misma localidad. Escalas 100 um.

with a diameter of 360 ym, light brown,

depressed dome-shaped, and smooth.

Teleoconch covered by numerous spiral

grooves (Fig. 14) which are about 12-13

at the beginning and, in the last whorl,

are about 48-50 up to the base. Umbili-

cus narrow, bordered by a sharp keel

and with small striae inside. Aperture

ovate-piriform, with an angulation in

the adapical part, and with the abapical

edge slightly protruding at the point

where the umbilical keel meets the colu-

mella. Peristome continuous, with

simple outer lip. Colour whitish.

Soft parts: The dry animal is appa-

rently unpigmented with very evident

and relatively large fecal pellets in its

intestine. Operculum (Fig.13) paucispi-

ral, thin and transparent. Radula (Figs.

15, 16) as in other species of the genus

(see PONDER, 1985).

Distribution: Only known from the

Canary Islands.

Remarks: This taxon was described as

subspecies of Cithna tenella Jeffreys,

1869, but is unrelated with this species

now classified in the Rissoidae as Bent-

honella tenella (see BOUCHET AND

WARÉN, 1993: 697-701).

It most resembles E. azoreana spec.

nov. but is more solid and slender (H/D

ratio 1.5 instead of 1.4). The Caribbean

species E. floridana is larger, more

slender, with one more whorl, a more

tumid profile of the spire and a narro-

wer umbilicus.

Iberus, 21 (2), 2003

Elachisina azoreana spec. nov. (Fig. 17-19)

Type material: Holotype (Fig. 17) and 11 paratypes from type locality (UNHN).

Type locality: Vila do Porto (depth 6-9 m), Santa Maria, Azores Archipelago.

Etymology: The specific name is after the archipelago where the species was found.

Description: Shell (Fig. 17) small,

ovate to ovate-conic, solid, opaque, with

convex whorls. Protoconch (Fig. 18) of

one whorl and !/s, transparent, dome-

shaped, smooth, about 350 ym in dia-

meter with a nucleus of 109 um. Teleo-

conch covered by numerous spiral

grooves which are about 11-13 at the

beginning, 25 over the end of the spire

and about 40 on the last whorl. Spiral

grooves (Fig. 19) very shallow and

narrow, with wider interspaces, and a

small thread visible inside in some

areas. Umbilicus narrow, with a faint

rim situated well inside and terminating;

the spiral sculpture, then commarginal

striae in its innermost part. Aperture

piriform, with a blunt angulation in the

adapical part, and with the abapical

edge very slightly protruding at the

point where the umbilical keel meets the

columella. Peristome continuous, with

simple outer lip. Colour white.

Dimensions: Holotype 1.5 mm high

and 1.1 mm in diameter.

Remarks: This species most resembles

E. canarica but the latter is narrower,

more solid, a little larger, and has a

more distinct keel delimiting the umbili-

cus and protruding over the apertural

edge. Also its microsculpture has micro-

channels in the interspaces between th-

reads which are not seen in E. azoreana.

The general shell morphology, and

particularly the conformation of the

umbilical area, resemble very much E.

tenuisculpta spec. nov. The main diffe-

rence is in the paucispiral protoconch,

but also the size is smaller and the spiral

grooves are wider than in E. tenuisculpta.

Elachisina canaliculata Rolán and Rubio, 2001 (Figs. 21-25)

Elachisina canaliculata Rolán and Rubio, 2001. Novapex, 2(4): 133-136. [Type locality: Baia Teodora,

North of Sal Rei, Boavista Island].

Type material: Holotype (Fig. 22) in MNCN (15.05 /44327). Paratypes in MNHN (1), DBUA (1),

CER (1), CFR (1).

Other material studied: Cape Verde Archipelago: Sal: 1 s, Mordeira, 5 m (destroyed during the

study). Boavista: 2 s, Sal Rei, 10 m; Santiago: 1 s, Praia, 8 m.

Description: Shell (Figs. 21-23) small,

ovoid, thin, somewhat transparent, with

2-3 whorls separated by a well defined

suture. Protoconch (Figs. 23, 25) globose,

smooth, with a little more than 1 whorl,

and a maximum diameter difficult to be

sized due to the gradual transition to the

teleoconch (258 um is mentioned in the

original description). Nucleus of the pro-

toconch 137 pm in diameter. Holotype

with 3 slightly convex teleoconch whorls

sculptured with spiral grooves (Fig. 24)

which are almost canal-like, and cover

the entire shell surface (numbering

between 22-25 on the last whorl, with the

final 3-4 grooves reaching into the umbi-

lical funnel). Umbilicus is narrow, with a

faint rim situated well inside and termi-

nating the spiral sculpture. Aperture

oval, with an angulation in the adapical

part, and with the abapical edge dis-

tinctly protruding at the point where the

umbilical keel meets the columella.

Peristome continuous, with simple outer

lip. Central area of the columella curved

and reflected towards the umbilicus.

Dimensions: Holotype 1.9 mm in

height.

Remarks: The widely spaced and

sharply delimited grooves differentiate

the present species from others in the

genus Elachisina, particularly from E.

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

Figures 11-17. Elachisina canarica (Nordsieck and Garcia-Talavera, 1979). 11, 12: shells from La

Restinga, Hierro (height 1.8 and 1.5 mm); 13: operculum (maximum diametre 0.6 mm). 14:

microsculpture (same shell as 11); 15: protoconch, same locality; 16, 17: radula. Scale bars, 13-15:

100 um; 16, 17: 20 um.

Figuras 11-17. Elachisina canarica (Nordsieck y Garcia- Talavera, 1979). 11, 12: conchas de La Restinga, Hierro

(altura 1,8 y 1,5 mm); 13: opérculo (diámetro máximo 0,6 mm). 14: microescultura (misma concha que 11).

15: protoconcha, misma localidad; 16, 17: radula. Escalas, 13-15: 100 qm; 16-17: 20 jm.

Iberus, 21 (2), 2003

Figures 18-20. Elachisina azoreana spec. nov. 18: holotype from Santa Maria, Azores (MN HN; height

1.5 mm); 19: microsculpture of the holotype; 20: protoconch of a paratype. Scale bars 100 um.

Figuras 18-20. E. azoreana. 18: holotipo de Santa Maria, Azores (MNAEN: altura 1,5 mm); 19:

microescultura del holotipo; 20: protoconcha de un paratipo. Escalas 100 ym.

canarica, E. azoreana, E. pelorcei and E. described below, have a protoconch

senegalensis. Elachisina floridana is larger, with 2 whorls at least and have more

with more whorls, a more tumid profile numerous spiral grooves; the former

of the spire and a narrower umbilicus. also differs in having a nearly closed

Elachisina pergrandis and E. tenuisculpta, umbilicus in the adult stage.

Elachisina tenuisculpta spec. nov. (Figs. 26-35)

Type material: Holotype (Fig. 26) deposited in MNHN. Paratypes: AMNH (1), BMNH (1), MNCN

(1, Fig. 16, n* 15.05/46460) CER (50), CER (1), CPR (1), all from the type locality.

Other material examined: Mauritania: 1 s, Baie de Cansado, 20” 50” N, 10 m(MNHN); 20s, 103, 10

f, in intertidal sediments, Banc d'Arguin (CER); 2 s, intertidal, Baie de l'Etoile, Nouadhibou (CER).

Senegal: 5 s, Dakar, on shipwreck “Le Tacoma”, 15 m (CJP); 1 s, Grand Thiouriba, 40 m, (CJP); 1 s,

2 j, 2 f, Madeleines, Dakar, 6-14 m (CER). Guinea Bissau: 2 s, 2 f, S Ilha do Mel, Exp. “Chalgui II”

sta. 7, 10? 41” N, 15” 44.5" W, 25 m (MNHN). Guinea Conakry: 1 j, W of Ile Kabak, “Sedigui I” sta.

154, 9 18' N, 14? 00” W, 24 m (MNHN); 1 j, W Ile Tannah, “Sedigui I” sta. 80, 9 12.37 N 13? 37' W,

16 m (MNHN); 1 j, W Rio Yomponi, “Sedigui II” sta. 688, 10 24? N, 14? 50” W, 22 m (MNHN); 1 j,

W Ouendi-Taboria, “Chalgui 7” sta. 41, 9 55 N, 14? 17” W, 17 m (MNHN); 2 s (CER). Ghana: 6 s,

2 j, Miamia, 38-40 m (CER); 3 s, 22 j, 14 f, Cape Three Points, 45-60 m (CER). Gabon: 5 c, between

Mayumba and Conkouati, “Congo”, sta. 796, 17-19 m (MNHN). Angola: near Ambrizete, 7? 07/ S,

12? 21' E, 80 m (MNHN); 154 s, Praia Etambar, Luanda, beach (MNHN); 4 s, 3 j, Ilha de Luanda,

10-20 m (MNHN); 4 s, off Luanda, 80-100 m, (CER); more than 130, off Mussulo, 90-100 m (MNHN);

1 s, Santo Antonio, Benguela (MNHN); 3 s, Caotinha, Benguela, infralittoral (MNHN); 1 s, Santa

Maria, Lucira, 40 m (MNHN); 3 s, 1 j, Sáo Nicolau (MNHN); 5 s, 5j, Praia Amelia, Namibe, 40-60

m (MNHN).

Type locality: Banc d'Arguin, Mauritania.

Etymology: The specific name alludes to the very fine sculpture of the shell.

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

Figures 21-25. Elachisina canaliculata Rolán and Rubio, 2001. 21: holotype from Sal, Cape Verde

Islands (UNCN; height 1.9 mm) 22, 23: paratype, same locality (UNHN, height 1.3 mm); 24:

microsculpture (same paratype as 22, 23). 25: protoconch, same paratype. Scale bars 100 um.

Figuras 21-25. Elachisina canaliculata Rolán y Rubio, 2001. 21: holotipo de Sal, Islas de Cabo Verde

(MNCN, altura 1,9 mm) 22, 23: paratipo, misma localidad (MNAN, altura 1,3 mm); 24: microes-

cultura (mismo paratipo que 22, 23). 25: protoconcha, mismo paratipo. Escalas 100 ym.

Description: Shell (Figs. 26, 27, 30, 32,

34-35) small, fragile, ovate to ovate-conic,

with convex whorls and a deep suture.

Protoconch of the type material (Fig. 29)

with 2 whorls, dome-shaped, smooth,

with a diameter of about 340 qm and a

very small nucleus of about 35 um. Proto-

conch (Figs. 29, 31, 33) in Angolan shells

a little larger with 460 qm and between 2

1/4 and 2/2 whorls. Teleoconch of 2-2 */4

convex whorls, transparent in fresh

shells, sculptured with very numerous

weak spiral grooves (Fig. 28), which are

10-12 in the beginning of the teleoconch,

15-18 in the second whorl and about 65-

70 in the last whorl. There are about 35-40

between the suture and the insertion of

the peristome, 25-30 between there and

the umbilicus border and 8-12 inside the

umbilicus). Spiral grooves 4 times narro-

wer than interspaces, being slightly

stronger near the umbilicus, and fre-

quently very attenuated or almost disap-

peared on the upper part of the last

whorl in adult shells. Umbilicus narrow,

hardly developed in juvenile shells; with

a faint rim situated well inside and termi-

nating the spiral sculpture, then commar-

ginal striae in its innermost part. Aper-

ture ovate-piriform, with a blunt angula-

tion in the adapical part, and with the

abapical edge protruding at the point

where the umbilical keel meets the colu-

mella. Peristome continuous, with Ple

outer lip. Colour white.

Dimensions: The holotype has 2.2

mm in heigth x 1.8 mm in diameter. The

largest shell reaches 3.0 mm in height.

Distribution: Known from Maurita-

nia to Angola, having been collected

Iberus, 21 (2), 2003

Figures 26-29. Elachisina tenuisculpta spec. nov. 26: holotype from Banc d'Arguin, Mauritania

(MNHN; height 2.2 mm); 27: paratype, same locality (MNCN; height 2.0 mm); 28: microsculp-

ture of the holotype. 29: protoconch of a paratype, same locality (CER). Scale bars 100 um.

Figuras 26-29. Elachisina tenuisculpta spec. nov. 26: holotipo de Banc d"Arguin, Mauritania

(MNHN, altura 2,2 mm); 27: paratipo, misma localidad (MNCN, altura 2,0 mm); 28: microescul-

tura del holotipo. 29: protoconcha de un paratipo, misma localidad (CER). Escalas 100 yum.

from Baie de 1'Etoile in Nouadhibou to Remarks: We have found some diffe-

Banc d'Arguin, and also in Senegal, rences between the shells from Maurita-

Guinea Bissau and Angola. nia and Senegal and those from Ghana

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

Figures 30-35. Elachisina tenuisculpta spec. nov. 30: shell from Miamia, Ghana (CER, height 1.15

mm); 31: protoconch, same locality, 32: shell from Guinea Conakry, trawled 60-80 m (CER;

height 1.09 mm); 33: protoconch of another shell from Guinea Conakry; 34-35: shells from off

Luanda, 80-100 m, Angola (CER; height 1.19 and 0.86 mm). Scale bars 100 um.

Figuras 30-35. Elachisina tenuisculpta spec. nov. 30: concha de Miamia, Ghana (CER, altura 1,15

mm); 31: protoconcha, misma localidad, 32: concha de Guinea Conakry, rastreada en 60-80 m (CER;

altura 1,09 mm); 33: protoconcha de otra concha de Guinea Conakry; 34-35: conchas de frente a

Luanda, 80-100 m, Angola (CER; altura 1,19 y 0,86 mm). Escalas 100 pom.

Iberus, 21 (2), 2003

to Angola. The specimens from the type

locality are a little larger, the number of

the spiral grooves is higher and the pro-

toconch has 2 spiral whorls, whereas

those from Angola have 2 */4 or more

whorls and, as a consecuence, a larger

diameter. Anyway, shells from Senegal

also had a protoconch similar to those

from Angola, so that we considered that

these differences could correspond to

intraspecific variability of a species with

a large distribution range. Therefore, we

have included all of them in the same

taxon.

Elachisina tenuisculpta resembles E.

catenata spec. nov. which also has a mul-

tispiral protoconch and a globose shape,

but the latter differs by a characteristic

sculpture of spiral rows of pits. Elachisina

pergrandis spec. nov. is also globose and

also has a multispiral protoconch, but has

much narrower and ragged spiral furrows,

and a nearly closed umbilicus in the adult.

E. azoreana, E. canarica, E. senegalensis

spec. nov. and E. pelorcei spec. nov. are

smaller, more elongate, with fewer

spiral grooves and have a paucispiral

protoconch.

Elachisina pergrandis spec. nov. (Figs. 36-40)

Type material: Holotype (Fig. 36), deposited in MNHN and 22 paratypes: 2 s, Cabo Ledo, Luanda,

10-40 m (MNHN); 20 s, 2 j, Corimba Bay, 10-20 m (MNHN), Angola. Other paratypes: MNCN(1,

n* 15.05/46461), CER (3), from type locality.

Other material examined: Senegal: 1 s, 13? 54 N, 16” 49 W, 7 m (MNHN). Guinea Conakry: W of

Ouendi-Taboria, “Chalgui 7” sta. 41, 9 55 N, 14” 17” W (MNHN). Ivory Coast: 7 s, region of Abidjan

(MNHN). Ghana: 3 s, 10 j, Cape Three Points, 35-65 m;, 5 j, Miamia, 38-40 m;, 3 j, Miamia, 35 m; 1 s,

2, 1 f, Miamia, 45-50 m. Angola: 6 j, Ambrizete, Bango lighthouse, 07” 20.19 S, 12? 55.09 E, inter-

tidal (UMNHN); 2 j, Corimba Bay, 10-20 m (MNHN).

Type locality: Palmeirinhas, Luanda, Angola.

Etymology: The specific name alludes to being one of the largest species in the genus.

Description: Shell (Figs. 36, 37, 39)

relatively large for the genus, ovate-

conic, quite solid, with convex whorls.

Protoconch (Fig. 40) of about 2 whorls,

dome-shaped, smooth. Teleoconch of a

little more than three whorls; these

whorls are convex, shiny, opaque, and

sculptured with some weak spiral

grooves, sometimes interrupted so as to

form a dashed pattern, and separated

from each other by intervals 10 times as

large as the grooves (Fig. 38). These are

about 10 in the beginning of the teleo-

conch, 12 in the subsequent whorl and

14 on the last spire whorl. The last

whorl has about 20-22 grooves, 14 in the

subsutural area, very closely set, wheras

the abapical area has a set of deeper

furrows separating flat, blunt spiral

cords around the umbilical chink. There

is only a very narrow chink in place of

the umbilicus. Aperture ovate piriform,

with an angulation in the adapical part,

and with the abapical edge strongly pro-

truding at the abapical termination of

the columella. Peristome continuous,

with simple outer lip, slightly notched

in the adapical angle (Fig. 36-37).

Dimensions: The holotype is 4.2 mm

in heigth and 2.8 mm in width; the

largest shells reach 4.5 mm in heigth.

Distribution: Known from Ghana to

Luanda, Angola.

Remarks: This species resembles

slightly E. bakeri (Strong, 1938) but the

latter is more elongate, the sutures are

not so deep, the spiral grooves are

always more regular and the interspa-

ces not so wide in the middle of the

whorls.

Among the other East Atlantic

species with multispiral protoconch, E.

tenuisculpta spec. nov. and E. catenata

spec. nov. differ in having a definite

umbilicus, and E. gubbiolííi spec. nov. in

being more solid, more conical, with a

peripheral angulation on the body

whorl.

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

Figures 36-40. Elachisina pergrandis spec. nov. 36: holotype from Palmeirinhas, Luanda, Angola

(MNHN; height 4.2 mm); 37: detail of the aperture of the holotype; 38: microsculpture of the

holotype; 39: apical view of a paratype (CER; maximum diametre 3.0 mm); 40: protoconch, same

paratype as 39. Scale bars, 37: 1mm, 38: 100 um, 40: 200 um.

Figuras 36-40. Elachisina pergrandis spec. nov. 36: holotipo de Palmeirinhas, Luanda, Angola

(MNAN, altura 4,2 mm); 37: detalle de la abertura del holotipo; 38: microescultura del holotipo; 39:

vista apical de un paratipo (CER; diámetro máximo 3,0 mm); 40: protoconcha, mismo paratipo que

39. Escalas, 37: Imm, 38: 100 um, 40: 200 ym.

Iberus, 21 (2), 2003

Figures 41-47. Elachisina pelorcei spec. nov. 41: holotype from Petit Thiouriba, Dakar, Senegal

(MNHN; height 1.6 mm); 42: paratype from Les Madeleines, Dakar, Senegal (MNCN; height

1.7 mm); 43: paratype from Grand Thiouriba (CER; height 1.7 mm); 44: protoconch of a juve-

nile from Petit Thiouriba (CER); 45: protoconch of a shell from Grand Thiouriba (CJP); 46, 47:

microsculpture (same paratype as 42). Scale bars, 44, 45: 200 um; 46, 47: 100 um.

Figuras 41-47. Elachisina pelorcei spec. nov. 41: holotipo de Petit Thiouriba, Dakar, Senegal

(MNHN, altura 1,6 mm); 42: paratipo de Les Madeleines, Dakar, Senegal (MNCN, altura 1,7 mm);

43: paratipo de Grand Thiouriba (CER; altura 1,7 mm); 44: protoconcha de un juvenil de Petit

Thiouriba (CER); 45: protoconcha de una concha de Grand Thiouriba (C]P); 46, 47: microescultura

(mismo paratipo que 42). Escalas, 44, 45: 200 um; 46, 47: 100 ym.

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

Figures 48-52. Elachisina senegalensís spec. nov. 48: paratype from La Tacoma, Dakar (MNCN;

height 1.4 mm); 49: holotype from Cap Vert, Dakar, Senegal (UNHN; height 1.2 mm); 50-51:

microsculpture (same paratype as 48); 52: protoconch of another paratype from La Tacoma

(CER). Scale bars 100 pm.

Figuras 48-52. Elachisina senegalensis spec. nov. 48: paratipo de La Tacoma, Dakar (MNCN, altura

1,4 mm); 49: holotipo de Cap Vert, Dakar, Senegal (MNHN, altura 1,2 mm); 50-51: microescultura

(mismo paratipo que 48); 52: protoconcha de otro paratipo de La Tacoma (CER). Escalas 100 ym.

Elachisina pelorcei spec. nov. (Figs. 41-47)

Type material: Holotype (Fig. 41) deposited in MNHN (from Dakar, Petit Thiouriba, 33 m). Paraty-

pes: in AMNH (1 s, Petit Thiouriba, 33 m, exCJP); BMNH (1, Dakar, Les Madeleines, 6-14 m), UANCN

(1 s, n* 15.05/46362, Dakar, Les Madeleines, 7-13 m, exCJP), CER (1 s, Dakar, Grand Thiouriba, 40

m, 2 s, Petit Thioutiba, 33 m, exCJP; 14 s, Les Madeleines, 6-14 m); CJP (7 s, Grand Thiouriba, 40 m;

25, 1j, Dakar, Goute Teni M'Both, 25 m, 4 s, Petit Thiouriba, 33 m; 2 s, Dakar, Almadies, 20-25 m),

CFR (1, Les Madeleines, 6-14 m) and CPR (1, les Madeleines, 6-14 m).

Other material examined: Senegal: 1 s, 30 m, Les Madeleines (broken during the study); 6 s, 1 j, 1

f, Petit Thiouriba, 35 m (CER).

Type locality: Dakar, Senegal.

Etymology: The species is named after Jacques Pelorce, French malacologist who collected the first

material of this species, and donated sediments where other shells were found.

Iberus, 21 (2), 2003

Description: Shell (Figs. 41-43) very

small, ovate to ovate-conic, relatively

solid, with convex whorls. Protoconch

(Figs. 44-45) of a little more than one

smooth, shining whorl, dome-shaped in

form and with a maximum diameter of

about 280-295 um and a nucleus with

106 pm. Teleoconch with 2 or 2 !/4

convex whorls, the first with 15 spiral

grooves, and the last whorl with about

28. Spiral grooves even, with regular

interspaces (Figs. 46-47), stronger near

the abapical part and extending into the

umbilicus. Umbilicus small, bearing

inside a sharp ridge which terminates

the external spiral sculpture. Aperture

ovate-piriform, with a blunt angulation

in the adapical part, and with the abapi-

cal edge very slightly protruding at the

point where the umbilical keel meets the

columella. Peristome continuous, with

an internal thickening of the outer lip,

and a beveled edge.

