COMITÉ DE REDACCIÓN

EDITOR

Ángel Guerra Sierra

EDITORES ADJUNTOS

Eugenia M* Martínez Cueto-Felgueroso

Francisco Javier Rocha Valdés

Gonzalo Rodríguez Casero

ComuTé EDITORIAL

Kepa Altonaga Sustacha

Eduardo Angulo Pinedo

Thierry Backeljau

Sigurd v. Boletzky

Jose Castillejo Murillo

Karl Edlinger

José Carlos García Gómez

Edmund Gittenberger

Serge Gofas

Ángel Antonio Luque del Villar

María Yolanda Manga González

Jordi Martinell Callico

Iberus

Revista de la

SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

Instituto de Investigaciones Marinas, CSIC, Vigo, España

Universidad de Oviedo, Oviedo, España

Instituto de Investigaciones Marinas, CSIC, Vigo, España

Universidad de Oviedo, Oviedo, España

Universidad del País Vasco, Bilbao, España

Institut Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica

Laboratoire Arago, Banyuls-surMer, Francia

Universidad de Santiago de Compostela, Santiago de Compostela, España

Naturhistorisches Museum Wien, Austria

Universidad de Sevilla, Sevilla, España

Notionaal Natuurhistorisch Museum, Leiden, Holanda

Muséum National d'Histoire Naturelle, Paris, Francia

Universidad Autónoma de Madrid, Madrid, España

Estación Agrícola Experimental, CSIC, León, España

Universidad de Barcelona, Barcelona, España

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

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

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

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

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

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

Joandoménec Ros ¡ Aragones Universidad de Barcelona, Barcelona, España

María del Carmen Salas Casanovas Universidad de Málaga, Málaga, España

Gerhard Steiner Universitát Wien, Austria

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

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

Anders Warén Swedish Museum of Natural History, Estocolmo, Suecia

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

notas breves. /berus edita un volumen anual que se compone de dos o más números.

INSTRUCCIONES PARA LOS AUTORES

Los manuscritos deben remitirse a: Dr. Ángel Guerra Sierra, Instituto de Investigaciones Marinas (CSIC),

c/ Eduardo Cabello 6, 36208 Vigo, España.

Los trabajos se entregarán por triplicado (original y dos copias). Se recomienda a los autores leer cuidadosa-

mente las normas de publicación que se incluyen en cada número de la revista.

SUBCRIPCIONES

Iberus puede recibirse siendo socio de la Sociedad Española de Malacología, en cualquiera de sus formas, O

mediante intercambio. Aquellos socios que deseen adquirir números atrasados deberán dirigirse al bibliotecario.

Los no socios deberán ponerse en contacto con BACKHUYS PUBLISHERS, P.O. Box 321, 2300 AH

Leiden, The Netherlands. Tel.: +31-71-51 70 208, Fax: +31-71-51 71 856, Correo Electrónico: backhuysCeuronet.nl

PORTADA DE Jberus

Iberus gualterianus (Linnaeus, 1758), una especie emblemática de la península Ibérica, que da nombre a la

revista. Dibujo realizado por Toza.

Iberus

REVISTA DE LA

SOCIEDAD ESPAÑOLA

DE MALACOLOGÍA

Vol. 15 (1) Oviedo, junio 1997

Dep. Leg. B-43072-81

ISSN 0212-3010

Diseño y maquetación: Gonzalo Rodríguez

Impresión: LOREDO, S. L. - Gijón

O Sociedad Española de Malacología Iberus, 15 (1): 1-4, 1997

How many species of Candidula (Gastropoda: Hygromii-

dae) in northern Portugal?

¿Cuantas especies de Candidula (Gastropoda: Hygromiidae) habitan

en el norte de Portugal?

Cristian R. ALTABA*

Recibido el 6-11-1996. Aceptado el 24-IV-1996

ABSTRACT

Two species of Candidula Kobelt, 1871 (Pulmonata: Hygromiidae) endemic to Portugal

are reported from the northern part of the country. C. belemensis (Servain, 1880) is consi-

dered a valid species similar to, but different from the widespread C. intersecta (Poiret,

1801). C. olisippensis (Servain, 1880) is distinctive, having a shell with very small, round

umbilicus, a long spermathecal duct, and a short flagellum.