Dimensions: The holotype is 1.6 mm

in height and 1.2 mm in width.

Distribution: Only known from

Dakar, Senegal.

Remarks: Elachisina azoreana, E. cana-

rica and E. tenuisculpta also have a pau-

cispiral protoconch but have more

fragile shells, with spiral sculpture more

dense and with smaller grooves and

wider umbilicus.

E. senegalensis, which is the most

similar species and lives sympatrically,

will be discussed below.

Elachisina senegalensis spec. nov. (Figs. 48-52)

Type material: Holotype (Fig. 49) deposited in MNHN (Cap Vert, Dakar, Senegal). Paratypes:

AMNH (1 s, Dakar, shipwreck of “La Tacoma”, 15 m, exCJP), BMNH (1, Dakar, Les Madeleines,

6-14 m), MNCN (1 s, n* 15.05/46463, “La Tacoma”, Fig. 48, ex CJP); in CER (1 s, “La Tacoma”; 12

s, Les Madeleines, 6-14 m); in CJP (2 s, “La Tacoma”, 15 m, 3 s, Les Madeleines, 7-13 m, ), CER (1,

Les Madeleines, 6-14 m), and CPR (1, Les Madeleines, 6-14 m).

Other material examined: Senegal: 1 s (broken during the study), “La Tacoma”, 15 m; 4 f, Les Made-

leines, 6-14 m (CER); 1 s, Dakar, on shipwreck, 12 m (CER); 2 j, Dakar, Tiwe, 35 m, (CER).

Type locality: Dakar, Senegal.

Etymology: The species is named after the area where it was collected.

Description: Shell (Figs. 48-49) very

small, ovate to ovate-elongate, fragile,

with convex whorls. Protoconch (Fig.

52) of a little more than one whorl,

smooth and shiny, dome-shaped in

form, and a maximum diameter of

about 270 um. Teleoconch with 2 or 2

1/4 convex whorls, the first with about

12 spiral grooves, the last whorl with

about 33. Spiral grooves (Figs. 50, 51)

even, with regular interspaces, a little

stronger towards the abapical part and

extending into the umbilicus. Umbili-

cus small, bearing inside a sharp ridge

which terminates the external spiral

sculpture, and with fine commarginal

striae further inside. Aperture ovate-

piriform, with an angulation in the ada-

pical part, and with the abapical edge

protruding at the point where the

umbilical keel meets the columella.

Peristome continuous, with simple

outer lip. Columella slightly curved, the

columellar edge of the aperture tending

to separate from the preceding whorl in

adults.

Dimensions: Holotype is 1.2 mm in

heigh and 0.8 mm in width. Some shells

can get 1.4 mm of maximum dimen-

sion.

Distribution:

Dakar, Senegal.

Remarks: Elachisina canarica and E.

azoreana are a little larger and more

solid; also they have the protoconch

wider, and have more spiral grooves. E.

pelorcei is larger, more solid, and

globose, with wider grooves, the umbili-

cus is more definite and the body whorl

does not tend to separate from the pre-

vious whorl. Both species are found

together, without intergrades.

Only known from

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

Figures 53-56. E. gubbiolii spec. nov. 53: holotype from Dahkla, Sahara (MNCN; height 5.4

mm); 54: side view of the aperture of a paratype (CER; scale bars 1 mm); 55: protoconch of a

paratype (MNHN; scale bar 200 um); 56: microsculpture of the holotype. Scale bars 100 um.

Figuras 53-56. E. gubbiolii spec. nov. 53: holotipo de Dabkla, Sahara (MNCN,; altura 5,4 mm); 54:

vista lateral de la abertura de un paratipo (CER; barra de escala 1 mm); 55: protoconcha de un para-

tipo (MNAEN,; barra de escala 200 um); 56: microescultura del holotipo. Escalas 100 ym.

Iberus, 21 (2), 2003

Elachisina gubbiolii spec. nov. (Figs. 53-56)

Type material: Holotype (Fig. 53) deposited in MNCN (n* 15.05/46464). Paratypes: in MNHN (1,

Fig. 55), CER (1, Fig. 54) and CFG (4), all from type locality and ex CFG.

Other material examined: Only known from the type material.

Type locality: Dakhla, Sahara, 50-60 m.

Etymology: The specific name is after Franco Gubbioli, of Malaga, Spain, who provided the mate-

rial of this species.

Description: Shell (Fig. 53) relatively

large for the genus, ovate-conic, solid, with

early spire whorls convex, the penultimate

whorl only slightly convex and the body

whorl bluntly angled at the periphery. Pro-

toconch (Fig. 55) of 2 whorls, dome-

shaped, smooth, with a small nucleus of

about 56 qm and a diameter of about 375

pm. Teleoconch of about 4 whorls sculp-

tured with weak spiral grooves, the first

with about 5, stronger and wider grooves,

the following with 6 grooves, then with

an increasing number of grooves in the

following. Spiral grooves on the first tele-

oconch whorl crossed by some folds

which parallel the prosocline growth lines;

spirals becoming weaker and narrower

(about 1/15 of intervals) on the last whorl.

There are about 32 grooves, 16 of them

between the suture and the insertion of

the aperture, and the remainder abapically

to these; both groups separated by a

smooth band, which is coincident with a

weak peripheral angle. Spiral grooves

fainter near the umbilicus, and are sepa-

rated by larger interspaces in the upper

part of the whorls (Fig. 56). Umbilicus

narrow and deep, bordered by a sharp

ridge overhanging an inner furrow. Aper-

ture rhomboid, with a sharp angle in the

adapical part, and with the abapical edge

sharply protruding and notched at the ter-

mination of the columella (Fig. 54). Colu-

mella strongly twisted, almost vertical on

its lower part.

Dimensions: The holotype is 5.4 mm

in heigth and 3.5 mm in width. The

largest shells can reach 7 mm in

maximum dimension.

Distribution: Only known from the

type locality.

Remarks: E. gubbiolíí spec. nov. is dis-

tinctive, and differs from the other living

species of the genus by its larger size, the

peripherical angulation, the suture less

deep and, over all, by its very protruding

columella and basal notch similar to a si-

phonal canal. It resembles the fossil species

Pseudocirsope burdigalica (Cossman and

Peyrot, 1919), from the Miocene of Aqui-

taine Basin, France but the latter differs in

a definitely stouter outline.

Elachisina catenata spec. nov. (Figs. 57-63)

Type material: Holotype (Fig. 57) and 10 paratypes from the type locality, deposited in MNHN.

Other paratypes: in MNCN (1, n? 15.05/46465, from Cacuaco, Angola); from Miamia, Ghana: in

AMNH (1), CER (6, Figs. 58, 59), CER (1) and CPR (1).

Other material examined: Ivory Coast: 2 s, Abidjan (MNHN). Ghana: 32 s, 28 j, 25 f, Cape Three

Points, 35-65 m (CER); 9 s, 30 j, 8 f, Miamia, 45-50 m (CER); 1 s, Miamia, 35-40 m (CER). Angola: 1

s, Ambrizete, 07” 00' S, 12” 20' E, 60 m (MNHN); 4 s, Luanda, Corimba Bay, 10-20 m (MNHN); 1 j,

off Mussulo, 50-70 m (MNHN); 10 j, 4 f, off Mussulo, 90-100 m (MNHN); 1 s, Palmeirinhas, 10 m.

Type locality: Mussulo, Luanda province, Angola.

Etymology: The specific name alludes the spiral sculpture which appears like formed by chains.

Description: Shell (Figs. 57-59) small,

fragile, ovate to ovate-conic, umbilicate.

Protoconch (Figs. 60-61) of a little more

than 2 whorls which are dome-shaped

and smooth, with 396 qm of diameter

and a nucleus of 54 qm. Teleoconch of

about 2-2 */2 whorls, uniformly convex,

with deeply impressed suture, sculptu-

red with numerous, weak, spiral grooves.

In first whorl there are 7 rows of irregu-

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

Figures 57-63. Elachisina catenata spec. nov. 57: holotype from Mussulo, Angola (UNHN; height

2.1 mm); 58: paratype from Miamia, Ghana (CER; height 1.3 mm); 59: paratype, same locality

(CER; maximum diametre 1.8 mm); 60: protoconch of another paratype from Miamia; 61: proto-

conch (same paratype as 59). 62: microsculpture (same paratype as 58); 63: microsculpture of the

holotype. Scale bars 100 um.

Figuras 57-63. Elachisina catenata spec. nov. 57: holotipo de Mussulo, Angola (MNHN, altura 2,1 mm);

58: paratipo de Miamia, Ghana (CER; altura 1,3 mm); 59: paratipo, misma localidad (CER; diámetro

máximo 1,8 mm); 60: protoconcha de otro paratipo de Miamia; 61: protoconcha (mismo paratipo que

59); 62: microescultura (mismo paratipo que 57); 63: microescultura del holotipo. Escalas 100 yum.

Iberus, 21 (2), 2003

Figures 64-66. Elachisina sp. 64-65: shell from Palmeira, Cape Verde Islands (CER; height 1.83

mm); 66: protoconch. Scale bar 100 um.

Figuras 64-66. Elachisina sp. 64-65: concha de Palmeira, Islas de Cabo Verde (CER; altura 1.83 mm);

66: protoconcha. Escala 100 yum.

lar, discrete pits (Figs. 62, 63); there are

about 30 spiral grooves in the last whorl,

and the pits become sunken inside the

grooves. Umbilicus deep, not very wide,

with fainter spiral grooves extending

inside; the innermost part fluted by com-

marginal striae. Aperture ovate piriform,

with a blunt angulation in the adapical

part, and with the abapical edge hardly

protruding at the point where the colu-

mella terminates. Peristome continuous,

slightly prominent in the abapical part.

Dimensions: The holotype is 2.13 mm

in heigth by 2.00 mm in diameter. The

material from Ghana is of smaller size, with

about 1.2-1.4 mm of maximum dimensions.

Distribution: Known from the Ivory

Coast and Ghana to Angola.

Remarks: The present species differs

from those previously mentioned by its

globose form, by the absence of an umbi-

lical keel, and especially by its distinctive

sculpture of spiral discontinuous grooves

formed by small pits.

Elachisina sp. (Figs. 64-66)

Material examined: 1 s, dredged in Palmeira, 30 m.

Description: Shell (Figs. 64, 65)

small, ovate to ovate-conic, narrowly

umbilicate, whorls convex sculptured

with numerous, weak, spiral

grooves. Protoconch (Fig. 66) with a

little more than one whorl, dome-

shaped, with a prominent cord

running from the nucleus, and

several ones which appear immedia-

tely after. Teleoconch similar to other

of the genus.

Dimensions: 1.4 in heigth and 1.0

mm in width.

Distribution: Only known from

Cape Verde archipelago, from where it

is probably an endemic species.

Remarks: The sculpture of the proto-

conch separates this species from the

other within the genus, which always

have a smooth surface. We prefer not to

name this species, considering that only

a single, damaged shell is available.

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

Table I. Main characters for differenciation between the species presented in this work. Canal.:

canaliculate.

Tabla 1. Principales caracteres para la diferenciación entre las especies estudiadas en este trabajo. Canal. :

acanalada.

S S y $

9 S SAM E

Protoconch WAN A 2 UV A A A

whorls brown brown

Protoconch 0 0 0 0 0

sculpture

pm of the IZAN 109 120

nudleus of

protoconch

pm on the IIS 0 350

protoconch

Teleoconch canal. — simple

grooves witha with

ihread several

inside threads

Number of striae 15 25 26 30 12

in last whorl up

to insertion of

aperture

Shell robust — robust

Maximum size 2.45 1.7 1.8 1,5 2.2

in mm

DISCUSION AND CONCLUSIONS

In the present work 12 species of the

genus Elachisina are recorded from the

Atlantic Ocean, of which only four were

previously known: Elachisina floridana is

the species present in the West Atlantic

coast; E. eritima from Santa Helena

Island is the only one known since the

19th century; E. canarica from the Cana-

ries and E. canaliculata from the Cape

Verde archipelago were described more

recently. Seven species are described in

the present work as new for science, and

one more is kept unnamed until better

material is obtained.

Three more species of the Atlanto-

Mediterranean area have been sugges-

ted as belonging to Elachisina, or the Ela-

chisinidae. One is Elachisina versiliensis

0 0 0 0 0 0

34 40 106 85 56 54 110

258 337-460 426 290 269 375 39% 39

simple simple canal. simple irregular simple simple simple formed simple

very Very

Narrow Narrow

very — by pits

narrow

40 26 1EZO 16 195620

robust medium medium fragile medium robust medium robust fragile fragile

A E LA le 87 UA] 1.3

Warén, Carrozza and Rocchini, 1990, a

junior synonym of Laeviphitus verduini

van Aartsen, Bogi and Giusti, 1989. This

species has a strongly cancellate proto-

conch (see BOUCHET AND WARÉN, 1993:

705) reminiscent of the Nystiellinae

(Epitoniidae) and very different from

the planktotrophic protoconch type

found in the Atlantic species of Elachi-

sina. Moreover, it lacks the characteristic

ridge bordering the umbilicus in all the

species we have seen. For these reasons

Laeviphitus was suggested in the original

publication as a member of the Epitonii-

dae. This was rebutted by OKUTANI, FU-

JIKURA AND SASAKI (1993) who described

an aditional species from a bathyal site

off Japan and found a taenioglossate ra-

dula. WARÉN AND BOUCHET (2001) des-

cribed another species L. desbryueresi

Iberus, 21 (2), 2003

Table II. Distribution of the species of Elachisina from West African coast. Car: Caribbean; Azo:

Azores; Can: Canary; CV: Cape Verde Islands; SH: Santa Helena; Sah: Sahara; Mau: Mauritania;

Sen: Senegal; GC: Guinea Conakry; Gha: Ivory Coast and Ghana; Gui: Gulf of Guinea; Ang;

Gabon and Angola.

Table 11. Distribución de las especies de Elachisina en la costa occidental africana. Car: Caribe; Azo:

Azores; Can: Canarias; CV: Cabo Verde; SH: Santa Helena; Sah: Sáhara; Mau: Mauritania; Sen:

Senegal; GC: Guinea Conakry; Gha: Costa de Marfil y Ghana; Gui: Golfo de Guinea; Ang: Gabón y

Angola.

Car A) Aco

E floridana E

E eritima E

E canarica Ñ

E. canaliculata

E azoreana n.sp.

E tenuisculpta n. sp.

E. pergrandis n. sp.

E. pelorcein. sp.

E. gubbioliin. sp.

E catenata n. sp.

E senegalensis n. sp.

E sp.

from the Mid Atlantic ridge and figured

the taenioglossate radula. They also for-

mally exclude it from Epitoniidae but

agree that, awaiting an anatomical

study, the systematic position of Laevip-

hitus remains uncertain.

“Cingula” globuloides Warén, 1972,

from the Boreal North Atlantic, was

transferred to Elachisina by WARÉN

(1996). This is biogeographically discre-

pant with all the other Elachisina, restric-

ted to tropical and warm temperate wa-

ters. The shell of *C.” globuloides lacks the

characteristic umbilical rim, and generic

placement is pending confirmation with

data on the living animals or radula.

Although most of the species have a

similar aspect, some characters were

found diagnostic with respect to the

other congeneric. Some of these charac-

ters are summarized in the Table 1.

The sizes of distribution ranges

appear to be very uneven among the

different species, although future

records may extend the current ranges.

The five insular species are reported

only from the respective islands or

archipelagoes and are probably

Sah Mau Sen GC Gha Gui Ang

k * * * *k

* * *

* *

endemic. From their paucispiral proto-

conch with about 1 whorl, they are infe-

rred to have non-planktotrophic larval

development. These are E. eretima from

St. Helena, E. canarica from the Canaries;

E. canaliculata and Elachisina sp from

Cape Verde archipelago; E. azoreana

from the Azores.

There are three species which appear

very localized on the mainland coast.

Elachisina gubbiolii, recorded from the

Sahara coast, can be inferred to have

planktotrophic development and its

apparently small range may be either an

artifact due to sampling bias and rarity,

or the result of ecological restriction

(VERMEI, 1989). The two sympatric

species E. pelorcei and E. senegalensis are

restricted to the Dakar area. This small

stretch of coastline is one of the few

extensive sites with a rocky shore to be

found on the West African coast, usually

overcast with sediments. In this respect,

it is ecologically an island and this is

reflected in the local species richness

and occurrence of short-range endemics

(particularly spectacular in the genus

Conus, see PIN AND LEUNG-TAK, 1995).

ROLÁN AND GOFAS: The family Elachisinidae in the temperate and tropical Atlantic

The other species (E. tenuisculpta and

E. pergrandis) are known from larger

areas along the West African coast as

can be expected from their pattern of

planktotrophic larval development.

There is nevertheless a large gap in the

Gulf of Guinea, which may reflect either

a poor sampling in that area, or an

adverse influence of the Niger Delta on

marine communities.

The distribution area of the species

is summarized in Table II.

Taking into account the new species

described herein, West Africa appears as

a center of high species richness for the

family. This point must nevertheless be

qualified, because Elachisinids are rather

featureless gastropods. Species-rich

areas such as the Indo-West Pacific may

harbour even more species, either undes-

cribed or misplaced in other molluscan

families.

The habitat and mode of life of Ela-

chisina species remains elusive. The

West African species described here

were never found alive, although some

species being represented by a rather

large number of shells, and other groups

in the same samples were represented

by a fair number of living specimens.

Judging from the observations in the

BIBLIOGRAPHY

BOUCHET, P. AND WARÉN, A., 1993. Revision of

the Northeast Atlantic bathyal and abyssal

Mesogastropoda. Bollettino Malacologico, sup-

plemento 3: 579-840.

DaLL, W. H., 1918. Changes in and additions

to molluscan nomenclature. Proceedings ot

the Biological Society of Washington, 31: 137-138.

NORDSIECK, F. AND GARCÍA-TALAVERA, F., 1979.

Moluscos marinos de Canarias y Madera (Gas-

tropoda). Aula de Cultura de Tenerife, 208

pp, 46 pls. /

OKUTANI, T., FUJIKURA, K. AND SASAKI, T., 1993.

New taxa and new distribution records of

deep-sea gastropods collected from or near

chemosynthetic communities in the Japa-

nese waters. Bulletin of the National Science Mu-

seum, Tokyo, ser. A, 19 (4): 123-143.

Pin, M. AND LEUNG-TAkK, K. D., 1995. Cones of

Senegal. La Conchiglia, supplement to issue

277: 4-55.

Caribbean, species of Elachisina may live

under rocks, in crevices. This is consis-

tent with the lack of colour pattern on

the shells of all species. In West Africa

however, the sampling of subtidal rock

was underrepresentative, because of the

lack of visibility and the lack of infras-

tructure for scuba-diving.

ACKNOWLEDGEMENTS

The authors extend thanks to the

MNHN which loaned material for our

study and the Project PARSYST which

allowed us to see the collections and biblio-

graphy in this Museum. We are grateful

to malacologists Colin Redfern, Winfried

Engl, Franco Gubbioli, Jacques Pelorce,

who sent material for study. Most SEM

photographs were taken by Jesús Méndez

of the CACTI, University of Vigo. We are

grateful to Pierre Lozouet (MNHN, Paris)

for access to relevant fossil material from

the Oligocene and Miocene of France and

for helpful advice on Tertiary elachisinids;

to Anselmo Peñas for fossil material from

La Pedrera, NE Spain.

The first author acknowledges the

grant PGIDTOOPXIS0121PR of the

Xunta de Galicia.

PONDER, W. F., 1985a. The anatomy and rela-

tionships of Elachisina Dall (Gastropoda: Ris-

soacea). Journal of Molluscan Studies, 51: 23-

34.

PONDER, W. F., 1985b. A review of the genera

of Rissoidae (Mollusca: Mesogastropoda:

Rissoacea). Records of the Australian Museum,

suppl. 4: 1-221.

REHDER, H. A., 1943a. New marine mollusks

from the Antillean region. Proceedings of the

United States National Museum, 93 (3161): 187-

203, pls. 19-20.

REHDER, H. A., 1943b. Corrections and ecolo-

gical notes on some recently described Flo-

rida marine shells. The Nautilus, 57 (1): 32-33.

SMITH, E. A., 1890. Report on the Marine Mo-

lluscan Fauna of St. Helena. Proceedings of

the Zoological Society, 18: 247-317, pls. 21-24.

Iberus, 21 (2), 2003

VAN AARTSEN, J.J., BOGI, C. AND GIUSTI F., 1989. WARÉN, A. AND BOUCHET, P., 2001. Gastropoda

Remarks on the genus Benthonella (Rissoi- and Monoplacophora from hydrothermal

dae) in Europe, and the description of Lae- vents and seeps new taxa and records. The Ve-

viphitus (nov.gen.) verduini (nov.spec.) (Epi- liger, 44 (2): 116-227.

toniidae). La Conchiglia, 246-249: 19-22. WARÉN, A., CARROZZA, F. AND ROCCHINI, R.,

VERMEN, G.]., 1989. Geographical restriction as 1990. Elachisina versiliensis, a new Medite-

a guide to the causes of extinction: the case rranean species of the family Elachisinidae

of the cold northern oceans during the Neo- (Prosobranchia, Truncatelloidea). Bollettino

gene. Paleobiology, 15: 335-356. Malacologico, 25: 335-339.

WARÉN, A., 1996. New and little known mo-

llusca from Iceland and Scandinavia. Part 3.

Sarsia, 81: 197-245.

O Sociedad Española de Malacología

Iberus, 21 (2): 91-98, 2003

Description of a new muricopsine species (Gastropoda:

Muricidae) from the Southwestern Indian Ocean

Descripción de una nueva especie de muricopsine (Gastropoda:

Muricidae) del suroesta del Océano Índico

Roland HOUART*

Recibido el 10-11-2003. Aceptado el 24-IV-2003

ABSTRACT

Favartia marianae n. sp. is described from Zululand, South Africa with range extension to

South Mozambique. lt is compared with Favartia maculata (Reeve, 1845), F. cecalupoi

Bozzetti, 1993, F. jeanae Bertsch and D'Attilio, 1980, and F. conleyi Hovart, 1999.

Murex mundus Reeve, 1849 is here proposed as a nomen dubium.

RESUMEN

Se describe Favartia marianae spec. nov. de Zululand, Suráfrica con una distribución hasta

el sur de Mozambique. La nueva especie se compara con Favartia maculata (Reeve, 1845),

F. cecalupoi Bozzetti, 1993, F. jeanae Bertsch and D'Attilio, 1980, and F. conleyi Houart,

1999. Se propone que el taxon Murex mundus Reeve, 1849 sea considerado nomen dubium.

KEY WORDS: Gastropoda, Muricidae, Muricopsinae, Southwestern Indian Ocean, Favartia n. sp.

PALABRAS CLAVE: Gastropoda, Muricidae, Muricopsinae, suroeste del Océano Índico, Favartia spec. nov.

INTRODUCTION

Since both reviews of Muricidae by

FAIR (1976) and by RADWIN and D'AT-

TILIO (1976), several muricids from Zul-

uland (South Africa) and Mozambique

have been described by VOKEs (1978),

HOUART (1986, 1990, 1994, 1995, 1998,

1999) and by PONDER and VOKES (1988).

Other new discoveries have also

extended the geographical distribution

of many species originally known from

South Africa or from other localities

throughout the Indian Ocean, to

Mozambique (unpublishea).

Another small muricid species

occurring off Zululand and Mozam-

bique, sent to me for identification,

remained unidentified in one of my

drawers for a couple of years. New

material obtained recently allowed a

better comparison with other species

and has led to its description as a new

species.

Abbreviations:

BM (NH): Natural History Museum,

London, U.K.

IRSNB: Institut royal des Sciences

naturelles de Belgique, Bruxelles,

Belgium.

MNHN: Muséum national d'Histoire

naturelle, Paris, France.

Iberus, 21 (2), 2003

Table I. Terminology employed in the present paper with acronyms used in the descriptions (after

MERLE, 2001): see Figures 1-3.

Tabla 1. Terminología empleada en el presente trabajo con los acrónimos utilizados en las descripciones

(según MERLE, 2001): véanse las Figuras 1-3.

SHOULDER

IP infrasutural primary cord (primary cord on shoulder)

adis adapical infrasutural secondary cord (adapical to IP)

abis abapical infrasutural secondary cord (abapical to IP -between IP and P1-)

CONVEX PART OF THE TELEOCONCH WHORL AND SIPHONAL CANAL

P] shoulder primary cord

P2-P6 primary cords of convex part of teleoconch whorl

s]-56 secondary cords

tad tertiary adapical cord

tab tertiary abapical cord

ADP adapical siphonal primary cord

MP median siphonal primary cord

APERTURE

1D infrasutural denticle

D1-05 denticles of the convex part of the teleoconch whorl

NM: Natal Museum, Pietermaritzburg, The acronyms are occasionally put

South Africa. between parentheses, meaning that the

Terminology: full list given in Figures 1- character was observed in a few cases

3 and Table 1. but not in all specimens.

SYSTEMATICS

Family MURICIDAE Rafinesque, 1815

Subfamily MURICOPSINAE Radwin and D'Attilio, 1971

Genus Favartia Jousseaume, 1880

Type species, by original designation: Murex breviculus Sowerby, 1834, Recent, Indo-West Pacific.

Favartia marianae n. sp. (Figs. 1, 5-8)

Type material: South Africa, N Zululand, off Jesser Point, 27*35.0' S, 32*41.8' E, 70 m, dredged

Meiring Naudé, 9.V1.87, holotype NMSA D8542/T1937.

Paratypes: South Mozambique, between Quissico and Zavora Point, 90-120 m, 2 C.P. Fernandes;

85-95 m, 1 R. Houart; 75-145 m, 1 Institut royal des Sciences naturelles de Belgique IG 29829 /515;

1 MNHN; 2]. Rosado, 3 R. Houart.

Type locality: South Africa, N Zululand, off Jesser Point, 27*35.0' S, 32%41.8' E, 70 m.

Distribution: North Zululand (South Africa) to South Mozambique, living at 70-90 m.

Etymology: This new species is named for Mariana, granddaughter of César P. Fernandes (Cascais,

Portugal).

Description: Shell small for the genus, late, lightly spinose. Spire high with

up to 11.17 mm in length (paratype C.P. 1.25-1.5 protoconch whorls (1.5 whorls

Fernandes) (holotype 7.15 mm), lanceo- in holotype) and up to 5 convex, weakly

HOUART: Á new muricopsine species from the Southwestern Indian Ocean

Figures 1-3. Spiral sculpture. 1: Favartia marianae n.sp.; 2: E conleyí Houart, 1999; 3: E cecalupoí

Bozzetti, 1993. Figures 4, 5. Protoconchs. 4: K jeanae Bertsch and D'Attilio, 1980, Sulu Sea,

Philippine Islands. Coll. R. Houart. Scale bar: 0.2 mm; 5: E marianae n.sp., South Mozambique,

between Quissico and Zavora Point, 90-120 m, paratype coll. R. Houart. Scale bar: 0.5 mm.

Figuras 1-3. Escultura espiral. 1: Favartia marianae spec. nov.; 2: E. conleyi Houart, 1999; 3: E.

cecalupoi Bozzetti, 1993. Figuras 4, 5. Protoconchas. 4: E. jeanae Bertsch y D'Attilio, 1980, Sulu Sea,

Philippine Islands. Coll. R. Houart. Scale bar: 0,2 mm; 5: E marianae n.sp., South Mozambique,

entre Quissico y Zavora Point, 90-120 m, paratipo coll. R. Houart. Escala gráfica: 0,5 mm.

Iberus, 21 (2), 2003

shouldered teleoconch whorls. Suture

impressed. Protoconch large and broad;

whorls rounded, smooth, glossy; termi-

nal varix thin, raised, weakly curved.

Axial sculpture of teleoconch whorls

consisting of moderately high, weakly

spinose varices, each with short, broad,

primary and secondary. spines. First

whorl with 6 or 7 varices (7 in holotype),

7 on second, 7 or 8 on third (8 in holo-

type), 6-8 on penultimate (6 in holo-

type), 5 on last whorl. Other axial sculp-

ture of low growth lamellae, forming

small scales on spiral cords.

Spiral sculpture of strong, low,

weakly squamose cords. First whorl

with visible P1-P2, occasionally starting

s1, second with P1, s1, P2, third and

fourth with adis, IP, abis, P1, s1, P2, P3

(P3 partly covered by next whorl), last

whorl with adis, IP, abis, P1, (tad), sl,

(tab AEZ ALS ARA ESAS OSO AUDE,

(MP). P1-P5, s1, and ADP ending as

short open spines on axial varices; sl

similar to primary cords; P6 strongly

reduced.

Aperture small, broadly ovate. Col-

umellar lip narrow, flaring, smooth,

occasionally with a small narrow knob

abapically, rim partially erect, adherent

at adapical extremity. Anal notch

shallow, broad. Outer apertural lip

weakly erect with weak denticles

within: ID obsolete or very shallow, D1-

D5 increasing in strength abapically.

Siphonal canal short, broad, dorsally

bent at tip, open.

Shell white, salmon or light orange,

axial ribs and siphonal canal paler in

coloured specimens; inside of aperture

white, light pink or light salmon. Oper-

culum and radula unknown.

Remarks: Favartia maculata (Reeve,

1845) (Figs. 9-11), known from through-

out the Indo-Pacific, differs in having

more indented whorls and a different

spiral sculpture morphology, the last

teleoconch whorl of F. maculata having

P1 and P2 never separated by a strong

s1; however the presence of a more or

less high s2 is observed in some speci-

mens. The convex part of the last teleo-

conch whorl of F. maculata usually

having following sculpture: P1, P2, (s2),

P3, (s3), P4, P5 (P6 absent or strongly

reduced). Moreover, the protoconch of F.

maculata is conical with 3.5 whorls and a

sinusigera type terminal varix.

F. cecalupoi Bozzetti, 1993 (Figs. 3,12)

has a broader last teleoconch whorl com-

pared to the previous whorls, lower axial

ribs, stronger primary spiral cords, and a

broader aperture. It also differs in having

higher, almost similar P1-P5. The spiral

sculpture morphology of the last teleo-

conch whorl in F. cecalupoi being; adis, IP,

(Right page) Figures 6-8. Favartia marianae n.sp.; 6: from South Africa, N. Zululand: off Jesser

Point, 27%35.0” S, 32%41.8” E, 70 m, holotype NM D8542/ 11937, 7.15 mm; 7-8: Mozambique,

between Quissico and Zavora Point, 90-120 m, paratype coll. C.P. Fernandes, 11.17 mm. Figures

9-11. Favartia maculata (Reeve, 1845). 9: Holotype of Murex salmonea Melvill and Standen,

1899, Torres Strait, Queensland, Australia, BM (NH) 1899.2.23.24, 12 mm; 10: shell from South

Africa, N Zululand, off Kosi River Mouth, 26%53.9” S, 32%55.5” E, 50 m, NM D6855, 18.9 mm;

11: syntype of Murex maculatus Reeve, 1845, unknown locality, BM (NH) 1972020, 16.1 mm.

Figure 12. Favartia cecalupoi Bozzetti, 1993, off Ras Hafun, 150 km south of Guarda Fadui,

Northeastern Somalia, 200-250 m, holotype IRSNB 27882/455, 14 mm.

(Página derecha) Figuras 6-8. Favartia marianae spec. nov.; 6: de Suráfrica, N. Zululand: fuera de

Jesser Point, 27%35.0" S, 32*%41.8” E, 70 m, holotipo NM D8542/ 11937, 7,15 mm; 7-8: Mozam-

bique, entre Quissico y Zavora Point, 90-120 m, paratipo col. C.P. Fernandes, 11,17 mm. Figuras 9-

11. Favartia maculata (Reeve, 1845). 9: holotipo de Murex salmonea Melvill y Standen, 1899, Torres

Strait, Queensland, Australia, BM (NH) 1899.2.23.24, 12 mm; 10: concha de South Africa, N Zulu-

land, fuera de Kosi River Mouth, 26"53,9” S, 32%55,5” E, 50m, NM D6855, 18,9 mm; 11: sintipo

de Murex maculatus Reeve, 1845, localidad desconocida, BM (NH) 1972020, 16,1 mm. Figura 12.

Favartia cecalupoi Bozzetti, 1993, fuera de Ras Hafun, 150 km al sur de Guarda Fadui, nordeste de

Somalia, 200-250 m, holotipo IRSNB 27882/455, 14 mm.

HOUART: A new muricopsine species from the Southwestern Indian Ocean

Iberus, 21 (2), 2003

abis, P1, P2, (s2), P3, (s3), P4, (s4), P5, P6

(reduced or absent), ADP, MP.

F. jeanae Bertsch and D'Attilio, 1980

(Figs. 4, 13-15) is more slender with a

higher spire, stronger axial ribs, and

conical multispiral protoconch consist-

ing of 3-3.5 whorls, ending with a

sinusigeral type terminal varix (Fig. 4).

F. jeanae resembles species currently

included in Caribiella Perriliat, 1972 from

Tropical America. However, I am of the

opinion that both taxa, Favartia and

Caribiella, are congeneric.

F. conleyi Houart, 1999 (Figs. 2, 17-19)

described from Guam but also occurring

in New Caledonia and the Society

Islands, differs in having a shell with

more shouldered wwhorls, more

squamose spiral cords, and a shorter

siphonal canal, strongly recurved at tip.

F. conleyi also has a different spiral

ACKNOWLEDGEMENTS

My deepest thanks to C. P. Fernandes,

Cascais, Portugal, J. Rosadoi, Almada,

Portugal, and R. N. Kilburn, Doctor

Emeritus at the Natal Museum for the

loan and gift of material, to J. Cilis, Insti-

BIBLIOGRAPHY

Farr, R. H., 1976. The Murex Book, an illustrated

catalogue of Recent Muricidae (Muricinae, Muri-

copsinae, Ocenebrinae). Sturgis Printing Co.,

Honolulu, Hawaii, 138 pp.

sculpture morphology in having IP, P1-

P5, (P6), ADP. Last whorl occasionally

with narrow sl and s2; IP, P1-P5 and

ADP strong, high; P6 reduced or absent.

The holotype of Murex mundus

Reeve, 1849 [new name for M. exiguus

Reeve, 1849 (not Broderip, 1833) ] also

resembles the new species. Although

this unique type specimen can easily be

ascribed to the genus Favartia, it is

beachworn, probably subadult, and

lacks the protoconch and the first teleo-

conch whorl. The actual identity of

Favartia munda (Reeve, 1849) remains

thus uncertain - it has been considered a

synonym of F. pelepili D'Attilio and

Bertsch, 1980 by VOKEs (1985) - and is

here maintained as a nomen dubium. 1

cannot apply to the new species that is

not known from the intensively

explored Philippines.

tut royal des Sciences naturelles de Bel-

gique, for SEM photograph of the proto-

conch of F. jeanae, and to John Wolff (Lan-

caster, USA) for his comments and cor-

rections of the English text.

HOUARrTr, R., 1986. Chicoreus (Naquetia) triqueter

vokesae subsp. nov., a new name for a misiden-

tified species (Gastropoda: Muricidae). Apex,

1 (3): 95-96.

(Right page) Figures 13-15. Favartia jeanae Bertsch and D'Attilio, 1980. 13, 14: Holotype of Murex

pumilus A. Adams, 1853 (not M. pumilus Broderip, 1833), China Seas, holotype BM (NH)

197466, 8.1 mm; 15. shell from Philippine Islands, Cebu, Punta Engaño, 110 m, coll. R. Houart,

9 mm. Figure 16. Murex mundus Reeve, 1849, Philippine Islands, holotype BM (NH) 1972018,

11 mm. Figures 17-19. Favartia conleyí Houart, 1999. 17: Guam, Pity Lagoon, among silty dead

coral, 1.5-2.5 m, holotype MNHN, 15.2 mm; 18-19: New Caledonia, 19%08” S, 16329” E, 65-

120 m, MNHN, 11.9 mm.

(Página derecha) Figuras 13-15. Favartia jeanae Bertsch and D'Attilio, 1980. 13, 14: holotipo de

Murex pumilus A. Adams, 1853 (not M. pumilus Broderip, 1833), mares de China, holotipo BM

(NA) 197466, 8,1 mm; 15: concha de Islas Filipinas, Cebu, Punta Engaño, 110 m, col. R. Houart, 9

mm. Figura 16. Murex mundus Reeve, 1849, Islas Filipinas, holotipo BM (NH) 1972018, 11 mm.

Figuras 17-19. Favartia conleyi Houart, 1999. 17: Guam, Pity Lagoon, entre sedimentos de coral

muerto, 1,5-2,5 m, holotipo MNHN, 15,2 mm; 18-19: concha de Nueva Caledonia, 19%08” S,

16329 E, 65-120 m, MNHN, 11,9 mm.

HOUART: Á new muricopsine species from the Southwestern Indian Ocean

Iberus, 21 (2), 2003

HOUART, R., 1990. New taxa and new records

of Indo-Pacific species of Murex and Haustel-

lum (Gastropoda, Muricidae, Muricinae). Bul-

letin du Muséum national d'Histoire naturelle,

Paris, 4” sér., 12, sect A, n” 2: 329-347.

HOUART, R., 1994. Description of a new species

of Typhinae (Muricidae) from the Western In-

dian Ocean. Annals of the Natal Museum, 35:

153-155.

HOUART, R., 1995. The Ergalataxinae (Gas-

tropoda, Muricidae) from the New Caledo-

nia region with some comments on the sub-

family and the description of thirteen new

species from the Indo-West Pacific. Bulletin

du Muséum national d'Histoire naturelle, Paris,

4e sér., 16, section A, n? 2-4: 245-197.

HOUART, R., 1998. Description of eight new

species of Muricidae (Gastropoda). Apex, 13

(3): 95-109.

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ternal denticles of the outer lip in the Muri-

cidae: terminology and methodological com-

ments. Novapex, 2 (3): 69-91.

PONDER, W. F. AND VOKES, E. H., 1988. Revision

of the Indo-West Pacific fossil and Recent

species of Murex s.s. and Haustellum (Mol-

lusca: Gastropoda: Muricidae). Records of the

Australian Museum, suppl. 8: 1-160.

RADWIN G. AND D'ATTILIO, A., 1976. Murex

shells of the world. An illustrated guide to the Mu-

ricidae. Stanford University Press, Stanford.

284 pp.

VOKESs, E. H., 1978. Muricidae (Mollusca: Gas-

tropoda) from the eastern coast of Africa.

Annals of the Natal Museum, 23 (2): 375-418.

VOKES, E. H., 1985. Those amazing Mactan mu-

ricids (and some of their friends). Concholo-

gists of America Bulletin, 13 (2): 29-34.

O Sociedad Española de Malacología

Iberus, 21 (2): 99-107, 2003

A new species of Phyllomeniidae (Mollusca Solenogastres:

Sterrofustia) from the South Shetland Islands, Antarctica

Una nueva especie de Phyllomeniidae (Mollusca Solenogastres:

Sterrofustia) de las Islas shetland del sur, antártida

Óscar GARCÍA-ÁLVAREZ* and Victoriano URGORRI*

Recibido el 14-X1-2002. Aceptado el 6-VI-2003

ABSTRACT

Ocheyoherpia bursata is a new species of Phyllomeniidae (Mollusca Solenogastres: Ste-

rrofustia) collected at a depth of 248 m, on a gravel bottom off Deception Island (South

Shetland Islands, Antarctica). The radula is distich, formed by pairs of hooked teeth (up to

48 pm long x 11 pm wide), each tooth with 5-6 lateral short denticles and a pair of long,

curved and fused distal denticles, these distal denticles are located in the same plane as

the radular tooth. Midgat with a pair of ventrolateral sacs posteriorly, which run ventraly

under the spawning duct. The new species is compared with other two species of the

genus. An amended diagnosis of the genus Ocheyoherpia is proposed.

RESUMEN

Ocheyoherpia bursata es una nueva especie de Phyllomeniidae (Mollusca, Solenogastres:

Sterrofustia) recogida a 248 m de profundidad en un fondo de gravas en la Isla Decep-

ción (Islas Sethland del Sur, Antártida). Rádula dística formada por pares de dientes gan-

chudos (hasta 48 ym de largo x 11 pm de ancho), cada diente con 5-6 dentículos latera-

les cortos y con un par de dentículos distales curvados y fusionados en su extremo distal,

estos dentículos distales están situados en el mismo plano que el diente radular. El intes-

tino medio presenta posteriormente un par de bolsas ventrolaterales situadas ventralmente

al conducto de desove. La nueva especie se compara con las otras dos del género. Se

propone una enmienda a la diagnosis del género Ocheyoherpia.

KEY WORDS: Ocheyoherpia bursata, Solenogastres, South Shetland Islands, Antarctica.

PALABRAS CLAVE: Ocheyoherpia bursata, Solenogastros, Islas Shetland del Sur, Antártida.

INTRODUCTION

During a sampling programme

under the BENTART'95 project, a speci-

men was collected off Deception Island

(South Shetland Island, Antarctica) that

proved to belong to a new species of

Solenogatres. Under the current classifi-

cation, the Solenogastres are grouped

into four orders based on the type of the

mantle sclerites. The mantle, in the

orders Pholidoskepia Salvini-Plawen,

1978, and Neomeniamorpha Pelseneer,

1906, has scales, whereas in the orders

* Departamento de Bioloxía Animal, Facultade de Bioloxía, Universidade de Santiago de Compostela, E-15782

Santiago de Compostela. España. E-mail: baoscarOusc.es; bavitucoOusc.es

Iberus, 21 (2), 2003

Sterrofustia Salvini-Plawen, 1978, and

Cavibelonia Salvini-Plawen, 1978,

mostly acicular sclerites are found. The

order Sterrofustia, to which the species

described here belongs, is characterized

by solid acicular sclerites unlike in the

order Cavibelonia where the acicular

sclerites are hollow. The order Sterrofus-

tia comprises nine species, grouped into

three families with seven genera. As in

all the families of the class Solenogas-

tres, the three families belonging to the

order Sterrofustia are based on the com-

binations of two features: the radula

type and the ventral foregut glandular

organs. The family Phyllomeniidae

Salvini-Plawen, 1978, to which this new

species belongs, is characterized by a

distich radula and subepithelial ventral

foregut glandular organs (type A after

SALVINI-PLAWEN, 1978) in ampoule-

shaped, whereas the other two familiaes

have epithelial ventral foregut glandular

organs (type B after SALVINI-PLAWEN,

1978).

RESULTS

MATERIAL AND METHODS

One specimen was collected during

the Spanish campaign (BENTART'95) to

study the Antarctic benthos at 248 m depth,

at station R-22, to the south of Deception

Island (South Shetland Islands, Antarc-

tica), using a rock dredge on a gravel bot-

tom. The specimen was fixed and preser-

ved in 70% alcohol. The sclerites were stu-

died by separation of small pieces of cuticle

from the central dorsal area of the body

and from the ventral groove. These pieces

were treated with 5% sodium hypochlo-

ride for 12 h in order to isolate the scleri-

tes. The sclerites were later rinsed with

distilled water, dried under a heater at

40" and mounted with synthetic resin.

For the anatomical study, the specimen

was decalcified in ethylenediaminetetra-

cetic acid (EDTA) solution for 12 h, cut in

paraffin in a series of 10 im cross-sections.

The staining method used was Azan (af-

ter Heidenhain) and the anatomy was re-

constructed from the serial cross-sections.

Order STERROFUSTIA Salvini-Plawen, 1978

Family PHYLLOMENIIDAE Salvini-Plawen, 1978

Ocheyoherpia Salvini-Plawen, 1978

Type species: Ocheyoherpia lituifera Salvini-Plawen, 1978

Amended diagnosis: Cuticle thik or thin

with solid, acicular and hooked sclerites.

Mouth opening in the atrium, directly or

via a posterior channel without sclerites.

Radula distich, teeth with a pair of curved

and fused distal denticles. Midgut with

diverticula. Without dorso-terminal sense

organ. Unpaired genital orifice. With copu-

latory spicules and associated gland.

Without respitatory folds.

Ocheyoherpia bursata new species

Type: The holotype (cut in serial sections) is deposited in the Museo Nacional de Ciencias Natu-

rales of Madrid, number MNCN 15.02/10.

Type locality: Deception Island (station R-22-BENTART'95) South Shetland Islands, Antarctica

(637 03' 26” S; 60 39 26” W) 248 m depth.