RESUMEN

Se citan para el norte de Portugal dos especies endémicas en este país pertenecientes al

género Candidula Kobelt, 1871 (Pulmonata: Hygromiidae). C. belemensis (Servain,

1880) es considerada como una especie válida similar, pero diferente, a la ampliamente

distribuidaC. intersecta (Poiret, 1801). C. olisippensis (Servain, 1880) es de fácil distin-

ción, al tener una concha con un ombligo redondeado y pequeño, un largo conducto de

la espermateca y un corto flagelo.

KEY WORDS: Gastropoda, Hygromiidae, Candidula, Portugal, distribution, taxonomy.

PALABRAS CLAVE: Gastropoda, Hygromiidae, Candidula, Portugal, distribución, taxonomía.

INTRODUCTION

The genus Candidula Kobelt, 1871

comprises several species of hygromiid

land snails living in Western Europe. In

the Iberian Peninsula this genus has un-

dergone a remarkable diversification,

particularly along the Atlantic drainages

(ALTONAGA, GÓMEZ, MARTÍN, PRIETO,

PUENTE AND RALLO, 1994). The taxo-

nomy of this genus is complex because

the diagnostic characters of the various

species are not conspicuous, either in

the internal anatomy or in the shells.

The species of Candidula living in Portu-

gal are still poorly known, yet this

country has at least four endemics (AL-

TIMIRA, 1969; GITTENBERGER, 1985, 1993).

This note is a contribution towards a re-

vision of the genus in the Iberian Penin-

sula (Ondina and Altaba, in prep.).

Two little-known, apparently rare

species of Candidula are reported here,

collected during a field trip in April

*Institut Mediterrani d'Estudis Avancgats (CSIC-UIB). Ctra. de Valldemossa, Km 7. 5, 07071 Palma de

Mallorca.

Iber IIIO

1994 in northern Portugal. These species

were described by SERVAIN (1880), and

subsequently ignored for over a century

(NOBRE, 1930, 1941; SErIxas, 1992). Con-

RESULTS

chological and anatomical data indicate

that the species studied here are indeed

distinct taxa. The specimens are kept in

the author's malacological collection.

Family HYGROMIIDAE Tryon, 1866

Genus Candidula Kobelt, 1871

Candidula belemensis (Servain, 1880) (Figs. 1, 2)

C. belemensis was known up to date

only from the original locality (Lisbon)

and three other sites further south (GrT-

TENBERGER, 1993). A single fresh shell

was found at the base of the old city

walls of Valenca do Minho, at the nort-

hern border of Portugal (UTM 29T

NG25; Figs. 1, 2). This locality represents

a considerable extension of the species”

known range. It is likely, however, that

C. belemensis lives also further north, in

Galicia, where specimens having a long

flagellum have been reported as C. inter-

secta by CASTILLEJO (1986).

This species is probably closely rela-

ted to C. intersecta (Poiret, 1801), which

ranges discontinuously throughout

Atlantic Europe, from southern Portugal

to southern Sweden (KERNEY AND CA-

MERON, 1979; GITTENBERGER, 1993; AL-

TONAGA et al., 1994). C. belemensis differs

conchologically from the latter in having

a more depressed shell with a broader

aperture, a wide and slightly eccentric

umbilicus, and a much shallower sculp-

ture of radial ribs and minute spiral

striae (GITTENBERGER, 1993).

Anatomically it differs from NW-Euro-

pean C. intersecta in the relatively longer

flagellum, which is about half the length

of the epiphallus (GITTENBERGER, 1993).

However, populations of C. intersecta from

NW-Iberia have a flagellum shorter than

half the length of the epiphallus, as is

typical of that species (MANGA GONZÁ-

LEZ, 1979). Thus, the available evidence

suggests that C. belemensis is not conspe-

cific with co-occurring C. intersecta.

Candidula olisippensis (Servain, 1880) (Figs. 3-6)

Two subadult specimens were col-

lected under bolders in the vicinity of

the Sanctuary of Sameiro, near Braga

(UTM 29T NE59; Figs. 3, 4). Although

the aperture edge of their shells was still

tender and became damaged, it bears a

conspicuous whitish internal rib. The

radulae have one central, and 22 and 24

lateral teeth. These numbers fall within

the range of other Candidula species

from northwestern Iberia (MANGA GON-

ZÁLEZ, 1979).

The genitalia (still immature) of both

specimens are shown in Figures 5 and 6.