Derivatio nominis: The specific name, bursata, refers to the midgut sacs observed in the posterior

part of the animal.

Diagnosis: Body measues 2.25 mm x

0.55 mm. Moderately thick cuticle

100

without keel, carina or protuberances.

Two types of solid oar-shaped scales,

GARCÍA-ÁLVAREZ AND URGORRI: A new species of Phyllomeniidae from the Antarctica

20 um

Figure 1. Mantle sclerites, and radular teeth of Ocheyoherpia bursata n. sp. A, F: long oar-shaped

scale; B, G: short and wide oar-shaped scale; C, H: blade-shaped acicular sclerite; D, l: Hooked

sclerite; E: blade-shaped scale from along the pedall groove; J: radular teeth

Figura 1. Escleritos del manto y diente radular de Ocheyoherpia bursata ». sp. A, E: escama larga en

forma de remo; B, G: escama corta y ancha en forma de remo; C, H: esclerito acicular en forma de hoja

de cuchillo; D, l: esclerito ganchudo; E: escama en forma de hoja de chuchillo del surco pedio; ]: diente

radular.

solid blade-shaped acicular sclerites and

solid hooked sclerites. Mouth opens

directly in the atrium. Pedal groove with

a fold that does not enter the pallial

cavity. Radula distich, teeth with 5-6

lateral denticles and one pair of curved

and fused distal denticles. Midgut with

pair of dorso-rostral caeca and posterior

pair of ventro-lateral sacs. Paired seminal

vesicles. With paired seminal receptacles.

Three pairs of copulatory spicules.

Description

General morphology: The specimen

measures 2.25 mm in length x 0.55 mm

101

Iberus, 21 (2), 2003

in width. The ends of the body are

rounded. The pedal groove is clearly

visible, with a single ciliate fold that

does not enter the pallial cavity.

Mantle: The cuticle is moderately

thick (30 to 35 ym), with epithelial papi-

llae at the base and 2-3 layers of scleri-

tes. There are two types of solid scales:

long oar-shaped scales (up to 192 qm x

19 um) (Fig. 1A, F) and short and wide

oar-shaped scales (up to 138 qm x 213

um) (Fig. 1B, G). And there are two

types of solid sclerites: blade-shaped

acicular sclerites (up to 119 um x 12.5

um) (Fig. 1C, H) and hooked sclerites

with a sharp point at the top of the hook

(o Sta 12 mu) dae 91D), 10)

Blade-shaped scales occur along the

pedal groove (up to 90 qm x 14.5 um)

(Fig. 18).

Pallial cavity: The pallial cavity is

small and lacks respiratory folds and

connects to the exterior through a

narrow ventro-terminal opening (Fig.

3A). The anus is narrow and opens into

the cavity dorsally. The unpaired genital

aperture is located in he dorso-frontal

wall of the pallial cavity.

Sense organs and nervous system: The

atrio-buccal cavity connects to the exte-

rior through a small opening. The lateral

and dorsal walls of the atrium have a

great number of individual and bifurca-

ted papillae. There is no dorso-terminal

sense organ. The cerebral ganglion is

voluminous (150 qm in width x 65 yum

in length), and is located dorsal to the

pharynx (Fig. 2A). The lateral ganglia

are small (40 um x 20 um) and are situa-

ted on either sides of the cerebral gan-

glion. Two ventral ganglia (40-45 qm in

diameter) are located latero-ventral to

the pharynx, and are found on the pos-

terior area of the pedal pit. A pair of

buccal ganglia (20 qm in diameter) are

located latero-dorsally to the pharynx.

No suprarectal commissure was located.

Digestive tract: The mouth opens into

the posterior area of the atrium (Fig.

2A). The buccal opening leads to a long

pharynx with thickened walls and

dorsal pharyngeal glands in its front

part. The radula is distich, formed by

pairs of hooked teeth (up to 48 ym long

102

x 11 um wide). Each tooth has 5-6 lateral

short denticles and a pair of long,

curved and fused distal denticles, the

distal denticles being located in the

same plane as the radular tooth (Fig. 1],

2C). The pharynx opens to a short

oesophagus, which leads to the midgut.

There is a long radular sac (60 um long),

ventral to the oesophagus. A pair of

subepithelial ventral foregut glandular

organs (type A after SALVINI-PLAWEN,

1978) are ampoule-shaped and open

laterally into the pharynx on either side

of the beginning of the radula (Fig. 2A,

C). The midgut has thick, glandular

walls with lateral constrictions due to

the dorso-ventral musculature. There is

a very wide dorso-rostral caecum which

frontally splits into two pouches (Fig.

2A, B). The posterior part of the midgut

is narrow. It is extended postero-late-

rally by in a pair of sacs (similar to those

described by HANDL (2002) in the poste-

rior body of Imeroherpia laubieri), These

run ventral to the spawning duct (Fig.

3A, C-E). Their dorsal walls have a glan-

dular appearance similar to that obser-

ved in the midgut. The rectum is narrow

and opens dorsally into the pallial

cavity.

Reproductive system: The gonads

were full of sperm and eggs. A pair of

large sperm fulled seminal vesicles lead

laterally into the gonopericardialducts,

which also contain sperm (Fig. 3A, B).

The pericardium is voluminous and also

contains sperm. The heart is located in

the dorsal wall of the pericardium. Two

short pericardialducts lead from the

posterior part of the pericardium, which

curve and turn towards the anterior part

of the pericardium, and then laterally to

lead into the mid part of the spawning

duct, when this is still unpaired (Fig.

3A, E). The spawning duct is short (225

um long), its front half is paired, its pos-

terior half is unpaired, wide and dorso-

ventrally flattened. It opens dorso-ros-

trally into the pallial cavity through an

unpaired genital pore. There are two

small seminal receptacles, which lead

into the dorso-rostral part of each spaw-

ning duct (Fig. 3A, C). There are three

pairs of copulatory spicules and one

GARCÍA-ÁLVAREZ AND URGORRI: A new species of Phyllomeniidae from the Antarctica

B, C: Microphotographs of the cross-sections of the anterior region of the body corresponding to

lines 1, 2. At: Atrial sense organ; Bg: Buccal ganglion; Cg: Cerebral ganglion; Dc: Dorsal caecum;

Lg: Lateral ganglion; Ma: Mantle; Mg: Midgut; Oe: Oesophagus; Ph: Pharynx; Pp: Pedal pit; Ra:

Radula; Rs: Radular sac; Vfg: Ventral foregut glandular organ; Vg: Ventral ganglion.

Figure 2. A. Organización esquemática de la parte anterior del cuerpo de Ocheyoherpia bursata n. sp.

B, C. Microfotografías de los cortes en sección de la región anterior del cuerpo correspondientes a las

líneas 1, 2. At: Órgano sensitivo atrial; Bg: Ganglio bucal; Cg: Ganglio cerebral; Dc: Ciego dorsal; Lg:

Ganglio lateral; Ma: Manto; Mg: Intestino; Oe: Esófago; Ph: Faringe; Pp: Foseta pedia; Ra: Rádula;

Rs: Saco radular; Vfg: Órgano glandular ventral de la faringe; Vg: Ganglio ventral.

pair of associated glands (Fig. 3A, D).

The copulatory spicules comprise two

groups of three located ventro-laterally,

each group includes a long spicule (ap-

proximately 475 um), a second shorter

spicule (approximately 295 jm), located

distally from the first, both extending to

the pallial cavity and a third short spi-

cule (approximately 100 im) located in

the area where the copulatory gland na-

rrows and terminates. The two copula-

tory glands are long (approximately 430

pm) and tubular (Fig. 3A, B), the glan-

dular walls are very similar in appea-

rance to the walls of the spawning duct.

Anteriorly copulatory glands are wide

(105 qm in diameter), posteriorly these

glands narrow in a duct which appears

to open on the posterior third of the lon-

gest copulatory spicule (Fig. 3A).

103

Iberus, 21 (2), 2003

DISCUSSION

Ocheyoherpia bursata n. sp. is placed in

the order Sterrofustia, because the scleri-

tes mantle are formed mostly by solid

acicles. The distich radula and the sube-

pithelial ventral foregut glandular organs

(type A after SALVINI-PLAWEN, 1978) in

ampoule-shaped, would place this spe-

cies within the family Phyllomeniidae.

SCHELTEMA (1999) however, differs and

does not include the genus Ocheyoherpia

in the family Phyllomeniidae because it

does not have true gonoducts which she

holds as being unique to true species of

genus Phyllomenia. Furthermore the ske-

letal sclerites are lacking in Ocheyoherpia

spp; and the radular teeth have the form

of a bar with denticles, unlike the disti-

chous hooks of the Phyllomeniidae. Ne-

vertheless we think, as SALVINI-PLAWEN

(1978, 2003), that the presence of true go-

noducts could represent a specialisation

within that genus. The sclerites mantle

are mostly formed by solid acicles like

genus Phyllomenia and O. bursata n. sp.

has two types of oar-shaped scales simi-

lar to those described in species of the ge-

nera Phyllomenia Thiele, 1913, and Li-

tutherpia Salvini-Plawen, 1978 (SALVINI-

PLAWEN, 1978). And the type of radula

presented by the genus Ocheyoherpia, see

Figure 1] here, Figure 93 (SALvINI-PLA-

WEN, 1978) in the original descriptions of

Ocheyoherpia lituifera Salvini-Plawen, 1978

and Figure 4 (SCHELTEMA, 1999) in the

original descriptions of Ocheyoherpia tra-

chia Scheltema, 1999, show that they have

radulae formed by pairs of hooked teeth

with smaller lateral denticles, ¡i.e., distich

radulae, like those of the Phyllomeniidae

and not bar-shaped radular teeth with

denticles.

O. bursata n. sp., has characters diag-

nostic of the genus in the sense of SAL-

VINI-PLAWEN (1978) and SCHELTEMA

(1999). Some characters, however, are

particular to this new species and do not

correspond with those of the other two

species of the genus (0. lituifera and O.

trachia) and lead us to specify the diagno-

sis of the genus. In the original diagnosis

of the genus (SALVINI-PLAWEN, 1978), it is

reported that types of sclerites: solid aci-

104

cles and hooks are present. SCHELTEMA

(1999) extends this diagnosis by adding a

third type: solid serrated acicular sclerites

at the distal tip, since they appear in two

species known at this time; but in our ob-

servations, the new species O. bursata has

no acicular sclerites with a serrated distal

tip. We believe that the original diagnosis

of the genus should prevail, i.e. there are

two types of solid sclerites: acicles and

hooked. In the diagnosis of the genus,

SALVINI-PLAWEN (1978) states that the

mouth opens in a channel with no scleri-

tes and which is linked to the atrium, ¡.e.,

the mouth is actually in the atrium, since

the channel is a continuation of the

atrium, as deduced from the description

and illustration. In O. bursata n. sp., we

note that the mouth is clearly in the pos-

terior part of the atrium, for this reason,

in the generic diagnosis, should indicate

that the mouth opens into the atrium, di-

rectly or via a posterior channel without

sclerites. The radula in the genus Ocheyo-

herpia is distich, as per the original diag-

nosis (SALVINI-PLAWEN, 1978). Our obser-

vations coincide with those of SCHEL-

TEMA (1999) in that the radular teeth

present a pair of characteristic curved

and fused distal denticles, a character

which should be added to the diagnosis

of the genus. In O. bursata n. sp., the dis-

tal denticles are located in the same plane

as the rest of the radular tooth and are

not curved upwards and back, as descri-

bed by SCHELTEMA (1999) in O. trachia, so

that this latter characteristic should not

be included in the diagnosis. Conse-

quently, we include an amended diagno-

sis of the genus Ocheyoherpia in the Re-

sults section.

Before this study two species of the

genus Ocheyoherpia were known, both

from Antarctic or sub-Antarctic waters

(SALVINI-PLAWEN, 1978; SCHELTEMA,

1999): O. lituifera Salvini-Plawen, 1978 is

known of the South Sandwich Islands,

South Georgia and the South Shetland

Islands; and O. trachía Scheltema, 1999 is

known of the Macquarie Island.

There are clear differences (Table 1)

between O. bursata n. sp. and the other

two species of the genus (SALVINI-PLA-

WEN, 1978; SCHELTEMA, 1999). O. bursata

GARCÍA-ÁLVAREZ AND URGORRI: A new species of Phyllomeniidae from the Antarctica

l dl ALTA ODIA, Se AAA

Figure 3. A: Schematic organization of the posterior part of the body of Ocheyoherpia bursata n. sp.

B, C, D, E: Microphotographs of the cross-sections of the posterior region of the body correspon-

ding to lines 1, 2, 3, 4. Cs: Copulatory spicule; Gl: Copulatory gland; Ht: Heart; Ma: Mantle;

Mg: Midgut; Ms: Midgut sac; Pc: Pallial cavity; Pd: Pericardioduct; Pr: Pericardium; Re: Rectum;

Sd: Spawning duct; Sr: Seminal receptacle; Vs: Seminal vesicle.

Figure 3. A: Organización esquemática de la parte posterior del cuerpo de Ocheyoherpia bursata n. sp.

B, C, D, E: Microfotografías de los cortes en sección de la región posterior del cuerpo correspondientes a

las líneas 1, 2, 3, 4. Cs: Espículas copulatrices; Gl: Gládula copulatríz; Ht: Corazón; Ma: Manto; Mg:

Intestino; Ms: Saco del intestino; Pc: Cavidad paleal; Pd: Pericardioducto; Pr: Pericardio; Re: Recto; Sd:

Conducto de desove; Sr: Receptáculo seminal; Vs: Vesícula seminal.

105

Iberus, 21 (2), 2003

Table I. Comparative table of characters for Ocheyoherpia species.

Tabla 1. Tabla comparativa de los caracteres de las especies de Ocheyoherpia.

———————<_—_—_——_—Á————_——__———

>0. lituifera 0. trachia 0. bursata n. sp.

Size 5 x 0.5 mm 6.5x 1.4 mm 2.25 x 0.55 mm

Carina No Yes No

Cuticula 35-60 ym 18-22 ym 30-35 pm >

Serrated acicles Yes Yes No

Dar-shaped scales No No Yes

Buccal opening In atrial channel -- In atrium

Radular teeth 64 x 10 ym 75x 18 pm 48 x 11 ym

Distal denticles upwards and Distal denticles upwards and Distal denticle in same plane

back. 5-6 lateral denticles back. 7-9 lateral denticles. as radular tooth. 5-6 lateral teeth

Posterior midgut sacs No No Yes

Seminal vesicles No Yes Yes

Copulatory glands Into a triangular tube Open into the grooved Open in the posterior

third part of longest

copulatory spicule

3 pairs, 3 different sizes

distal end of smaller

copulatory spicule

2 pairs, 2 different sizes

formed by 3 small

copulatory spicules

Copulatory spicules 4 pairs, 3 small, 1 large

(SALVINIEPLAWEN, 1978; SCHELTEMA,

n. sp. has a pair of midgut sacs extending

under the spawning duct, which are not

present in O. lituifera and O. trachia. As re-

gards the mantle sclerites, O. lituifera and

O. trachia have distally serrate acicular

sclerites which O. bursata n. sp. lacks, ne-

vertheless, O. bursata n. sp. has two types

of oar-shaped scales which are not pre-

sent in the other two species. The radula

in O. bursata n. sp. is smaller (48 um long

in O bursata n. sp. by 64 yum long in O. li-

tuifera and 75 ym long in O. trachia). 1t

also has the pair of fused distal denticles

in the same plane as the radular tooth,

this clearly differs from the arrangement

of the fused distal denticles in O. lituifera

and O. trachia (SCHELTEMA, 1999), where

the denticles curve upwards and back

and lie in a different plane to that of their

base (see Figs . 4D, F and 5B in SCHEL-

TEMA, 1999). The copulatory glands in O.

bursata n. sp. open on the posterior third

part of the longest compulatory spicule,

however in O. lituifera it opens into a

triangular tube formed by the tree small

copulatory spiculaes and in O. trachia it

opens into the grooved and ridged distal

end of the smaller copulatory spicule. O.

bursata n. sp. further differs (Table I) from

each of the other two species in the genus

1999): O. bursata n. sp. differs from O. li-

tuifera in that it has a thin cuticle (30-35

pum in O. bursata n. sp. and 35-60 pm in O.

lituifera) and differs from O. trachia in that

the cuticle is thicker (30-35 im as compa-

red wich 18-22 um). O. trachia has a ca-

rina which is not present in O. bursata n.

sp. O. bursata n. sp. has well developed

seminal vesicles, which are not described

for O. lituifera. O. bursata n. sp. has 3 pairs

of copulatory spicules but only 2 pairs

are described for O. trachia and 4 pairs for

O. lituifera

ACKNOWLEDGEMENTS

We are grateful to Prof. Dr. Fernando

Cobo Gradín at the University of San-

tiago de Compostela and to Prof. Dr.

Luitfried v. Salvini-Plawen at the Uni-

versity of Vienna for their invaluable

help. This paper is part of the BENTART

research projects (ANT94-1161-E,

ANT95-1011; ANT97-2097-E) and the

Integrated Spanish-Austrian Co-opera-

tion Actions (HU1997-0002; HU2000-

0010). Our thanks also to lan Emmett for

the English translation.

GARCÍA-ÁLVAREZ AND URGORRI: A new species of Phyllomeniidae from the Antarctica

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Foregut and reproductive tract anatomy of three species of

the Strombina-group (Buccinoidea: Columbellidae)

Anatomía del tubo digestivo y del tracto reproductor en tres especies

del grupo Strombina (Buccinoidea: Columbellidae)

Helena FORTUNATO*

Recibido el 13-1X-2002. Aceptado el 16-VI-2003

ABSTRACT

This paper reports for the first time anatomical data for three species of the Strombina-

group sensu Jung, 1989: Bifurcium bicanaliferum, Sincola (Dorsina) gibberula and Clav-

istrombina clavulus. Anatomical data for the foregut and reproductive tracts were deter-

mined from gross dissections mainly of living specimens. Both the anterior digestive tract

and the reproductive systems of the three species are similar in their general plan and con-

form with what is known for columbellids and neogastropods in general.

Differences among the three species are mostly within accessory structures. None of them

presents either a bursa copulatrix or a prostate gland. The ingestive gland is also absent. C.

clavulus males present an open reproductive system with a diverticulum communicating the

vas deferens with the body cavity. This species also presents a separated albumen gland,

whereas in both S. gibberula and B. bicanaliferum only an internal septum separates this

gland from the capsule gland. Bifurcium and Sincola both appear during the early Miocene

of Dominican Republic, which could bear on their anatomical similitude, whereas Clavistrombina

is a Recent genus of the eastern Pacific. More studies of other species of these genera are

needed in order to better understand possible evolutionary connections of these body plans.

RESUMEN

Se describe por vez primera datos anatómicos de tres especies del grupo Strombina sensu

Jung, 1989: Bifurcium bicanaliferum, Sincola (Dorsina) gibberula y Clavistrombina clavulus. Los

datos anatómicos se determinaron a partir de disección de especimenes vivos. Los tractos

digestivo y reproductor de las tres especies estudiadas son similares en su estructura general y

se corresponden con los conocidos para columbellidos y neogasterópodos en general.

Las diferencias se concentran en las estructuras accesorias. Ninguna especie presenta bolsa

copulatriz, próstata o glándula ingestiva. Los machos de C. clavulus tienen un sistema repro-

ductor abierto con un divertículo que comunica el vaso deferente con la cavidad del cuerpo.

Esta especie tiene una glándula del albumen separada, mientras que en $. gibberula y en

B. bicanaliferum sólo un septo la separa de la de la cápsula. Tanto Bifurcium como Sincola

aparecieron durante el Mioceno temprano en la República Dominicana, lo cual puede

explicar su similitud anatómica, mientras que Clavistrombina es una especie reciente del Pací-

fico este. Se precisan más estudios sobre otras especies de estos géneros para una mejor

comprensión de las posibles conexiones evolutivas entre estas características anatómicas.

KEY WORDS: Gastropods, Strombina-group, anatomy, foregut, reproductive tract

PALABRAS CLAVE: Gasterópodos, grupo Strombina, anatomía, tracto digestivo, tracto reproductor.

* Center for Tropical Anthropology and Paleoecology, Smithsonian Tropical Research Institute, P.O. Box 169,

Balboa, Panama R P, e-mail: fortunaeBancon.si.edu

109

Iberus, 21 (2), 2003

INTRODUCTION

Neogastropods are generally consid-

ered the most highly evolved proso-

branch gastropods. This large order,

with more than 1000 recent and fossil

genera and subgenera (TAYLOR AND

SoHL, 1962), is characterized by a shell

with an elongate siphonal canal and

rachiglossate or toxoglossate radula.

The classification and biology of neogas-

tropods has been the focus of many

authors as it contains several extremely

important groups, both economically

and biologically (ADAMS AND ADAMS,

1858; THIELE, 1929; RIisBEC, 1954;

FRETTER AND GRAHAM, 1962; PONDER,

1973; HARASEWYCH, 1984; BRIGHT AND

ELLIS, 1990; KOOL, 1993).

Neogastropods have a rather similar

anatomical organization. This was

shown to be true for the reproductive

systems of the Muricidae, Buccinidae,

and Nassaridae (FRETTER, 1941; FRETTER

AND GRAHAM, 1962). Families such as

Olividae, Columbellidae, Fasciolaridae

and Turridae are less studied (MARCUS

AND MARCUS, 1959, 1960, 1962; SMITH,

1967; HOousToN, 1976; KANTOR, 1991;

SYsOEvV, 1991). The foregut of some of

these groups shows signs of simplifica-

tion, as in the Muricidae (GRAHAM,

1949), whereas others have features of

the Archaeogastropoda (Brown, 1969;

PONDER, 1972), or have become secon-

darily complex (PONDER, 1970).

The family Columbellidae, one of

the groups traditionally included in the

buccinoid neogastropods, appears dur-

ing the Eocene (RADwInN, 1977a). Al-

though relatively younger than most

other neogastropod taxa, which appear

in the Cretaceous, is one of the most di-

verse and abundant, with almost four

hundred recent species (TAYLOR, MORRIS

AND TAYLOR, 1980). Contrasting with

this, there aren't that many works deal-

ing with the anatomy of this group, as

stated above. THIELE (1929) and RADWIN

(1977a, 1977b) use radular features to

subdivide the family into two subfami-

lies. RISBEC (1954) and especially MAR-

CUS AND MARCUS (1962, 1964) give the

first anatomical descriptions of several

110

columbellid species. Later, HOUSTON

(1976) and HOUSTON AND HATFIELD

(1981) described a couple more species.

Several other species were studied in re-

lation to evolution of herbivory in gas-

tropods (HATFIELD,1979; HARASEWYCH,

1990; KANTOR AND MEDINSKAYA, 1991;

MEDINSKAYA, 1992; MEDISNKAYA, 1993;

GURALNICK AND DE MAINTENON, 1997;

DE MAINTENON, 1999).

The Panamic province has a rich

assemblage of columbellids many of

which have a very interesting evolution-

ary history related to the rise of the

Central American Isthmus . This is the

case with the Strombina-group sensu

Jung, 1989. The group consists of five

genera and includes most taxa classified

as Strombina by earlier workers. JuNG

(1989) splits this genus and proposes

several new genera. Despite the split-

ting, the group is still known in the

malacological jargon as the Strombina-

group, following JUNG's (1989) designa-

tion. The group needs extensive system-

atic revision (DE MAINTENON, 1994) and

the relations of these genera to other

tropical American columbellids and to

each other is still poorly unknown

(RADWIN, 1977a, 1977b; JunG, 1989).

Nevertheless, preliminary cladistic

analyses based on shell morphology

and anatomy strongly support the

hypothesis of a separate columbellid

clade composed by these five genera, as

well as the validity of JunG's (1989)

major genera and subgenera (FORTU-

NATO AND JUNG, 1995).

The Strombina-group is amongst the

most abundant and diverse Neogene

gastropods, and is represented by more

than 30 living species in the eastern

Pacific but only four in the Caribbean,

where they suffered a massive extinc-

tion at the end of the Pliocene (JUNG,

1989; JACKSON, JUNG, COATES AND

COLLINS, 1993; JACKSON, JUNG AND FOR-

TUNATO, 1996; FORTUNATO, 1999).

The Strombina-group has been used

as a model taxon to study changing

species morphology and diversity dur-

ing the gradual emergence of the Isth-

FORTUNATO: Foregut and reproductive tract anatomy of three Strombina-group species

Figure 1. Portraits of the species studied. A: Bifurcium bicanaliferum (B. G. Sowerby I, 1832); B: Sincola

(Dorsina) gibberula (B. G. Sowerby 1, 1832); C: Clavistrombina clavulus (B. G. Sowerby 1, 1834).

Figura 1. Especies estudiadas. A: Bifurcium bicanaliferum (B. G. Sowerby L, 1832); B: Sincola

(Dorsina) gibberula (B. G. Sowerby l, 1832); C: Clavistrombina clavulus (B. G. Sowerby l, 1834).

mus of Panama (JACKSON, JUNG AND

FORTUNATO, 1996; FORTUNATO, 1998,

1999). In spite of this, very little is

known of the biology of these gas-

tropods (CIPRIANI AND PENCHASZADEH,

1993; CIPRIANI, FORTUNATO AND Ro-

DRIGUEZ, 1996; FORTUNATO, PEN-

CHASZADEH AND MILOSLAVICH, 1998;

FORTUNATO, 2002). There are no anatom-

ical studies of any of its species, except

for some references to the type of radu-

lae (RADWIN, 1978; HOUBRICK, 1983;

EMERSON, 1993). The objective of this

paper it is to present the gross anatomy

of the digestive and reproductive sys-

tems of three species: Bifurcium bicanal-

iferum (G. B. Sowerby 1, 1832), Sincola

(Dorsina) gibberula (G. B. Sowerby 1,

1832) and Clavistrombina clavulus (G. B.

Sowerby 1, 1834). B. bicanaliferum is the

only living species of this genus other-

wise known from Miocene deposits of

the Caribbean. S. gibberula is one of the

three extant species of this genus, also

known from Miocene deposits of the

Caribbean. C. clavulus is the only species

of this monotypic genus and has no

known fossil record (JUNG, 1989). All

three species inhabit the shallow water

(up to 40 m deep) coasts of the eastern

Pacific. Whereas the first two species

can be found in muddy and sandy

beaches, the third prefers rocky environ-

ments. In spite of the fact that all three

species have a very wide distribution,

ranging from the Gulf of California

1418]

Iberus, 21 (2), 2003

Table I. Summary of dimensions of several organs for the three species discussed here. Measure-

ments were made with fixed specimens using light microscopy.

Tabla I. Resumen de las dimensiones de varios órganos de las tres especies estudiadas. Las medidas fueron

hechas a partir de especímenes fijados con un microscopio óptico.

Length Length of

of penis proboscis Length

Genus and species (mm) (mm) (mm)

Bifurcium bicanaliferum 11 10 12

Sincola gibberula 7 5 15

Clavistrombina clavulus 12 ] 10

Radulae Opercula

N.of Sizeof N.of Sizeof Lengih Width

lateral — loteral — medial medial (mm) (mm)

teeth — teeth (pm) plate — plate (pm)

360 79 180 30 1250 0Al

450 53 225 20 1,5 |

418 68 209 26 4 2

through Peru, C. clavulus is much less

abundant than the other two species. All

three species are scavengers and have

planktotrophic larvae (FORTUNATO ET

AL., 1998; FORTUNATO, 2002).

MATERIAL AND METHODS

All work was done with live collected

specimens. Collections were done by

hand during low tides and by trawling

from small boats. Both live and preserved

in 5% buffered formalin material was

used for this study. About 10 specimens

of both sexes from each studied species

were dissected. Soft parts were removed

from the shells with a vise after the

animals have been relaxed with menthol

crystals added to the seawater. Before

RESULTS

1. Morphology of the anterior ali-

mentary tract

dissecting, material was submerged in

Methylene blue to delineate tissues and

organs for gross anatomy study. Histo-

logical sections were done to study the

proboscises. Standard histological tech-

niques were used, the tissues infiltrated

with paraffin and sectioned at 7 yum.

Transverse sections of of the proboscis

were stained with Hematoxylin/eosin

(HUMASON, 1962). Measurements of

penises, proboscises, and radular teeth

were done in fixed specimens using light

microscopy and a millimeter ocular .

Radulae were removed and cleaned with

warm 10% KOH until completely free of

tissue. Opercula were treated with 37%

hydrogen peroxyde. Both radulae and

opercula were coated with gold-palla-

dium, and examined under a JEOL HMS-

5300LV Scanning Electron Microscope.

Bifurcium bicanaliferum (Fig. 1A)

The operculum (Fig. 2A) is yellow-

ish, rounded, and small (1 mm long and

1.2 mm wide) (Table I). It is very thin

and transparent with a proteinaceous

consistency. The growth rings are evenly

spaced and it has an antero-posterior

located nucleus.

The animal has a long, very mobile pro-

boscis (prb) tapering towards a small slit-

1412

like mouth opening (mo) (Fig. 4). When

resting, the basal part of the proboscis

retracts into a tubular fold of the body wall,

the proboscis sheath. The proboscis mea-

sures about 10 mm (in a fixed animal)

(Table D and is of pleurembolic type, typi-

cal of a scavenger and predatory life style.

The mouth leeds to the buccal cavity

which occupies the first portion of the

FORTUNATO: Foregut and reproductive tract anatomy of three Strombina-group species

Figure 2. Scanning electron micrographs of opercula. A: Bifurcium bicanaliferum; B: Sincola

(Dorsina) gibberula; C: Clavistrombina clavulus. Scale bar.50 um.

Figura 2. Microfotografías al MEB de opérculos. A: Bifurcium bicanaliferum; B: Sincola (Dorsina)

gibberula; C: Clavistrombina clavulus. Escalas 50 ym.

proboscis. Here the odontophore and

the radular sac are located. The

rachiglossate type radula (1+1+1) (Fig.

3A) is about 12 mm in length (Table D.

The radular ribbon is typically collum-

bellid in shape, narrow, and has two

lateral rows of 180 teeth. The median

plate, or rachidian tooth, is almost rec-

tangular, long and has rounded edges.

Each median plate is 30 pm in length.

The lateral teeth have a sigmoid shape

and are separated from the rachidian by

a large inter-space. Each lateral tooth

measures 79 ym in length. They have a

wide base, a shaft and two hooked

cusps at the tip. A single blunt hook

appears in the middle of the tooth shaft.

The radula and several layers of the sur-

bradular membrane together form the

radular sheath at the end of which the

radula is secreted.

The anterior oesophagus, (Fig. 4;

aoe), starts at the posterior end of the

buccal cavity after separating from the

radular sac. Inside the proboscis, the

anterior oesophagus has a ventral posi-

tion. Once it leaves the proboscis, it con-

tinues towards the neural ring. This

section of the oesophagus is adhered up

to its middle through a connective tissue

layer to the proboscis sheath. Just after

the neural ring, the anterior oesophagus

expands in diameter forming the Valve

of Leiblein (vle) or oesophageal bulb.

The valve has a reddish natural col-

oration and a pear shape. The

oesophageal bulb is surrounded by the

large salivary glands (sgl). These are

paired, yellowish glands and they are

not attached to the valve. These glands

are lobed and the lobes are held

together by strong strands of tissue. The

salivary ducts (sd) leave the glands and

enter the lateral walls of the anterior

oesophagus just before the expansion

that represents the oesophageal bulb.

The ducts follow the anterior oesopha-

gus and discharge their secretions in the

buccal mass. The oesophageal bulb

signals the beginning of the mid-

oesophagus (moe). This bulbous expan-

sion prevents the food from being

sucked forward.

The beginning of the mid-oesopha-

gus is narrow. After traversing the

neural ring, it makes an S-shape curve

after which it begins to engross. The

large Gland of Leiblein (gle), or foregut

gland, enters directly into the mid-

oesophagus without a delimited duct.

This unpaired gland has a V-shape and

is dark brown. After the insertion of the

foregut gland, the oesophagus continues

to enlarge becoming the posterior

oesophagus (poe) that ends in the

stomach.

Iberus, 21 (2), 2003

The proboscis is very long and its wall

is composed of a thin outer layer of cuticle

(cut) (Fig. 5). The cells here are cubic with

a big central nucleus. The next layer is

composed of longitudinal muscles (Iml)

followed by a layer of helical muscle tissue

(hml). The subradular cartilages (sc),

located within the radular sac (rs), have

big cells with a central nucleus. The ante-

rior 0esophagus (aoe) is lined with a layer

of columnar epithelium interspersed with

mucous ciliated cells. The salivary ducts

(sd) are located laterally relative to the

anterior oesophagus.

Sincola (Dorsina) gibberula (Fig. 1B)

The operculum of this species (Fig.

2B) is very similar to the anterior one. lt

is oval shaped, small for the overall size

of the animal (Imm long, 1.5 mm wide)

(Table D, and yellowish. Its consistency

is proteinaceous and it is somewhat thin

and transparent. The growth rings are

evenly spaced and it has an antero-

lateral located nucleus.

Figure 6 shows a short (5 mm)

pleurembolic type proboscis (prb) (Table

D that ends in a small slit-like mouth

opening (mo). The proboscis in this

species is almost translucent. Its basal

part retracts into a tubular fold of the

body wall, the proboscis sheath.

The mouth leeds to the buccal mass

where the odontophore and the radular

sac are located. The rachiglossate type

radula (1+1+1) (Fig. 3B) is about 15 mm

in length (Table D). The typically colum-

bellid radular ribbon is narrow, and has

two lateral rows of 225 teeth. The

rachidian tooth is almost square in

shape, narrow, with rounded edges. It

measures 20 um. The sigmoid lateral

teeth are separated from the median

plate by a large inter-space. Lateral teeth

have a wide base and a shaft with two

hooked cusps at the tip. Lateral teeth

measure 53 um. The hook in the middle

of the tooth shaft is better delineated

here than in the previous species. The

radula is constantly being secreted at

the end of the radular sheath, formed by

the radula itself and the layers of the

subradular membrane.

The anterior oesophagus (Fig. 6; ace)

begins at the posterior end of the buccal

mass and runs dorsally inside the pro-

boscis. The Valve of Leiblein (vle) or

oesophageal bulb consists of a small

expansion of the oesophagus just before

114

the neural ring has a reddish natural

coloration and a pear shape and is sur-

rounded by the large salivary glands

(sgl). These paired, creamy glands are

not connected in any way to the valve.

These glands are lobed and the lobes are

held together by strong strands of

tissue. The long and curved salivary

ducts (sd) leave the glands and enter

directly into the anterior oesophagus

through the posterior lateral walls of the

proboscis. The ducts run laterally inside

the oesophagus and discharge their

secretions into the buccal mass. The

mid-oesophagus (moe) begins after the

ocesophageal bulb. The mid-oesophagus

is thick in most of its length. The Gland

of Leiblein (gle), or foregut gland, enters

into the mid-oesophagus through a

small, narrow duct. This is a small, light

brown, unpaired organ, with a conic

shape. The posterior oesophagus (poe),

that starts after the insertion of the

foregut gland, ends in the stomach.

The proboscis walls have an outer

layer of cuticle (cut) (Fig. 7). The cells of

the cuticle are big, cubic, and have a

prominent central nucleus. The next

layer is composed of helical muscles

(hml) followed by a layer of longitudi-

nal muscle tissue (Iml).

The presence of abundant connec-

tive tissue (ct) is noticeable at this level.

The subradular cartilages (sc), located

within the radular sac (rs), have big cells

with a prominent nucleus. Remnants of

radular teeth can also be observed

inside the radular sac. The salivary

ducts (sd) are located on both sides of

the anterior oesophagus (aoe). The later

has an internal layer of columnar

epithelium interspersed with mucous

ciliated cells.

FORTUNATO: Foregut and reproductive tract anatomy of three Strombina-group species

A Py E y

Figure 3. Scanning electron micrograph of radulae. A: Bifurcium bicanaliferum; B: Síncola

(Dorsina) gibberula, C: Clavistrombina clavulus. Scale bar 100 um.

Figura 3. Microfotografías al MENB de rádulas. A: Bifurcium bicanaliferum; B: Sincola (Dorsina)

gibberula; C: Clavistrombina clavulus. Escalas 100 ym.

Clavistrombina clavulus (Fig. 1C)

The operculum (Fig. 2C) of this

species is quite different from the two

described before. it has a lenticular

shape, brownish color, and is four mil-

limeters long and two millimeters wide

(Table D. It is relatively thick with a cor-

neous consistency. The growth rings are

evenly spaced and it has an antero-pos-

terior located nucleus, and exhibits a

ridge in the middle which gives it a very

uncommon aspect.

Figure 8 shows a short proboscis

(prb) that ends in a small slit-like mouth

opening (mo). At rest, the basal part of

the proboscis retracts into the proboscis

sheath formed by a tubular fold of the

body wall. The proboscis measures

seven millimeters (in a fixed animal)

(Table D and is of pleurembolic type.

The mouth ends in the buccal mass

located in the first section of the pro-

boscis, and where the odontophore and

the radular sac are located. The radula is

10 mm in length (Table D and is

rachiglossate (1+1+1) (Fig. 3C). The

radular ribbon has a collumbellid shape,

is narrow, and is composed of two

lateral rows of 209 teeth. The median

plate, or rachidian tooth, is almost

square, with rounded edges. The

median plate is 26 ym in length. The

lateral teeth are sigmoid, and separated

from the median plate by a very large

inter-space. These teeth are 68 ym in

length. The lateral tooth is composed of

a narrow base, a shaft with a single

hook in the middle, and two hooked

cusps at the tip. The radular teeth are

secreted at the end of the radular sheath.

The later is formed by the radula itself

and several layers of the surbradular

membrana bundled together.

The short anterior oesophagus (Fig.

8; aoe) starts at the posterior end of the

buccal cavity after separating from the

radular sac. It runs laterally and

expands into the oesophageal bulb or

Valve of Leiblein (vle) shortly after

leaving the proboscis. The valve is

small, somewhat elongated, and

translucent. The paired salivary glands

(sgl) have a yellowish color and sur-

round the oesophageal bulb. The lobes

of the salivary glands are quite big.

They are held together by connective

tissue. The salivary ducts (sd) are long

and curved. They leave the glands and

enter the lateral walls of the proboscis,

almost at one third of its length. They

continue on the inside of the proboscis

until they reach the buccal cavity where

they discharge their secretions. The

oesophageal bulb delimits the begin-

ning of the mid-oesophagus (moe).

The mid-oesophagus is quite narrow

up to the section where the short duct of

the Gland of Leiblein (gle) enters it. This

gland is dark brown and elongated,

115

Iberus, 21 (2), 2003

accompanying the posterior vesophagus

(poe) almost in its entirely length. The

posterior oesophagus thickens slightly

before entering in the stomach.

The foot (ft) is very thick and short

(Fig. 9). The short proboscis is quite

2. Morphology of the reproductive

systems

muscular. The walls have a thick outer

layer of cuticle (cut). Inside the pro-

boscis (prb) large mucous glands (glm)

can be seen. The anterior oesophagus

(aoe) has on both sides the large open-

ings of the salivary ducts (sd).

Bifurcium bicanaliferum (Figs. 10, 13)

The bright yellow ovary (Fig. 10; ov)

is located laterally relative to the diges-

tive gland from which is separated by a

layer of connective tissue. The oviduct

(9vd) is relatively short; it runs parallel

to the body wall and enters the albumen

gland (alb) located near the kidney. The

capsule gland (cgl) follows. At first

glance, the two glands are almost indis-

tinguishable from each other. Neverthe-

less, transversal and longitudinal sec-

tions of both organs show the presence

of a small internal septum dividing both

glands. Histological sections show the

presence of more glandular tissue in the

section corresponding to the albumen

gland than after the internal septum.

The vestibule (vsb) is very short and is

located at the end of the capsule gland,

followed by the female aperture (fop)

which drains directly to the mantle

cavity next to the anus.

The testis (Fig. 13; tes), share with

the digestive gland the posterior part of

the visceral cavity. The testis are located

laterally to the digestive gland and from

it the short testicular duct (ted) runs

parallel to the stomach; after a short

while it becomes extremely convoluted

and becomes the seminal vesicle (sev).

In the posterior region of the body mass,

the seminal vesicle straightens out again

and becomes the vas deferens (vd), a

very thin and quite long duct that runs

parallel to the rectum and the body wall

until it reaches the base of the penis

(pen). It enters the penis becoming the

penial duct (pd), which runs centrally

inside it. It is a convoluted tube that

opens at the tip of the male organ. In

resting animals, the penis is curved

backwards along the right side of the

body and rests in the penial pouch. The

penis measures 11 mm long and is thick-

ened in its middle part, tapering

towards the tip. Its ventral side is some-

what wrinkled and dorso-ventrally flat-

tened.

Sincola (Dorsina) gibberula (Figs. 11, 14)

The yellowish colored ovary (Fig. 11;

ov) is lateral to the digestive gland, and

separated from these by connective

tissue. Follows a short and somewhat

sinuous oviduct (ovd) that runs parallel

to the body wall and enters the albumen

gland (alb). A small external membrane

connects the albumen and the capsule

glands (cgl). Histological sections show

the presence of a relatively deep internal

septum dividing the two regions. The

tissues present in both are also different,

the first being much more glandular

116

than the second. Both organs are located

near the kidney. The vestibule (vsb) is

well delineated in this species and runs

from the anterior region of the capsule

gland parallel to the rectum until it ends

in the female opening (fop) located con-

tiguous to the anus.

The testis (Fig. 14; tes) are located in

the visceral cavity near the digestive

gland, in a lateral position. The testicu-

lar duct (ted) is relatively long, runs par-

allel to the stomach and, soon after

leaving the testis, starts to convolute,

FORTUNATO: Foregut and reproductive tract anatomy of three Strombina-group species

Figures 4, 5. Bifurcium bicanaliferum. 4: macromorphology of the anterior alimentary tract; 5:

transverse section of proboscis. Figures 6, 7. Sincola (Dorsina) gibberula. 6: macromorphology of

the anterior alimentary tract; 7: transverse section of proboscis. Figures 8, 9. Clavistrombina clavu-

lus. 8: macromorphology of the anterior alimentary tract; 9: transverse section of proboscis. Abbre-

viations. aoe: anterior oesophagus; ct: connective tissue; cut: cuticle; ft: foot; gle: gland of Leiblein;

glm: gland of mucus;hml: helical muscle; lml: longitudinal muscle; mo: mouth; moe: middle

oesophagus; poe: posterior oesophagus; prb: proboscis; rs: radular sac; sc: subradular cartilages; sd:

salivary duct; sgl: salivary gland; vle: valve of Leiblein. Scale bars, 4, 6, 8: 2 mm; 5, 7, 9: 250 um

Figuras 4, 5. Bifurcium bicanaliferum. 4: macromorfología del tracto alimentario anterior; 5: sección

transversal de la probóscide. Figuras 6, 7. Sincola (Dorsina) gibberula. 6: macromorfología del tracto

alimentario anterior; 7: sección transversal de la probóscide. Figuras 8, 9. Clavistrombina clavulus. 8:

macromorfología del tracto alimentario anterior; 9: sección transversal de la probóscide. Abreviaturas.

ave: esófago anterior; ct: tejido conectivo; cut: cutícula; ft: pie; gle: glándula de Leiblein; glm: glándula

del mucus;hml: múscula helicoidal; lml: múscula longitudinal; mo: boca; moe: esófago medio; poe:

esófago posterior; prb: probóscide; rs: saco radular; sc: cartílagos subradulares; sd: conducto salivar; sel:

glándula salivar; vle: válvula de Leiblein. Escalas, 4, 6, 8: 2 mm; 5, 7, 9: 250 ym

1517

Iberus, 21 (2), 2003

forming the seminal vesicle (sev). This

has a whitish coloration and is located

laterally to the stomach. At the posterior

end, the gonadal duct becomes straight

again forming the vas deferens (vd). The

later is thick, not very long and runs

parallel to the rectum along the body

walls and enters the base of the penis

(pen). Here it forms the penial duct (pd)

that runs along and inside the penis,

opening at its tip. The penis of this

species is quite short, only seven mil-

limeters long, and thickened throughout

most of its way. Its surface is smooth

and flattened in the dorso-ventral

region. At rest, it is tucked in the penial

pouch located in the dorsal region of the

body wall.

Clavistrombina clavulus (Figs. 12, 15)

The female ducts of this species (Fig.