The flagellum is distinct, proximally na-

rrow, and very short, measuring about

1/4 the length of the epiphallus, which is

somewhat shorter and thinner than the

penis. The bursa is small, oblong and in-

distinctly united to its duct, which lies

appressed to the spermoviduct throug-

hout and is exceedingly long, measuring

almost twice the length of the penis and

epiphallus combined. There are no glan-

dulae mucosae, their location being oc-

cupied instead by a moderate swelling

at the proximal end of the vagina. The

dart sac appears partially developed,

being only a large medial swelling of the

vagina with no trace of a dart, but exhi-

biting two large internal longitudinal

folds adjacent to two shallow invagina-

tions, the lower one bordered by a few

papillae. This internal structure can be

interpreted as an immature stage of that

described for other species of Candidula

ALTABA: How many species of Candidula in northern Portugal?

Figures 1-4. Shells of Candidula from northern Portugal. 1, 2: C. belemensís from Valenga do Minho

(CRA 4895); 3, 4: C. olisippensis from Sameiro, near Braga (CRA 4866-1). Scale bar 5 mm.

Figuras 1-4. Conchas de Candidula del norte de Portugal. 1, 2: C. belemensis de Valenga do Minho

(CRA 4895); 3, 4: C. olisippensis de Sameiro, cerca de Braga (CRA 4866-1). Escala 5 mm.

Figures 5, 6. Proximal genitalia of two immature specimens of Candidula olisippensis.

Abbreviations. A: atrium; E: epiphallus; F: flagellum; M: penial retractor muscle; P: penis; S: bursa

copulatrix; SD: duct of the bursa; SO: spermoviduct; V: vagina; VD: vas deferens. Scale bar 2 mm.

Figuras 5, 6. Genitalia proximal de dos especimenes inmaduros de Candidula olisippensis.

Abreviaturas. A: atrio; E: epifalo; E: flagelo; M: músculo retractor peneal; P: pene; S: bursa copulatrix;

SD: conducto de la bursa; SO: espermoviducto; V: vagina; VD: vaso deferente. Escala 2 mm.

Iberus, 15 (1), 1997

(HAUSDORE, 1988, 1991), confirming the

generic assignation by GITTENBERGER

(1993), who examined a dried specimen

in poor condition.

Several nominal species described

by LOCARD (1899) are probably sy-

nonyms of Helix olisippensis Servain

1880. The general shape of the shell in

this species (or species complex) is fairly

variable, yet it is always fairly thin, with

ACKNOWLEDGEMENTS

My wife Catalina Ponsell helped in

the field and provided the necessary

conditions for the preparation of the

manuscript. We are indebted to Or-

lando Moreira and his family for in-

comparable hospitality in Portugal and

BIBLIOGRAPHY

ALTIMIRA, C., 1969. Notas malacológicas. Pu-

blicaciones del Instituto de Biología Aplicada,

46: 91-113.

ALTONAGA, K.; GÓMEZ, B.; MARTÍN, R.; PRIETO,

C.R.; PUENTE, A.I. AND RALLO, A., 1994. Es-

tudio faunístico y biogeográfico de los moluscos

terrestres del norte de la Península Ibérica. Eusko

Legebiltzarra, Vitoria-Gasteiz, 505 pp.

CASTILLEJO, J., 1986. Caracoles terrestres de Ga-

licia. Familia Helicidae (Gastropoda, Pul-

monata). Monografías de la Universidad de San-

tiago de Compostela, 122: 1-65.

GITTENBERGER, E., 1985. The taxonomic status

of Xeroplexa Monterosato, 1892 (Pulmonata:

Helicidae: Helicellinae), a surprise. Iberus, 5:

59-62.

GITTENBERGER, E., 1993. Digging in the grave-

yard of synonymy, in search of Portuguese

species of Candidula Kobelt, 1871 (Mollusca:

Gastropoda Pulmonata: Hygromiidae). Zo-

ologische Meddedelingen, 67 (17): 283-293.

HAUSDORE, B., 1988. Zur Kenntnis der syste-

matischen Beziehungen einiger Taxa der He-

licellinae Ihering 1909. Archiv fiir Mollusken-

kunde, 119 (1/3): 9-37.

HAUSDORE, B., 1991. Uber zwei Candidula-Ar-

ten von der súdlichen Balkanhalbinsel (Gas-

tropoda: Hygromiidae). Archiv fur Mollus-

kenkunde, 120 (4/6): 119-129.

rounded or slightly angular periphery,

very narrow roundish umbilicus, spire

formed by flattened whorls separated

by an indented suture, radial sculpture

moderately developed, and microspiral

striae well marked (GITTENBERGER,

1993). Candidula olisippensis is appa-

rently endemic to Portugal. The current

finding represents the northern limit of

its known range.

Galicia. Paz Ondina, Carlos Prieto, Án-

gel A. Luque, and two anonymous

reviewers kindly provided helpful com-

ments. This work was partially suppor-

ted by Research Project PB93-0055 of

DEE:

KERNEY, M. P. AND CAMERON, R. A. D.,, 1979.

A field guide to the land snails of Britain and

north-west Europe. Collins, London, 288 pp.

LOCARD, A., 1899. Conchyliologie Portugaise.

Les coquilles terrestres, des eaux douces et

saumátres. Archives du Muséum de Lyon, 7

(1): i-xi, 1-303.

MANGA GONZALEZ, Y., 1979. Sobre las espe-

cies del género Candidula Kobelt, 1871, (Gas-

tropoda, Stylommatophora) en la provincia

de León. Revista Ibérica de Parasitología, 79: 455-

465 + 1 pl.

NOBRE, A,, 1930. Moluscos terrestres, fluviais e das

aguas salobras de Portugal. Porto, 259 pp, 18 pls.

NOBRE, A., 1941. Moluscos terrestres e fluviais.

Fauna de Portugal, 2. Memórias e Estudos do

Muséu de Zoología da Universidade da Coimbra,

124: 1-277, pls. i-iv, 1-30.

SEIxAS, M., 1992. Gasterópodes terrestres da

coleccao do Museu Bocage. Arquivos do Mu-

séu Bocage, Nova Série, 2 (10): 155-255.

SERVAIN, G., 1880. Étude sur les mollusques re-

cueillis en Espagne et en Portugal. Saint Ger-

main, Paris, 172 pp.

O Sociedad Española de Malacología Iberus, 151% 9=-22, 1997

Nuevos datos sobre la distribución de la superfamilia Heli-

coidea Rafinesque, 1815 (Gastropoda, Pulmonata, Stylom-

matophora) en el oeste de Galicia

New records about the distribution of the superfamily Helicoidea

Rafinesque, 1815 (Gastropoda, Pulmonata, Stylommatophora) in

the West of Galicia

Paz ONDINA, Jesús HERMIDA y Adolfo OUTEIRO*

Recibido el 21-IV-1996.Aceptado el 30-IV-1996

RESUMEN

En este trabajo se realiza un estudio faunístico de las especies de la superfamilia Helicoi-

dea Rafinesque, 1815, encontradas en el oeste de Galicia (provincias de A Coruña y

Pontevedra). Para cada especie se incluyen citas previas y localidades de captura con las

coordenadas U.T.M. 10x10 km. Teniendo en cuenta nuestros hallazgos y los de los auto-

res consultados en la bibliografía se han elaborado los mapas de distribución correspon-

dientes, en sistema U.T.M. de 10x10 km. Se cita por primera vez para el área de estudio

Mengoana brigantina (da Silva Mengo, 1867).

ABSTRACT

A faunistic study of species of superfamily Helicoidea Rafinesque, 1815, in the west of

Galicia (La Coruña and Pontevedra provinces) has been realized. For each species the

previous records and the coordinates U.T.M. 10x10 km of the localities where the species

have been found, are included. Taking our own findings into consideration, and the data

from bibliography, a map showing the distribution of each species has been drawn up,

using U.T.M. 10x10 km system. Mengoana brigantina [da Silva Mengo, 1867) is recor-

ded for the first time in this area.

PALABRAS CLAVE: Gastropoda, Pulmonata, Helicoidea, distribución, Galicia.

KEY WORDS: Gastropoda, Pulmonata, Helicoidea, distribution, Galicia.

INTRODUCCIÓN

El conocimiento de la fauna de

moluscos terrestres de la Península

Ibérica presentaba, hasta no hace mucho

tiempo, un gran retraso respecto a

Europa. Fue a finales del siglo XIX y

principios del XX cuando se desarrolló

más intensamente la actividad en este

campo, y cuando comienza a otorgár-

sele validez taxonómica a ciertas estruc-

turas anatómicas, especialmente el

*Departamento de Biología Animal. Facultad de Biología. Universidad de Santiago de Compostela. 15706

Santiago de Compostela. La Coruña, España.