12) show several differences from the

two previous species. The ovary (ov) is

yellow and has a lateral position relative

to the digestive gland, from which it is

separated by a layer of connective

tissue. The long oviduct (ovd) is sinuous

and divided into two parts. It runs par-

allel to the body wall, curves and enters

the completely delimited albumen

gland (alb), which is located laterally to

the kidney. After traversing the albumen

gland, the gonadal duct leaves it, curves

again and enters the capsule gland (cgl).

The later has a bean shape and is

located anteriorly to the kidney. From its

posterior section starts the long and

plain vestibule (vsb), which runs paral-

lel to the rectum and opens near the

anus in the female opening (top).

Both the testis (Fig. 15; tes) and the

digestive gland are located posteriorly

in the visceral mass. A short testicular

DISCUSSION

Both the anterior alimentary tract

and the reproductive systems of the

three studied species are similar in their

general plan. In what concerns the ante-

rior alimentary tract, the most important

difference lies in the insertion of the sali-

vary ducts, which in B. bicanaliferum oc-

curs in the anterior oesophagus just be-

fore the Valve of Leiblein, whereas in

both S. gibberula and C. clavulus it hap-

pens in the posterior region of the pro-

boscis. B. Bifurcium also presents a much

longer proboscis and anterior oesopha-

gus, as well as a larger oesophageal bulb

118

duct (ted) curves once after leaving the

testis, and soon becomes coiled forming

the seminal vesicle (sev). Both the testic-

ular duct and the seminal vesicle are

located parallel to the stomach. After the

seminal vesicle, the duct becomes

straight again forming a long conduct,

the vas deferens (vd) which runs paral-

lel to the rectum and ventrally gives

origin to a short diverticulum (dv). The

later connects with the mantle cavity

through an opening (omc). Immediately

after the diverticulum, the vas deferens

enters the base of the penis (pen) and

becomes the penial duct (pd), running

centrally and appears as a convoluted

and long tube which opens at the tip of

the penis. The penis is very long, over

12 mm in length, smooth, and flattened

dorso-ventrally. As in the two previous

species, the penis rests in the penial

pouch located in the posterior dorsal

region of the body.

than the other two species, in spite of

the fact that the animal in itself is

smaller in size (range size for B. bicanal-

iferum: 9-13 mm, S. gibberula: 9-12 mm;

C. clavulus: 14-25 mm).

At the level of the reproductive sys-

tems, C. clavulus shows several differ-

ences mostly within the accessory struc-

tures. Of particular interest is the exis-

tence of a well defined albumin gland in

females and a mantle opening (the diver-

ticulum) in the males. The later is of spe-

cial interest as it bears on the evolution

of the open reproductive system. This

FORTUNATO: Foregut and reproductive tract anatomy of three Strombina-group species

Figures 10-12. Macromorphology of the female ducts. 10: Bifurcium bicanaliferum; 11: Sincola

(Dorsina) gibberula; 12: Clavistrombina clavulus. Figures 13- 15. Macromorphology of the male

ducts. 13: Bifurcium bicanaliferum, 14: Sincola (Dorsina) gibberula, 15: Clavistrombina clavulus.

Abbreviations: alb: albumen gland; cgl: capsule gland; dv: diverticulum; fop: female opening; omc:

opening in mantle cavity; ov: ovary; ovd: oviduct; pd: penial duct; pen: penis; ted: testicular duct;

tes: testis; sev: seminal vesicle; vd: vas deferens; vsb: vestibule. Scale bars. 10, 11: 3 mm; 12: 5 mm;

13, 14: 4 mm; 15: 10 mm.

Figuras 10-12. Macromorfología de los conductos femeninos. 10: Bifurcium bicanaliferum,; 11:

Sincola (Dorsina) gibberula; 12: Clavistrombina clavulus. Figuras 13- 15. Macromorfología de los

conductos masculinos. 13: Bifurcium bicanaliferum; 14: Sincola (Dorsina) gibberula; 15: Clavis-

trombina clavulus. Abreviaturas: alb: glándula del albumen; cgl: glándula de la cápsula; du: divertí-

culo; fop: abertura femenina; ov: ovario; omc: abertura en la cavidad del manto; ovd: oviducto; pd: con-

ducto peneal; pen: pene; ted: conducto testicular; tes: testis; sev: vesícula seminal; vd: vaso deferente; usb:

vestíbulo. Escalas. 10, 11: 3 mm; 12: 5 mm; 13, 14: 4 mm; 15: 10 mm.

condition is typical in most mesogas-

tropods and is considered to be the

primitive state (JOHANSSON, 1942; FRET-

TER, 1946). In spite of the fact that most

neogastropods present a fused reproduc-

tive duct, the presence of an opening to

the mantle cavity occur in several colum-

bellids, olivids, muricids, buccinids, and

turrids (MARCUS AND MARCUS, 1959,

1962; SmITH, 1967; Houston, 1976;

HOUSTON AND HATFIELD, 1981). It would

appear that this means a retention of the

primitive state for this species.

The columbellid reproductive system

combines very primitive features, like a

gonopericardial-pallial communication,

with advanced ones as it is the pouch for

the resting penis. Based on this, MARCUS

AND MARCUS (1962) divide the whole

family into two groups: a) males with a

seminal vesicle and no prostate, and

females with an albumen gland and sperm

storing organ; b) absence of seminal vesicle

and presence of prostate in males com-

bined with absence of albumen gland and

sperm storing organ in females (sperm is

kept in the pericardium).

None of the species studied presents

either a bursa copulatrix or a prostate

gland. This means that they wouldn't

very well fit neither of the morphologi-

cal divisions defined above. Interesting

is also the fact that in spite of having an

open male duct, C. clavulus has a higher

compartmentalized reproductive sys-

tem, whereas the other two species have

a more generalized, with less distinct ac-

cessory structures, system. Whereas this

means a more or less advanced condi-

tion it is difficult to say.

1115)

Iberus, 21 (2), 2003

The greater similitude between Bi-

furcium and Sincola is especially interest-

ing. These genera first appear in the

early Miocene in fossil deposits of the

Dominican Republic, Caribbean Sea. On

the other hand, Clavistrombina is a recent

genus of the eastern Pacific. Whereas

these differences in times of origination

could bear on the internal anatomy of

the taxawis difficult to know. Many more

studies of other species of these and

other genera of the group are still

needed in order to be able to make defi-

nite conclusions about the functional

and ecological relationships of the re-

productive systems and their evolution.

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AKNOWLEDGEMENTS

A. Velarde, J. Jara, and E. Rodríguez

assisted in the field and in the labora-

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photographs. A special word of thanks

to F. Rodríguez who did the line draw-

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O Sociedad Española de Malacología Iberus, 21 (2); 123-132, 2003

The genus Schwartziella in Senegal (Gastropoda, Rissoidae)

El género Schwartziella en Senegal (Gastropoda, Rissoidae)

Emilio ROLÁN* and José María HERNÁNDEZ**

Recibido el 17-11-2003. Aceptado el 24-VI-2003

ABSTRACT

The species of the genus Schwartziella from Senegal are revised; five species are illus-

trated, of which one is new to science. S. crassior, usually considered a synonym of S.

africana is now considered a valid species. Two species are considered similar to others

from the Cape Verde archipelago. The shell characters of all the species and some infor-

mation on soft parts, opercula and radulae of some of them are shown.

RESUMEN

Se revisan las especies del género Schwartziella recolectadas en Senegal, mostrando

cinco especies, de las cuales una de ellas es nueva para la ciencia. Se da validez al

taxon S. crassior, hasta ahora considerado una forma de S. africana. Dos especies se

consideran similares a otras descritas en el Archipiélago de Cabo Verde. Se muestran

gráficamente los caracteres de la concha, y alguna informacion sobre partes blandas,

opérculo y rádula de algunas de las especies estudiadas.

KEY WORDS: Rissoidae, Schwartziella, Senegal, new species.

PALABRAS CLAVE: Rissoidae, Schwartziella, Senegal, especie nueva.

INTRODUCTION

DAUTZENBERG (1912) described Ris-

soina africana from Senegal and men-

tioned the variety crassior being repre-

sented by two figures, plate 2, figs. 5-6

and 7-8 respectively. The types of those

taxa are in the MNHN (Figs. 1, 2).

GOFAS (1999) made a revision of the

African species of Rissoidae and con-

cluded that both taxa were the same

species. Schwartziella africana and Ris-

soina punctostriata, descrita por TALAV-

ERA (1975) were the sole species of this

subfamily considered present on the

West African coast. ROLÁN AND LUQUE

(2000) revised the subfamily Rissoininae

from the Cape Verde Archipelago

showing the existence of an important

diversification of this subfamily with 29

species, Ma malmlyitintheiisgenus

Schwartziella, including 26 new species.

In sediment material collected in

Dakar by Jacques Pelorce, and by both

authors, a large quantity of shells of

Schwartziella were examined; and the

present work presents the information

obtained from this study. Some speci-

mens had opercula and remains of soft

parts, but it was very difficult to obtain

radulae from all the species because in

many cases the soft parts were damaged

and the radula was not found or was in

very bad condition.

* Investigador del Museo de Historia Natural, Campus Universitario Sur, 15782, Santiago de Compostela.

** Capitán Quesada 41, 35460 Gáldar, Gran Canaria, Spain.

Iberus, 21 (2), 2003

Figures 1, 2: Schwartziella africana and S. africana var. crassior, both 3.0 mm, Dakar (MNHN).

Figuras 1, 2: Schwartziella africana y S. africana var. crassior, ambas de 3,0 mm, Dakar (MNAHN).

Abbreviations

MNCN Museo Nacional de Ciencias

Naturales, Madrid

MNHN Museum National d'Histoire

Naturelle, Paris

SYSTEMATICS

NNM Natuurhistorischen Museum,

Leiden

CER collection of Emilio Rolán

CJH collection of José María Hernán-

dez

CJP collection of Jacques Pelorce

Family RISSOIDAE Gray, 1847

Subfamily RISSOININAE Stimpson, 1865

Genus Schwartziella Nevil, 1881

Schwartziella africana (Dautzenberg, 1912) (Figs. 1, 3, 4, 12, 17, 22, 26, 27)

Rissoina africana Dautzenberg, 1912. Mission Gruvel. Ann. Inst. Océanogr., 5 (3): 48-49, pl. 2, figs.

5-6. [Type locality: Baie de Hamn, Senegall.

Type material: Holotype (Fig. 1) in MNHN, 3 mm.

Other material examined: 728 shells (of which about 40 with operculum and remaining soft parts),

from several localities in Dakar, Senegal (for locality and collection details, see Table I, where sym-

patric species are also listed).

Description: See DAUTZENBERG (1912)

and GOFAS (1999), where a general de-

scription of the shell is given. We add the

following details: The shell (Figs. 1, 3, 4)

is whitish. The protoconch (Fig. 17) is of

the non-planktotrophic type with a di-

124

ameter of about 315 im and of one spiral

whorl; its surface has irregular flat eleva-

tions. The teleoconch has five whorls with

a slightly curved profile; the axial ribs are

opisthocline, almost rectilinear and the in-

terspaces are of similar size; there are 15-

ROLÁN AND HERNÁNDEZ: The genus Schwartziella in Senegal (Gastropoda, Rissoidae)

Table I. Material studied from the Dakar area in the CER, CJP and CJH.

Tabla 1. Material estudiado del área de Dakar en las colecciones CER, CJP y CJH.

S.africana

Le Tacoma 6-12 m (CER) 61

Le Tacoma 13 m, Epave (CER) 152

Le Tacoma 15 m Epave (CER) 167

Ngor, 14 m, in Palythoa (CER) 81

Madeleines 15-30 m (CER) 36

East Madeleines 6-14 m (CER) 36

NE Madeleines, Oune, 18 m (CER) 20

Petit Thiouriba, 33 m (CER) 3

Grand Thiouriba, 30 m (CER) 7

Cap Vert, Pa Maquete, 33 m (CER) 21

Between Gorée and Dakar, 20 m (CER)

Ngor, in beach sediments (CER) 50

Dahomey, 27-30 m (CER) in Bouquet

S of Gorée, 5-8 m (CER) 10

Grand Thiouriba, 40 m (CJP) 10

Tacoma Epave, 13 m (CP) 22

Les Madeleines, 10-30 m (CP) 3

S of Gorée 3-6 m (UH) 8

Petit Thiouriba, 30-35 m (UH) |

Seminoles, 20-25 m (UH) 1

S of Gorée, 3-6 m (UH) 37

Oceanium Diver Center, 0-6 m (UH) 2

Total 728

S. crassior

IATA |

S. yragoae S.cf. puncticulata — S. cf. hoenselaari

224 6

528 12

375 21

3 24 7

53 23 3

140 ]

13 1

8 1

6 5

7 8 5

5 4 11

3 l

11 5

1617 115 31

17 ribs on the last whorl (Fig. 12). When

the shell is fresh, it is rather translucent

and the soft parts or sand can be seen in

its interior. Microsculpture (Fig. 22) of ir-

regular threads visible mainly on the in-

tervals between the ribs.

Dimensions: most of the material stud-

ied is between 2.8 and 3.5 mm high; in

some populations, shells can reach 3.7 mm.

Periostracum evident and yellowish

to orange in color.

Operculum (Figs. 26, 27) whitish,

transparent and with a marginal nucleus.

Radula (not illustrated) was exam-

ined by optical microscope and showed

no difference from that of $. cf. puncticu-

lata (see below).

Distribution: Known only from

Senegal. In the Dakar area, it is one of

the most common species being present

in nearly all of the collecting locations.

Remarks: S. africana was the first Ris-

soininae found on the West African

coast as mentioned by DAUTZENBERG

(1912).

We have made measurement of the

the larger shells of our material from

several lots and they always were

smaller than 3.7 mm. The size referred

in GOFAS (1999) as up to 5.25 mm, may

be due of the inclusion of shells of other

species in this taxon which usually does

not reach this size.

S. africana differs from any other

from European and Cape Verde Islands

species mainly because of its trans-

parency, scarcely curved profile,

opisthocline axial ribs, and its fine

microsculpture.

From other Senegal species living

sympatrically, Table 1! shows the most

important differences.

125

Iberus, 21 (2), 2003

Table IL. Characters of the species from Dakar.

Tabla II. Caracteres de las especies de Dakar.

S 7

2 S S

o NE E 5

'S SS ES 3 o ÉS] S S

SS S S ES S SX S

S. africana 316 28-37 solid 15-17 almost opisthocline same slightly

rectilinear as ribs convex +

S. crassa 291 2.5-3.4 verysolid 11-13 almost orthocline widerthan slightly

rectilinear the ribs COnvex ++

S. yragoue spec. nov. 312 2.4-3.4 lesssolid 8-11 S-curved strongly doublethat convex +++

opisthocline ofribs

S. cf. puncticulata 315 3.8-4.1 verysolid 12-14 almost orthoclineor similar strongly

rectilinear very slightly toribs convex on

opisthocline upper part +++

S. cf. hoenselaari 328 25-30 fragile 8-10 almost orthocineor widerthan slightly

rectilinear — very slightly theribs convex +

opisthocline

Schwartziella crassior (Dautzenberg, 1912) (Figs. 2, 5, 6, 13, 18, 24)

Rissoina africana var. crassior Dautzenberg, 1912. Mission Gruvel. Ann. Inst. Océanogr., 5 (3): 48, pl.

2, figs. 7-8. [Type locality: Baie de Hamn, Senegall.

Type material: Holotype (Fig. 2) in MNHN, 3 mm.

Material examined: 82 shells from the Dakar area (see Table 1 for further explanation).

Description: DAUTZENBERG (1912)

dedicates only one paragraph to this

species which he considers as simply a

form of S. africana. GOFAS (1999) also

considers this species as a form of S.

africana. Thus, a redescription is neces-

sary.

Shell (Figs. 2, 5, 6) length up to 3.4

mm, maximum width 1.5 mm, solid,

elongate-conic, not transparent.

Protoconch (Fig. 18) of 1 whorl of

about 290 pm maximum diameter, of

non-planktotrophic type, with no spiral

sculpture; transition to teleoconch

abrupt. Microsculpture formed by very

numerous irregular flat prominences.

Teleoconch of 4-4 */2 whorls, slightly

convex, not angulated below the suture;

126

last whorl convex; suture shallow but

evident. Color whitish.

Axial sculpture (Fig. 13) consisting

of prominent rounded narrow ortho-

cline (slightly opisthocline on early

whorls), very distantly spaced axial

ribs; numbering 11-13 on the last

whorl. Spiral sculpture almost invis-

ible at low magnification. Microsculp-

ture (Fig. 24) with irregular threads

visible mainly in the interspaces

between ribs.

Aperture D-shaped, small to

medium-sized; inner lip thick; columel-

lar side weakly concave; anterior

channel absent; outer lip strongly

opisthocline with a very thick external

varix; peristome narrow with about 4

ROLÁN AND HERNANDEZ: The genus Schwartziella in Senegal (Gastropoda, Rissoidae)

500 um

Figures 3-16. Shells of Schwartziella species from Dakar, Senegal. 3, 4: S. africana, 3.1, 3.3 mm; 5,

6: S. crassior, 2.8, 2.4 mm; 7, 8: S. yragoae, holotype, 3,3 mm (MNCN) and paratype, 2.8 mm

(MNHN); 9: S. cf. puncticulata, 4.1 mm; 10, 11: S. hoenselaari, 2.5, 2.5 mm; 12: $. africana,

apical view; 13: $. crassior, apical view; 14: S. yragoae, paratype (CER), apical view; 15: S. cf. punc-

ticulata, apical view; 16: S. hoenselaari, apical view.

Figuras 3-16. Conchas de las especies de Schwartziella de Dakar, Senegal. 3, 4: S. africana, 3,1, 3,3

mm; 5, 6: S. crassior, 2,8, 2,4 mm; 7, 8: S. yragoae, holotipo, 3,3 mm (MNCN) y paratipo, 2.8 mm

(MNHN); 9: S. cf. puncticulata, 4,1 mm; 10, 11: S. hoenselaari, 2,5, 2,5 mm; 12: S. africana, vista

apical; 13: S. crassior, vista apical; 14: S. yragoae, vista apical; 15: S. cf. puncticulata, vista apical;

16: S. hoenselaari, vista apical.

127

Iberus, 21 (2), 2003

parallel lines toward the inner part of

the aperture.

Distribution: Known only from some

localities in Dakar area. It is generally

uncommon and was not found in all

localities (see Table 1 for the places of

collecting).

Remarks: This species is consistently

different from S. africana, but only by

examining a large quantity of shells it is

possible to appreciate these constant dif-

ferences. In comparison with S. africana,

S. crassior is smaller, wider, and not

transparent; the axial ribs are orthocline

and less numerous.

The similar species from the Cape

Verde Islands may be differentiated: S.

typica Rolán and Luque, 2000 has a

spiral cord on the protoconch and differ-

ent microsculpture; S. luisi Rolán and

Luque, 2000 has spiral cords; S. minima

Rolán and Luque, 2000 is smaller and

has more prominent spiral sculpture; S.

hoenselaari Rolán and Luque, 2000 has a

spiral sculpture with smooth interspaces

and the axial ribs are lower in number.

Schwartziella yragoae spec. nov. (Figs. 7, 8, 14, 19, 29, 30, 32)

Type material: Holotype (Fig. 7) of 3.3 mm height, deposited in MNCN 15.05/46610; paratypes in

the following collections: MNHN (1); CER (25); CJH (10) and CJP (10).

Other material examined: 1617 shells (about 30 with remains of the soft parts and operculum),

from the Dakar area (see Table 1 for additional explanation).

Type locality: Dakar, Senegal.

Etymology: The species is named after Cristina Garcia Yrago, wife of the junior author, for her

many years of patience and cooperation.

Description: Shell (Figs. 7, 8) length

up to 3.5 mm, maximum width 1.3 mm,

not very solid, elongate-conic, not trans-

parent.

Protoconch (Fig. 19) of a little more

than 1 whorl of about 310 pum of

maximum diameter, of non-plank-

totrophic type, without any spiral sculp-

ture; transition to teleoconch abrupt.

Microsculpture formed by very numer-

ous irregular flat prominences.

Teleoconch of 4-5 strongly convex

whorls, not angulated below the suture;

last whorl strongly convex; suture deep

and evident. Color whitish.

Axial sculpture (Fig. 14) consisting

of prominent sharp, narrow, S-curved,

strongly opisthocline separate axial ribs,

which seem to extend from whorl to

whorl, being less prominent near the

base. Their number is 9-11 on the last

whorl. Spiral sculpture not evident

without magnification. Microsculpture

formed by irregular threads visible

mainly in the interspaces between the

ribs.

Aperture D-shaped, medium sized;

inner lip thick; columellar side weakly

128

concave; anterior channel nearly absent;

outer lip strongly opisthocline, with

thick external varix; with a few parallel

lines toward the inner part of the aper-

ture.

Operculum (Figs. 29, 30) whitish,

transparent, with marginal nucleus.

Radula (Fig. 32) with a rachidian

tooth with a pair of basal denticles, and

a scarcely prominent central cusp with

5-7 small cusps at each side. Inner mar-

ginal teeth finely denticulate; outer mar-

ginal teeth without any denticles on the

external margin.

Distribution: The species is known

from Dakar, where it is the most

common species collected at practically

all the collecting points.

Remarks: S. yragoae spec. nov. is very

different from all the species known

from Cape Verde Islands and Senegal

for the following particular characters:

the S-curved, strong, sharp and narrow

axial ribs, numbering 11 or fewer on the

last whorl, and the very convex whorls

and deep suture. This species lives sym-

patrically with the other species present

in Senegal without intergrades.

ROLÁN AND HERNÁNDEZ: The genus Schwartziella in Senegal (Gastropoda, Rissoidae)

Figures 2 Pro toconehs: 17:

S. africana; 18: S. crassior; 19: S. yragoae spec. nov.; 20: S. cf.

puncticulata; 21: S. cf. hoenselaari. Figures 22-25: Microsculpture. 22: S. africana; 23: S. cf. puncti-

culata; 24: S. crassior, 25: S. cf. hoenselaari.

Figuras 17-21. Protoconchas: 17: S. africana; 18: S. crassior; 19: S. yragoae spec. nov.; 20: S. Ej

puncticulata; 21: S. cf. hoenselaari. Figuras 22-25: Microescultura. 22: S. africana; 23: S. cf. puncti-

culata; 24: S. crassior; 25: S. cf. hoenselaari.