Iberus, 15 (1), 1997

aparato genital, tras comprobar que

especies con conchas muy parecidas

albergaban animales de organización

anatómica diferente.

También en Galicia se inician, en este

período, los estudios malacológicos con

las aportaciones de autores como

GRAELLS (1846), SEOANE (1866), MACHO

VELADO (1871, 1878) e HIDALGO (1875).

Posteriormente otros autores han reali-

zado estudios en zonas geográficas con-

cretas como ALTIMIRA (1969); SACCHI y

VIOLANI (1977) y ROLÁN y OTERO (1988).

Con mención específica a los helícidos

cabe destacar el trabajo de CASTILLEJO

(1981; 1986), que cubre gran parte del

territorio gallego, recogiendo muestras

en un total de 87 cuadrículas de 10x10

km U.T.M.

A pesar de estos estudios existen dis-

continuidades en la distribución de gran

parte de las especies, motivo por el cual

con el presente estudio queremos contri-

buir a ampliar el conocimiento faunís-

tico de los gasterópodos terrestres perte-

necientes a la superfamilia Helicoidea

Rafinesque, 1815, aportando nuevos

datos de distribución para las provincias

de A Coruña y Pontevedra.

MATERIAL Y METODOS

Durante el período 1986-1989 se ha

recolectado material malacológico pro-

cedente de las 176 cuadrículas U.T.M. de

10x10 km, en las que se han dividido las

provincias de A Coruña y Pontevedra

(Fig. la). En las distintas cuadrículas

visitadas se han examinado detenida-

mente, de día y de noche, los distintos

hábitats donde suelen resguardarse,

recogiéndose además, muestras de suelo

y hojarasca de distintos biotopos, que

posteriormente se lavaron y tamizaron

para separar los ejemplares.

Todos los gasterópodos capturados

se sometieron al proceso habitual de

muerte por anoxia sumergiéndolos en

agua, para facilitar de este modo su

disección, conservándose posterior-

mente en alcohol de 70*.

A partir de los datos obtenidos, se

han elaborado los mapas de distribución

de cada especie (Figs. 1b-g, Figs. 2a-1) en

cuadrículas U.T.M. de 10x10 km, indi-

cándose tanto las localidades aportadas

en este trabajo (+) como las procedentes

de la bibliografía (*), así como las locali-

dades en las que se encontraron única-

mente conchas vacías (O).

La colección malacológica se encuen-

tra depositada en el Departamento de

Biología Animal (Facultad de Biología,

Universidad de Santiago de Compos-

tela).

RESULTADOS

Se han identificado un total de 6873

ejemplares pertenecientes a 18 especies.

Para cada especie se incluyen los

siguientes apartados: citas previas,

material estudiado (en el que se indica

el número de ejemplares capturados en

cada cuadrícula y aquellas localidades

donde se encontraron conchas vacías),

un breve resumen de su distribución

geográfica y algunas observaciones de

interés para aquellas especies en que lo

hemos considerado necesario. El listado

de las localidades junto a su correspon-

diente código, coordenadas U.T.M.,

Ayuntamiento y fecha de muestreo se

pueden observar en la Tabla 1.

Superfamilia HELICOIDEA Rafinesque, 1815

Familia XANTHONYCHIDAE Strebel y Pfeffer, 1880

Elona quimperiana (Férussac, 1821) (Fig. 1B)

Citas previas: CAZIOT (1915) como H. quimperiana; CASTILLEJO (1986); OTERO y TRIGO (1989).

Material examinado (107 ejemplares). 39: 1; 58: 1; 93: 9; 94: 3; 100: 7; 102: 1; 103: 8; 104: 12; 125: 9;

130: 1; 141: 3; 143: 2; 144: 1; 145: 5; 146: 15; 147: 14; 148: 4; 156: 6; 157: 5.

ONDINA £7 4Z.: Distribución de la superfamilia Helicoidea en el oeste de Galicia

Tabla I. Listado de las localidades, provincia, U.T.M. en 10 x 10 Km y fecha de recogida de las

muestras, junto a su correspondiente código. C: A Coruña; L: Lugo; O: Ourense; P: Pontevedra.

Table I. List of the locations, provinces, U.I.M. 10x10 Km and sampling date, with its code. C: A

Coruña; L: Lugo; O: Ourense; P: Pontevedra.