Iberus, 21 (2), 2003

Schwartziella cf. puncticulata Rolán and Luque, 2000 (Figs. 9, 15, 20, 23, 28, 31, 33)

Schwartziella puncticulata Rolán and Luque, 2000. Iberus, 18 (1): 58, figs. 85-87.

Type material: Holotype in NNM. Paratypes, see ROLÁN AND LUQUE (2000).

Material examined: 5 live specimens, Dahomey, 10 Km south of Dakar, 27 m;, 115 shells (five alive

collected) from the Dakar area (see Table I for additional explanation).

Description: See ROLÁN AND LUQUE

(2000). Shells from the Dakar population

are figured (Figs. 9, 15), as well as the

protoconch (Fig. 20) and microsculpture

(Fig. 23).

The animal (Fig. 33) was observed

alive: it is whitish in color, with narrow

and elongate tentacles, the male has a

curved penis.

Operculum (Figs. 28) whitish and

transparent.

The radula (Fig. 31) has a rachidian

tooth with a pair of basal denticles, and

a prominent central cusp with 5-6 small

cusps at each side. Lateral tooth with

about 10 cusps at the inner border; inner

marginal teeth finely denticulate; outer

marginal teeth without any denticles on

the external margin.

Distribution: If the present population

is the same species as that described from

Cape Verde Islands, it is known only

from that archipelago and Senegal.

Remarks: We did not find important

differences between these shells from

Dakar and the Cape Verde material, there-

fore we included them tentatively in that

taxon. The shells from Dakar are slightly

smaller (up to 4.1 mm, against 4.5 in the

Cape Verde shells); also the protoconch is

smaller (about 315 ym against 360 for the

Cape Verde shells). The shells frequently

appear decollated at the apex, which is

not the case in the Cape Verde popula-

tions. Finally, the microsculpture is more

pitted in the Cape Verde shells than in the

Dakar ones, where there are threads.

But all those differences are not very

important and thus we consider proba-

bly both conspecific. Our only doubt

arises how a species with non-plank-

totrophic development might be present

in such widely separate places, but hu-

man action may be considered through

the frequent traffic between the islands

and the main coast.

Schwartziella cf. hoenselaari Rolán and Luque, 2000 (Figs. 10, 11, 16, 21, 25)

Schwartziella hoenselaari Rolán and Luque, 2000. Iberus, 18 (1): 60, figs. 88-92.

Type material: Holotype in NNM. Paratypes, see ROLÁN AND LUQUE (2000).

Material examined: 31 shells from the Dakar area (see Table I for more explanations).

Description: See ROLÁN AND LUQUE

(2000). Shells from Dakar area are figured

(Figs. 10, 11, 16), as well as the protoconch

(Fig. 21) and the microsculpture (Fig. 25).

Distribution: lf the present is the

same species as described from Cape

Verde, only known from this archipel-

ago and Senegal.

Remarks: We did not find important

differences between the shells from Dakar

and those from Cape Verde, thus we

included them tentatively in this taxon

with the Cape Verde material. The shells

from Dakar are slightly narrower than the

130

holotype but not from other shells repre-

sented in the original description. The pro-

toconch is smaller in the type material

(about 300 ym against 315 for the Dakar

shells), and the microsculpture is a little

different, in the type material having alter-

nating lines without microlines. But all

these are scarcely important differences

and so we consider probably both con-

specific. We have the same doubt as in the

preceding species about how a non-plank-

totrophic species may be present in such

widely separated places, but we consider

the same explanation for this presence.

ROLÁN AND HERNÁNDEZ: The genus Schwartziella in Senegal (Gastropoda, Rissoidae)

Figures 26-30. Opercula. 26, 27: S. africana; 28: S. cf. puncticulata; 29, 30: S. yragoae spec. nov.

Figures 31, 32: Radulae: 31: S. cf. puncticulata; 32: S. yragoae spec. nov. Figure 33. Animal

drawing of 5. cf. puncticulata.

Figuras 26-30. Opérculos. 26, 27: S. africana; 28: S. cf. puncticulata; 29, 30: S. yragoae spec. 101.

Figuras 31, 32: Radulas: 31: S. cf. puncticulata; 32: S. yragoae spec. nov. Figura 33. Dibujo del

animal de S. cf. puncticulata.

Iberus, 21 (2), 2003

CONCLUSIONS

In Dakar, there are several species of

Schwartziella. In the present work, five

are described. This is not a final number

of the Dakar species, because some

shells collected in small quantity and

not mentioned in the present work

could be different.

The species studied in Dakar are dif-

ficult to separate on the basis of addi-

tional characters like protoconch or

microsculpture, because they are very

similar to each other. Only the shell

characters are constantly different

(general shape, profile, higth/width

ratio, features of the axial ribs).

The presence and abundance of the

species studied was very variable in

sediments from different localities in

Dakar. Probably, this means that each

species has its own particular habitat,

and the abundance of the shells of each

BIBLIOGRAPHY

DAUTZENBERG, P., 1912. Mission Gruvel sur la

cóte occidentale d'Afrique (1909-1910): Mo-

llusques marins. Annales de l'Institut Océa-

nographique, 5 (3): 1-111, pls. 52-54.

GOFAS, S., 1999. The West African Rissoidae

(Gastropoda: Rissooidea) and their similari-

ties to some European species. The Nautilus,

113 (3): 78-101.

132

species in sediments varies according to

the presence of this habitat in each area.

The different species have some

degree of sympatry in each locality, as it

is evident by sampling in several locali-

ties; the differences of the characters

between these species are constant

without intergrades.

ACKNOWLEDGMENTS

The authors thank Jacques Pelorce,

Le Grau du Roi, France, for his collect-

ing in Dakar where important informa-

tion was recorded and also for the sedi-

ments and for the material sent for the

present study. To Jesús Méndez of the

Centro de Apoyo Científico y Tec-

nológico a la Investigacion (CACTI) of

the University of Vigo for the SEM pho-

tographs. To the MNHN for the loan of

the type material.

ROLÁN, E. AND LUQUE, A. A., 2000. The subfa-

mily Rissoininae (Mollusca: Gastropoda: Ris-

soidae) in the Cape Verde Archipelago (West

Africa). Iberus, 18 (1): 21-94.

TALAVERA, F. G., 1979. Moluscos de sedimen-

tos de la plataforma continental de Mauri-

tania. Boletín del Instituto Español de Oceano-

grafía, 192: 3-18.

O Sociedad Española de Malacología —____——— lberus, 21 (2): 133-139, 2003

Aspectos bioecológicos de Gemma gemma (Totten, 1894)

(Pelecypoda: Veneridae) en la laguna de Chacopata,

Venezuela

Biological aspects of Gemma gemma (Totten, 1894) (Pelecypoda:

Veneridae) on Chacopata lagoon, Venezuela

Ildefonso LIÑERO ARANA* y Marelis MATA

Recibido el 9-1X-2002. Aceptado el 6-VIL-2003

RESUMEN

Gemma gemma [(Totten, 1894) es un pelecipodo endofaunal ovovivíparo, generalmente

no mayor de 5 mm de longitud, y la especie más abundante en la laguna de Chacopata,

situada en la costa nororiental de Venezuela (10% 39” 00” - 10? 41' 00” N y 63* 47'

30” - 60” 49' 50” O). En esta laguna, se tomaron mensualmente 9 réplicas de sedimento

durante 12 meses, a partir de noviembre de 1995. La densidad mensual promedio fue de

2176 + 4032,36 ind.m-2, siendo enero, febrero y noviembre los meses que presentaron

los valores más elevados (12039, 8658 y 3919 ind.m-2, respectivamente). Los especíme-

nes colectados presentaron tallas comprendidas entre 0,450 y 4,250 mm. El periodo de

liberación de los embriones ocurre desde septiembre hasta marzo, observándose dos

periodos de reclutamiento uno en noviembre 95 - enero 96 y otro de febrero a abril 96.

A diferencia de lo que ocurre en latitudes altas de la costa de Estados Unidos, las mayo-

res densidades se presentan durante los meses de menor temperatura, posiblemente

debido al efecto estresante de este factor durante los meses más cálidos.

ABSTRACT

Gemma gemma (Totten, 1894) is an endofaunal ovoviviparous and dioecious pelecy-

pod, generally not greater than 5 mm of length, and the species most abundant in the

lagoon of Chacopata, located in the Northeast coast of Venezuela (10? 39” 00” - 10*

41' 00” N and 63” 47' 30” - 60% 49" 50” W). In this lagoon, 9 monthly sediment repli-

cates were taken during 12 months, from November 1995. The monthly density average

was of 2176 + 4032,36 ind.m2, being January, February and November the months

with the topmost values (12039, 8658 and 3919 ind.m2, respectively). The lengths of

the collected specimens range between 0,450 and 4,250 mm. Brood release period

takes place from September to March, being observed two recruitment periods, one from

November 95 to January 96 and other from February 96 to April 96. Against what it

occurs in high latitudes of the United States coast, the greater densities were observed

during the months of smaller temperature, possibly due to the stress of this factor during

the warmest months.

PALABRAS CLAVE: Gemma gemma, pelecípodo, macrofauna, laguna

KEY WORDS: Gemma gemma, pelecypod, macrofauna, lagoon.

* Instituto Oceanográfico de Venezuela, Universidad de Oriente, Cumaná, Venezuela. e-mail: ilineroWcantv.net

1583

Iberus, 21 (2), 2003

INTRODUCCIÓN

Gemma gemma (Totten, 1894) es un

pelecípodo endofaunal ovovivíparo, de

dimensiones reducidas, generalmente

no mayor de 5 mm de longitud, que

habita en fondos poco profundos desde

Nueva Escocia hasta Florida, Texas y

Las Bahamas, y ha sido introducida en

las Antillas por aves limícolas migrato-

rias (ABBOTT, 1974). Este bivalvo es la

especie más abundante en la Laguna de

Chacopata, sus poblaciones se distribu-

yen en parches y puede alcanzar densi-

dades de hasta más de 86000 ind.m-2 en

algunas zonas, constituyendo una frac-

ción importante de la dieta de aves limí-

colas migratorias y residentes (LINERO,

1994). Debido a la importancia numérica

y ecológica de esta especie en la laguna,

se consideró de interés realizar un

estudio sobre diversos aspectos de su

bioecología.

MATERIAL Y MÉTODOS

La Laguna de Chacopata está

situada en la costa nororiental de Vene-

zuela, aproximadamente entre 10% 39”

00” y 10? 41' 00 N y 63? 47' 30” y 607 49'

50” O, es un cuerpo de agua de aproxi-

madamente 4 km de longitud máxima

en sentido NE-SO, y de 2 km de anchura

máxima. No posee aportes de agua

dulce, excepto los provenientes de las

escorrentías durante la época de lluvias.

Presenta una angosta comunicación

(aproximadamente 20 m de anchura)

con el Mar Caribe. Las orillas están bor-

deadas de manglares, excepto en la

costa sur, donde son escasos y poco

desarrollados. La profundidad media es

de unos 50-60 cm, presentando las

mayores profundidades en las cercanías

de la bocana (3-4 m). En la zona surcen-

tral de la laguna se establecieron 4 esta-

ciones equidistantes 50 m a lo largo de

un transecto lineal, con una profundi-

dad comprendida entre 40 y 60 cm. Con

frecuencia mensual, durante 12 meses, a

partir de noviembre de 1995, se tomaron

9 réplicas de sedimento con nucleadores

de PVC de 5 cm de diámetro y 15 cm de

134

longitud. En cada muestreo se tomaron

registros de la temperatura y muestras

adicionales de sedimento para su análi-

sis granulométrico y de contenido de

materia orgánica.

El análisis granulométrico se realizó

por tamizado y el contenido de materia

orgánica se estimó por la pérdida de

peso de la muestra seca (100 *C, 24 h),

después de combustión a 500 *C duran-

te 24 h (LÓPEZ-JAMAR, 1981).

Las muestras de sedimento fueron

lavadas a través de una serie de tamices,

siendo el último de 250 qm de apertura

de malla. Los especímenes retenidos

fueron fijados en formalina al 8% en

agua de mar. A cada ejemplar se le

determinó la longitud antero-posterior y

dorso-ventral y el número y tamaño de

los huevos en las hembras ovadas.

Se realizó análisis de varianza y

prueba a posteriori SNK (SOKAL Y

ROHLE, 1980), para determinar el efecto

de los meses y de las estaciones sobre

los parámetros biológicos. Cuando los

datos no cumplían los supuestos del

ANOVA, se aplicó la prueba no paramé-

trica de Kruskal-Wallis, seguida de la

prueba a posteriori correspondiente. Las

relaciones entre las longitudes antero-

posterior y la dorso-ventral, así como

entre el número de huevos y la talla de

los ejemplares se analizaron a través de

análisis de regresión.

RESULTADOS

La temperatura del agua en el fondo

estuvo comprendida entre 26 *C (enero

y abril) y 30 *C (agosto), con un prome-

dio de 27,8 *C. Los sedimentos de las

estaciones mostraron un predominio de

la fracción arena, predominando las

arenas fina y media; mientras que la

fracción fina, correspondiente a limo-

arcilla, mostró porcentajes comprendi-

dos entre 7 y 14%. Las partículas más

gruesas, guija y gránulo, son casi en su

totalidad de origen biogénico, princi-

palmente restos de exoesqueletos de

gasterópodos y de bivalvos, y muy par-

ticularmente de G. gemma. El contenido

de materia orgánica de los sedimentos

LINERO-ARANA Y MATA: Aspectos bioecológicos de Gemma gemma en Venezuela

1400

1200

1000

800

600

400 *

DENSIDAD (NÚMERO DE INDIVIDUOS.m

NADIE TE

AIN AA SO

MESES

Figura 1. Evolución mensual de la densidad de Gemma gemma.

Figure 1. Monthly evolution of Gemma gemma density.

de las estaciones mostró valores entre

6,16 y 12,98%

La densidad mensual promedio fue

de 2176 + 4032,36 ind.m2, siendo enero,

febrero y noviembre los meses que pre-

sentaron los valores más elevados

(12039, 8658 y 3919 ind.m2, respectiva-

mente); en los demás meses la densidad

estuvo comprendida entre 28 (mayo) y

538 en marzo (Fig. 1). La densidad

mensual promedio por estación estuvo

comprendida entre 1382 ind.m”2 (esta-

ción 2) y 2862 ind.m”? en la estación 1

G= 2176 + 628,89). No se encontraron

diferencias significativas de la densidad

entre las estaciones de muestreo, pero sí

entre los meses (p < 0,005).

Los especímenes colectados presen-

taron tallas comprendidas entre 0,450 y

4,250 mm. Una fracción importante de la

población (63%) mostró talla promedio

de 2,305 + 0,688 mm. No se pudo realizar

un análisis de la distribución mensual de

tallas debido a que los valores de densi-

dad variaron significativamente entre

meses, por lo que se estudió la distribu-

ción trimestral de frecuencias de tallas en

los dos primeros trimestres de estudio,

debido a que en los otros dos el número

de especímenes colectados fue bajo. De

acuerdo a este análisis (Fig. 2) en los dos

periodos (noviembre 95 - enero 96 y fe-

brero — abril 96) se aprecian dos modas,

aunque menos marcadas en el segundo

trimestre, lo que evidencia la existencia

de dos periodos de reclutamiento. En el

tercer y cuarto trimestre no se colectaron

individuos con talla inferior a 1,2 mm, lo

que aunado al escaso número de indivi-

duos, indica ausencia de reclutamiento.

La correlación entre la longitud

antero-posterior (LAP) y la dorso-

ventral (LDV) fue altamente significa-

tiva (r = 0,975, p < 0,001) (Fig. 3). La

ecuación predictiva que representa la

relación entre estas dos variables fue

LDV = 0,04 + 1,10LAP.

La correlación entre la longitud

antero-posterior de las hembras ovadas

6) y el número de huevos por hembra

(y) fue significativa (r = 0,52, p < 0,01), y

la ecuación obtenida fue y = -12,88 +

62 (Elo):

Se diferenciaron tres etapas de desa-

rrollo de los huevos, la etapa 1 corres-

ponde a huevos esféricos, de color

blanco, con un diámetro entre 99 y 184

pm (x= 131,16 um); la etapa Il incluye

huevos ovalados de aspecto cristalino y

diámetro entre 201 y 279 pm (<=

275,87um), y la etapa III corresponde a

embriones con características similares a

las del adulto y diámetro entre 309 y 500

pm (<= 365,84 yum).

185

Iberus, 21 (2), 2003

708

No DE INDIVIDUOS

]

0,55 -

LONGITUD ANTERO-POSTERIOR (mm)

DOWN VO OO OO O NO ONO A O

DOODaNnaan DODODDODnNaDaaa

A NANA AN DNOa ana

a ARS II AA AAA AAAAAOS

1,15

LONGITUD ANTERO-POSTERIOR (mm)

Figura 2. Distribución de frecuencias de tallas de Gemma gemma. A: periodo noviembre 95 — enero

96; B: periodo febrero 96 — abril 96.

Figure 2. Distribution of sizes frequencies of Gemma gemma. A: period November 95 — January 96;

B: period February 96 — April 96.

DISCUSIÓN

La densidad de G. gemma en la

laguna de Chacopata es baja en compa-

ración con valores reportados para otras

latitudes (entre 2 y 4.106), según THomp-

SON (1982). Esta baja densidad puede

ser debida a varias causas, unas relacio-

nadas con aspectos bióticos y otras con

aspectos físico-químicos. Entre las pri-

meras se pueden citar principalmente el

efecto de la depredación; a este respecto,

SANDERS, GOUDSMIT, MILLS Y HAMPSON

(1962) y GREEN Y HOBSON (1970),

señalan, en Barnstable Harbor, la depre-

dación de G. gemma por los crustáceos

Crangon septumspinosus y Limulus polyp-

hemus, respectivamente. Excluyendo las

aves limícolas, debido a la profundidad

de las estaciones, en la laguna existen

potenciales depredadores de G. gemma,

entre ellos varias especies de peces ben-

tófagos, poliquetos y crustáceos, como

Farfantepenaeus spp., Callinectes spp. y

Pagurus spp. (LIÑERO, 1994).

Por otro lado, la temperatura es uno

de los factores abióticos que puede influir

en las bajas densidades; esta presunción

se basa en el hecho de que las mayores

densidades se obtuvieron en los meses

menos cálidos; GREEN Y HOBSON (1970),

en experimentos controlados, señalan que

136

la temperatura superior letal para el 50%

de los individuos de G. gemma es 35 *C.

Durante este estudio la temperatura

máxima registrada fue de 30 *C en agosto,

pero es necesario señalar que las medi-

ciones de este parámetro fueron realiza-

das entre las 09:00 y las 10:00 h aproxi-

madamente; y en horas próximas al medio-

día, entre las 11:00 y 14:00 h, la tempera-

tura del agua de la laguna es mayor, por

efecto de la intensa insolación, baja pro-

fundidad y escaso movimiento del agua,

en particular durante los meses más

cálidos (mayo-septiembre).

En la laguna esta especie habita

prácticamente en la superficie del sedi-

mento, y las elevadas temperaturas

podrían constituir el principal factor

abiótico que limite la abundancia de

esta especie, bien sea por alcanzar

niveles letales o por producir un menor

número de embriones, aun cuando

GREEN Y HOBSON (1970) reportan que G.

gemma puede eludir temperaturas extre-

mas de la superficie del fondo, enterrán-

dose al menos hasta 4 cm en el sedi-

mento, estrategia que podría presentar

también en la laguna.

En zonas someras estos organismos

pueden ser arrastrados por las corrientes

producidas durante los periodos de

mayor intensidad del viento hasta fondos

LINERO-ARANA Y MATA: Aspectos bioecológicos de Gemma gemma en Venezuela

LONGITUD DORSO-VENTRAL (mm)

159)

0 1 2

y = 0,04 + 1,10x

3 4 5

LONGITUD ANTERO-POSTERIOR (mm)

Figura 3. Regresión entre la longitud dorso-ventral y la longitud antero-posterior de Gemma gemma.

Figure 3. The regression of dorsal-ventral and the antero-posterior lengths on Gemma gemma.

donde la intensidad de la corriente sea

menor (THOMPSON, 1982). En la laguna,

como en el resto de la costa caribeña de

Venezuela, la amplitud de las mareas es

débil, de aproximadamente 30 cm, por lo

que las corrientes en la laguna son

debidas principalmente a la acción de los

vientos alisios, que se presentan con

mayor intensidad en la época de sequía

(noviembre-abril). Esta característica,

aunada a la distribución temporal de la

temperatura, podría explicar la variación

de las densidades en la población estu-

diada. Es probable que exista un trans-

porte de individuos de zonas someras (<

30 cm) hacia fondos de mayor profundi-

dad, donde la intensidad de la corriente

se debilite por efecto de la profundidad.

Esto ocurriría durante el periodo noviem-

bre-abril, durante el cual individuos de

poblaciones someras serían arrastrados

hacia fondos más profundos. Durante los

meses más cálidos, la intensidad del

viento decrece notablemente, por lo que

el transporte no se produciría o los orga-

nismos podrían migrar a más profundi-

dad en el sedimento, según lo señalado

por GREEN Y HOBSON (1970).

En la laguna existen dos especies de

algas filamentosas (Chaetomorpha sp. y

Cladophora sp.) que alcanzan gran desa-

rrollo durante los meses más cálidos.

Estas algas, adheridas precariamente

principalmente a restos de conchas de

gasterópodos y pelecípodos, durante los

meses secos y de alta intensidad de los

alisios, son desprendidas y arrastradas

hasta los márgenes de la laguna, donde

se acumulan y descomponen. En esas

aglomeraciones de algas se han obser-

vado numerosos ejemplares de G.

gemma entre los frondes filamentosos, lo

cual puede ser atribuido a que indivi-

duos de este pelecípodo son arrastrados

por las corrientes y retenidos por las

algas, o que son removidos del fondo

por las algas cuando éstas son desplaza-

das por las corrientes en zonas de poca

profundidad y quedan atrapados en los

numerosos y tupidos filamentos, o a la

acción conjunta de ambas acciones.

La longitud del periodo de liberación

de los embriones parece estar influen-

ciada por la latitud y/o la temperatura

(THOMPSON, 1982). Este autor compara

características de la reproducción y el

desarrollo de G. gemma en San Francisco

Bay, California (37* 45”), con estudios rea-

lizados en Barnstable Harbor, Massachu-

setts (41? 42%) y en Union Beach, New

Jersey (40? 27”). En San Francisco Bay, la

duración del periodo de liberación de los

embriones es de 7 meses, en Barnstable

Harbor es de 3 meses, y en Union Beach

de 5 meses. En la Laguna de Chacopata

se colectaron hembras ovadas en los

137

Iberus, 21 (2), 2003

y = -12,88 + 9,62x

NÚMERO DE HUEVOS

LONGITUD (mm)

Figura 4. Regresión del número de huevos de Gemma gemma según la talla.

Figure 4. The regression of egg number on Gemma gemma size.

meses de enero a marzo y de septiembre

a noviembre, lo que aunado a la distribu-

ción trimestral de tallas permite afirmar

que el periodo de liberación de los

embriones de G. gemma ocurre desde sep-

tiembre hasta marzo, comprendiendo los

meses de menores temperaturas y, al

igual que en San Francisco Bay, el

periodo de liberación de los embriones

comprende 7 meses.

En la Figura 4 se aprecia que en prác-

ticamente todas las tallas se presentan

hembras con un número de huevos rela-

tivamente bajo, inferior a 20, y que sola-

mente el 7% de las hembras poseen más

de 30 huevos, correspondiendo el

máximo de 54 para una hembra de 3,99

mm. Estos resultados difieren notable-

mente de los obtenidos por GREEN Y

HOBSON (1970), quienes en la figura 8 de

su trabajo presentan al 64% de las

hembras con más de 30 huevos, y dos de

ellas con hasta 100 huevos. Al igual que

en el trabajo de Green y Hobson, en este

estudio la talla mínima de producción de

embriones se encuentra cercana a los 1,7

mm. La diferencia en el número de

huevos / hembra entre estos dos estudios

puede ser debida a que los huevos de G.

gemma en Massachussets sean de menor

tamaño que los de la Laguna de Chaco-

pata, tomando en consideración que el

tamaño de los individuos en las dos

poblaciones son similares; sin embargo,

138

no se dispone de referencias sobre el

tamaño de los huevos de este pelecípodo

en otras latitudes que puedan confirmar

esta hipótesis, aunque Sullivan (1948,

citado por BRADLEY Y COOKE, 1959) men-

ciona que cuando los huevos van a ser

liberados miden 340 pum.

THOMPSON (1982) atribuye el mayor

éxito de las poblaciones de G. gemma en

San Francisco Bay con relación a pobla-

ciones de latitudes mayores, al clima

cálido de esa bahía, lo cual se traduce en

periodos reproductivos de mayor dura-

ción y mayor número de reclutamientos,

obteniendo las mayores densidades en

julio y agosto, durante el periodo de

máximo reclutamiento y las menores en

los meses que preceden inmediatamente

al reclutamiento. En la Laguna de Chaco-

pata ocurre lo contrario, las menores den-

sidades se producen en los meses más

cálidos, por lo que podría deducirse que

para esta especie su intervalo de tempe-

ratura Óptima de reproducción y desarro-

llo se encuentra por debajo de 30 *C.

AGRADECIMIENTOS

Los autores desean expresar su agra-

decimiento a los revisores anónimos por

las observaciones realizadas al manus-

crito, las cuales permitieron mejorarlo

notablemente.

LINERO-ARANA Y MATA: Aspectos bioecológicos de Gemma gemma en Venezuela

BIBLIOGRAFÍA

BRADLEY, W. H. Y COOKE, P., 1959. Living and

ancient populations of the clam Germma gemma

in a Maine coast tidal flat. Fishery Bulletin, 58:

306-337.

GREEN, R. H. Y HOoBsOoN, K. D., 1970. Spatial

and temporal structure in a temperate inter-

tidal community, with special emphasis on

Gemma gemma (Pelecypoda: Mollusca). Eco-

logy, 51 (6): 999-1011.

LINERO, L, 1994. Variations spatio-temporelles de

la structure des peuplements benthiques dans

une lagune cótiere tropicale. These Ph. D. Uni-

versité du Québec á Rimouski, Canadá, 177

12128

LÓPEZ-JAMAR, E. 1981. Spatial Distribution of the

Infaunal Benthic Communities of the Ría de

Muros, North-West Spain. Marine Biology,

63: 29-37.

SANDERS, H. L., GoupsmIT, E. M., MILLS, E. L.

Y HAMPSON, G. E., 1962. A study of the in-

tertidal fauna of Barnstable Harbor, Massa-

chusetts. Limnology and Oceanography, 7: 63-

79.

THOMPSON, J. K., 1982. Population structure of

Gemma gemma (Bivalvia: Veneridae) in South

San Francisco Bay, with a comparison to

some Northeastern United States estuarine

populations. Veliger, 24 (3): 281-290.

159

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Iberus, 21 (2): 141-143, 2003

Notas Breves

Nueva información sobre Paladilhiopsis septentrionalis

(Mollusca, Prosobranchia)

New information on Paladilhiopsis septentrionalis (Mollusca,

Prosobranchia)

Emilio ROLÁN* y Beatriz ARCONADA**

Recibido el 9-XI1-2002 Aceptado el 10-VI-2003

INTRODUCCIÓN

ROLÁN Y RAMOS (1995) describen

Paladilhiopsis septentrionalis en base a

conchas recolectadas en las provincias

de Santander, Vizcaya, Álava y Burgos;

en ningún caso se encontraron ejempla-

res vivos o con restos de partes blandas

debido a que, probablemente, se trataba

de una especie estigobionte.

En recientes muestreos realizados

por el primer autor en Asturias, se vol-

vieron a encontrar conchas de esta

especie en varias localidades, e incluso

se consiguió la captura de un ejemplar

vivo, por lo que se aporta nueva infor-

mación sobre esta especie.

MATERIAL ESTUDIADO

Coordenadas UTM:

TP50: Trubia (Oviedo), en el camino

de Trubia a Caces, que sale desde la

estación del ferrocarril, antes de la

fábrica del Nalón, y que va entre este río

y la vía del tren, en un pequeño naci-

miento en una pared, a 2,8 Kms de la

estación de Trubia y a 1,7 Kms de la des-

viación a Pintoria: 1 ejemplar vivo, 18

conchas y 22 fragmentos.

TP50: Carretera de Trubia a Proaza,

en la desviación a Castañedo del Monte,

. apoco del comienzo de la misma, en un

barranco al lado de una fuente, en sedi-

mentos: 1 concha.

QJ30: Carretera de Grado a Tameza,

en sedimentos de la fuente frente al río:

1 concha.

RESULTADOS

Descripción: Concha (Figs. 1-5): Ver

ROLÁN Y RAMOS (1995). Las conchas

estudiadas ahora en las nuevas estacio-

nes tenían una apariencia muy similar a

las presentadas en el trabajo de descrip-

ción, lo mismo que la protoconcha (Fig.

6) y su escultura (Figs. 7-8). En el único

ejemplar vivo observado, la concha era

transparente, al contrario de lo que

ocurre en las conchas vacías, que se

vuelven opacas y de color blanco. Las

* Investigador del Museo de Historia Natural, Campus Universitario Sur, 15782, Santiago de Compostela.

** Museo Nacional de Ciencias Naturales, J. Gutierrez Abascal, 2, 28006 Madrid.

141

Iberus, 21 (2), 2003

Figuras 1-9. Paladilhiopsis septentrionalis. 1-5: conchas, entre 1,2 y 1,5 mm, recolectadas en Trubia,

Asturias; 6: protoconcha; 7, 8: microescultura de la protoconcha; 9: microescultura de la teloconcha.

Figures 1-9. Paladilhiopsis septentrionalis. 1-5: shells, between 1.2 and 1.5 mm, collected in Trubia,

Asturias; 6: protoconch; 7, 8: microsculpture of the protoconch; 9: microsculpture of the teleoconch.

142

ROLÁN Y ARCONADA: Nueva información sobre Paladilhiopsis septentrionalis

dimensiones de las conchas del material

estudiado estaban entre 1,2 y 1,5 mm.

Microescultura de la teloconcha (Fig. 9):

Similar a la de la descripción original,

aunque algo más atenuada.

Animal: Se pudo observar que era

casi totalmente transparente, con un

tono lechoso muy ligero. La rádula, de

color blanco, podía verse perfectamente

por transparencia. Los tentáculos finos y

alargados. No se apreciaban ojos en la

base de los mismos. Lamentablemente,

el tubo se quedó casi sin líquido y el

animal se secó bastante, con lo que no se

pudieron observar detalles anatómicos

en su disección. Sí se pudo apreciar que

se trataba de una hembra. La gonada

que se veía en el último tramo del

cuerpo era de color naranja y con forma

de saco. El ctenido aparecía poco desa-

rrollado, con dos lamellas minúsculas.

BIBLIOGRAFÍA

ROLÁN, E. Y RAMOS, M. A. 1995. Una nueva es-

pecie de Hydrobiidae (Mollusca, Proso-

branchia) del norte de la Península Ibérica.

Iberus, 13 (2): 119-127.

CONCLUSIONES

El área de distribución de P. septen-

trionalis, aunque incluía varias provin-

cias, era en realidad muy pequeña (ver

ROLÁN Y RAMOS, 1995). Con la presente

cita esta distribución se extiende notable-

mente hacia el oeste, hasta Asturias. Por

otra parte, el hallazgo de un ejemplar

vivo, muestra que aun siendo una

especie estigobionte, su distancia a la

superficie no es muy grande, y por ese

motivo pueden aparecer ocasionalmente

ejemplares vivos arrastrados por el agua.

AGRADECIMIENTOS

Los autores agradecen a Jesús

Méndez, del CACTI, Universidad de

Vigo, las fotografías realizadas el MEB.

143

Ia.

Meal: ALA AT

; En 0 da

ERRATUM

In the article

FRANCK BOYER, 2003. The Cystiscidae (Caenogastropoda) from upper reef for-

mations of New Caledonia”. Iberus, 21 (1): 241-272.

a wrong numbering was introduced within the second plate of figures. The

numbering attached to the pictures must be changed as following;

n* 20 becomes n? 18

n* 18 becomes n* 19

n* 21 becomes n? 20

n* 19 becomes n? 21

The numbering used in the text and in the captions are right.

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vez deberá citarse su autor y fecha de su descripción. En el caso de artículos sistemáticos, cuando se den las sinonimias

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critas, y la localidad tipo si es conocida entre corchetes, según el siguiente esquema (préstese especial cuidado a la pun-

tuación):

Dendrodoris limbata (Cuvier, 1804)

Sinonimias

Doris limbata Cuvier, 1804, Ann. Mus. H. N. 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.

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e 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 publicado más de

un trabajo en un año se citarán con letras: (Davis, 1989a; Davis, 1989b). No deberá emplearse op. cit. La lista de refe-

rencias 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 minús-

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número total de páginas. Deberán evitarse referencias a Tesis Doctorales u otros documentos inéditos de difícil con-

sulta. 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.

Miscelánea Zoolgica, 3 (5): 21-51.

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las figuras, que no deberán bajar de los 2 mm. En figuras compuestas, cada parte deberá etiquetarse con letras mayús-

culas, el resto de las letras deberán ser minúsculas. No deberán hacerse referencias a los aumentos de una determi-

nada ilustración, ya que éstos cambian con la reducción, por lo que debe emplearse una escala gráfica. 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 carví. A: animal desplazándose; B: detalle de un rinóforo; C: branquia.

Las abreviaturas empleadas en las ilustraciones deberán incluirse en la hoja de pies de figura.

Los autores interesados en incluir láminas en color deberán abonarlas a precio de coste (30.000 ptas por página). Por

lo demás, deberán ajustarse a los mismos requisitos que los indicados para las figuras.

Si se pretende enviar gráficas o ilustraciones en formato digital es IMPRESCINDIBLE contactar previamente con el

Editor

e Las Tablas se presentarán en hojas separadas, siempre con numeración romana (1, II, IIT,...). Las leyendas se inclui-

rán en una hoja aparte acompañándose de una traducción al inglés. Deberán evitarse las tablas particularmente com-

plejas. Se recomienda reducir el número y extensión de ilustraciones, láminas o tablas al mínimo necesario.

e Los artículos que no se ajusten a las normas de publicación serán devueltos al autor con las indicaciones de los cam-

bios necesarios.

e 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 manuscrito. 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 modificaciones que considere oportunas. Si el trabajo es

aceptado, el autor deberá enviar una copia impresa del mismo corregida, acompañada por una versión en disco flexi-

ble (diskette), utilizando procesadores de texto en sus versiones de DOS o Macintosh. La fecha de aceptación figura-

rá en el artículo publicado.

e Las pruebas de imprenta serán enviadas al autor responsable, EXCLUSIVAMENTE para la corrección de erratas, y

deberán ser devueltas en un plazo máximo de 15 días. Se recomienda prestar especial atención en la corrección de las

pruebas.

e De cada trabajo se entregarán gratuitamente 50 separatas. Aquellos autores que deseen un número mayor, deberán

hacerlo constar al devolver las pruebas de imprenta, y NUNCA POSTERIORMENTE. El coste de las separatas adi-

cionales será cargado al autor.

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 will be published as Supplements. Authors wishing to publish monographs

should contact the Editor. Manuscripts are considered on the understanding that their contents have not appeared or

will not appeared, elsewhere in substantially the same or any abbreviated form.

+ Manuscripts and correspondence regarding editorial matters must be sent to: D. Gonzalo Rodríguez Casero, Editor

de Publicaciones, Apartado 156, 33600 Mieres, Asturias, Spain.

+ Manuscripts may be written in any modern language.

* When a paper exceeds 20 pages, extra pages will be charged to the author(s) at full cost.

+ 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. When a paper has joint

authorship, one author must accept responsability for all correspondence.

* The authors must include a list of at least 4 possible referees; the Editor can choose any others if appropriate.

e Papers should conform the following layout:

First page. This must include a concise but informative title, with mention of family of higher taxon when appropri-

atte, and its Spanish translation. It will be followed by all authors” names and surnames, their full adress(es), an

abstract (and its Spanish translation) not exceeding 200 words which summarizes not only contents but results and

conclusions, and a list of Key Words (and their Spanish translation) under which the article should be indexed.

Following pages. These should content the rest of the paper, divided into sections under short headings. Whenever

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.

e Notes should follow the same layout, without the abstract.

e 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 authori-

ty including the year. In systematic papers, when synonyms of a taxon are given, they must be cited IN FULL, includ-

ing 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 limbata Cuvier, 1804, Ann. Mus. H. N. Paris, 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. Ifa full list

of references of the taxon is to be given immediately below it, the same layout should be followed (also excluding those

nowhere else cited from the Bibliography list).

Only Latin words and names of genera and species should be underlined once or be given in ¿talics. No word must

be written in UPPER CASE LETTERS. SÍ 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 include all of them

[Smith, Jones % Brown (1970)], thereafter use et al. [Smith ez al. (1970)]. Ifan 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 include 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 CAP-

ITALS. The references need not be cited when the author and date are given only as authority for a taxonomic name.

Titles of periodicals must be given IN FULL, not abbreviated. For books, give the title, name 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 minimum. 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. EF.

(Ed.): Prosobranch Phylogeny, Malacological Review, suppl. 4: 129-166.

Ros, J., 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas.

Miscelánea Zoológica, 3 (5): 21-51.

* Figures must be original, in Indian ink on draughtsman's tracing paper. Keep in mind page format and column size

when designing figures. 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 recomended. It is desirable to print figures with their legend

below, so authors are asked to take this into account when preparing full page figures. If computer generated graph-

ics are to be included, they must be printed on high quality white paper with a laser printer. Photographs must be of

good contrast, and should be submitted in the final size. When mounting photographs in a block, ensure spacers are

of uniform width. Remember that grouping photographs of varied contrast results in poor reproduction. Take

account of necessary reduction in lettering drawings; final lettering must be at least 2 mm high. In composite draw-

ings, each figure should be given a capital letter; additional lettering should be in lower-case letters. A scale line is

recomended 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 separate sheets, and numbered with consecutive Arabic num-

bers (1, 2, 3,...), without separating “Plates' and “Figures”. Legends for Figures must be typed in numerical order on

a separate sheet, and an Spanish translation must be included. Follow this example (please note the punctuation):

Figure 1. Neodoris carví. A: animal crawling; B: rinophore; C: gills.

If abbreviations are to be used in illustrations, group them alphabetically after the Legends for Figures section.

Authors wishing to publish illustrations in colour will be charged with additional costs (30,000 ptas, 180 euros per

page). They should be submitted in the same way that black and white prints.

If the authors want to send Figures in digital format, CONTACT the Editor first.

* Tables must be numbered with Roman numbers (1, IL, IIL,...) and each typed on a separate sheet. Headings should

be typed on a separate sheet, together with their English translation. Complex tables should be avoided. As a general

rule, keep the number and extension 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 referees. Based of

these evaluations, the Editorial Board will decide on acceptance or rejection. Anyway, authors will receive a copy of

the referees* comments. Ifa 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 man-

uscript must then be returned to the Editor, together with a floppy disk containing the article written with a DOS or

Macintosh word processor. Dates of reception and acceptance of the manuscript will appear in all published articles.

* Proofs will be sent to the author for correcting errors. 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 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.

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La Socienbap ESPAÑOLA DE MALACOLOGÍA

Junta Directiva desde el 14 de noviembre de 2000

Presidente Emilio Rolán Mosquera

Vicepresidente Diego Moreno Lampreave

Secretario Luis Murillo Guillén

Tesorero Jorge J. Otero Schmitt

Avda. de las Ciencias s/n, Campus Universitario, 15706 Santiago

de Compostela, España

Editor de Publicaciones Gonzalo Rodríguez Casero

Apartado 156, 33600, Mieres del Camino, Asturias, España

Bibliotecario Rafael Araujo Armero

Museo Nacional de Ciencias Naturales, CSIC, c/ José Gutierrez

Abascal 2, 28006 Madrid, España

Vocales Ramon M. Álvarez Halcon

Benjamín Gómez Moliner

Eugenia María Martínez Cueto-Felgueroso

Jesús Souza Troncoso

José Templado González

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, c/ José Gutierrez Abascal 2, 28006 Madrid, España.

CUOTAS PARA 2003:

Socio numerario (en España): 33,06 euros

(en Europa) 36,06 euros

(fuera de Europa): 42,07 euros

Socio estudiante (en España): 18,03 euros

(en el extranjero): 24,04 euros

Socio Familiar: (sin recepcion de revista) 3 euros

Socio Protector: (mínimo) 42,07 euros

Socio Corporativo (en Europa): 42,07 euros

(fuera de Europa): 48,08 euros

INSCRIPCIÓN: 6,01 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.

Members living in foreing countries can deduce 6 euros if paid before 15 April.

Cada socio tiene derecho a recibir anualmente los números de /berus, Reseñas Malacológicas y Noticiarios que

se publiquen.

ÍNDICE

Iberus 21 (2) 2003

CABRAL, J. P. Characterization and multivariate analysis of Patella intermedia, Patella ulyssiponensis

and Patella vulgata from Póvoa de Varzim (Northwest Portugal)

Caracterización y análisis multivariante de Patella intermedia, Patella ulyssiponensis and

Patella vulgata de Póvoa de Varzim (noroeste de Portugal) ici 1-17

SINGH, S. K. AND SINGH, A. Toxic effect of Thevetia peruviana and Alstonia scholaris latices on the

freshwater snail Lymnaea acuminata

Toxicidad de los látex de Thevetia peruviana y Alstonia scholaris sobre el molusco dulceacuí-

cola Eymanaci acuminata a AN 19-27

BOYER, E A sibling species of Gibberula cordorae (de Jong and Coomans, 1988) in the Leeward

Antilles

Una especie gemela de Gibberula cordorae (de Jong and Coomans, 1988) en las Antillas de

SADO O o TO 9-36

SALVINI-PLAWEN, L. v. Contributions to West-Mediterranean Solenogastres (Mollusca) with three

new species

Contribuciones al conocimiento de los Solenogastres (Mollusca) del Mediterráneo occidental,

COM EZES NUEVAS ESPECIOS A 37-60

Díaz Díaz, O. Y LIÑERO-ARANA, l. Poliquetos asociados a [sognomun alatus (Gmelin, 1791)

(Bivalvia: Isognomonidae) en la costa nororiental de Venezuela.

Polychaete worms associated to lsognomun alatus (Gmelin, 1791) (Bivalvia: Isognomonidae)

from. northeastern coast of Venezuela ni a o 61-65

ROLAN, E. AND GOFAS, S. The family Elachisinidae (Mollusca, Rissooidea) in the temperate and

tropical Atlantic

La familia Elachisinidae (Mollusca, Rissooidea) en el Atlántico templado y tropical .... 67-90

HOUART, R. Description of a new muricopsine species (Gastropoda: Muricidae) from the South-

westert: Indian Ocean

Descripción de una nueva especie de muricopsine (Gastropoda: Muricidae) del suroesta del

Oeéaño Indico a ES A 91-98

GARCÍA-ÁLVAREZ, O. AND URGORRI, V. A new species of Phyllomeniidae (Mollusca Solenogas-

tres: Sterrofustia) from the South Shetland Islands, Antarctica

Una nueva especie de Phyllomensidae (Mollusca Solenogastres: Sterrofustia) de las Islas shetland

Al 31m. ALTEA: ¿E NI ANS ES NA 99-107

FORTUNATO, H. Foregut and reproductive tract anatomy of three species of the Strombina-group

(Buccinoidea: Columbellidae)

Anatomía del tubo digestivo y del tracto reproductor en tres especies del grupo Strombina (Buc-

cmordea: Columbelldae) ¿3 Na a 109-122

ROLÁN, E. AND HERNÁNDEZ, J. M. The genus Schwartziella in Senegal (Gastropoda, Rissoidae)

El género Schwartziella en Senegal (Gastropoda, Rissoidae) con. nono. 123-132

LINERO ARANA, l. Y MATA, M. Aspectos bioecológicos de Gemma gemma (Totten, 1894) (Pelecy-

poda: Veneridae) en la laguna de Chacopata, Venezuela

Biological aspects of Gemma gemma (Totten, 1894) (Pelecypoda: Veneridae) on Chacopata

lagoon, Venezudla A TO E ÓN 133-139

ROLAN, E. Y ARCONADA, B. Nueva información sobre Paladilhiopsis septentrionalis (Mollusca, Pro-

sobranchia)

New information on Paladilhiopsis septentrionalis (Mollusca, Prosobranchia) ..... 141-143

ISSN 0212-3010