Código

O0XD0 NXNOo0hb0N —

Localidad /Ayuntamiento

Caldelas de Tui, Tui

Cristelos, Tomiño

Louredo, Mos

Coruxo, Vigo

Pintos, Pontevedra

Dorrón, Sanxenxo

A Xesta, A Lama

O Covelo, O Covelo

Caldas de Reis, Caldas de Reis

Sobrido, Ribeira

Noal, Porto do Son

Sionlla, Santiago

Negreira, Negreira

Ponte Sarandón, Vedra

Arca, A Estrada

Ferreiros, Vila de Cruces

Prado, Lalín

Santa María, Rodeiro

Amance, Golada

Herbón, Padrón

Budiño, Porriño

Piñeiro, O Covelo

Salvaterra, Salvaterra do Miño

Arbo, Arbo

loureza, Oia

Baredo, Baiona

Amoedo, Pazos de Borbén

Magdalena, Cangas

Leiro, Ribadumia

San Vicente do Grove, O Grove

Serrapio, Cerdedo

Nigoi, A Estrada

Vilatuxe, Lalín

lagoa, Dozón

Barrio, Vila de Cruces

Ventosa, Golada

Catoira, Catoira

Boiro, Boiro

Lapido, Ames

Sergude, Boqueixón

Costa de Mougás, Oia

Camposancos, A Guarda

Provincia

ADA OO A A A A AS AAA A A SAO ASISTAN O O) A A a a) e) A) a) A) a)

Coord. UTM

29TNG35

29TNG15

29TNG37

29TNG17

29TNG39

29TNG19

29TNG59

29TNG57

29TNH21

29TNHO1

29TNHO3

29TNHA5

29TNH25

29TNH43

29TNH41

29TNHÓ64

29TNH62

29TNH82

29TNH84

29TNH23

29TNG36

29TNG58

29TNGA45

29TNG56

29TNG14

29TNG16

29TNG38

29TNG18

29TNH20

29TNHOO

29TNH40

29TNH42

29TNH61

29TNH81

29TNH63

29TNH83

29TNH22

29TNHO2

29TNH24

29TNH44

29TNGO5

29TNG13

Fecha

26-09-86

27-09-86

28-09-86

29-09-86

30-09-86

01-10-86

02-10-86

03-10-86

04-10-86

05-10-86

06-10-86

04-12-86

05-12-86

06-12-86

07-12-86

08-12-86

09-12-86

10-12-86

11-12-86

12-12-86

13-12-86

10-04-87

11-04-87

Iberus, 15 (1), 1997

Tabla I. Continuación.

Table I. Continuation.

Código Localidad/Ayuntamiento Provincia Coord. UTM

44 Mondariz, Mondariz P 29TNG47

A5 Lavadores, Vigo P 291NG27

46 Pontecaldelas, Pontecaldelas p 29TNG49

A7 Postemirón, Vilaboa P 29TNG29

48 Triñnáns, Boiro (E 29TNH11

49 Oleiros, Ribeira S 29TMH91

50 Portobravo, Lousame a 29TNH13

51 César, Caldas de Reis P 29TNH31

2 Carcacía, Padrón E 29TNH33

53 Carballeda, Lalín P 29TNH72

54 Baíña, A Golada P 29TNH74

DS) Quintás, Touro E 29TNH54

56 Rellas, Silleda Cc 291NH52

7 Louro, Muros € 29TMH93

58 Ordoeste, A Baña E 29TNH15

DY A Peregrina, Santiago € 291NH35

60 Illas Cíes, Vigo P 29TNGO7

61 Barrantes, Ribadumia P 29TNH10

62 Fontáns, Barro Pp 29TNH30

63 Forcarei, Forcarei P 29TNH5 1

64 Lamela, Silleda P 29TNH53

65 Muimenta, Lalín P 29TNH73

66 A Xesta, Lalín P 29TNH7 1

67 Baroña, Porto do Son a 29TMH92

68 Pontenafonso, Noia € 29TNH14

69 Carreira, Ribeira E 29TMH9O

70 Bures, Catoira € 29TNH 12

71 Santa Mariña de Barcala, A Estrada. P 291TNH32

Ez Oitavén, Fornelos de Montes P 29TNG48

US Ponteareas, Ponteareas P 29TNG46

7A Moaña, Moaña P 29TNG28

75 Mosende, Porriño P 29TNG26

76 Tomiño, Tomiño P 29TNG24

7 Chavella, Oia P 29TNGO4

78 Cabo Silleiro, Baiona P 29TNGO06

79 Santiago, Santiago € 29TNH34

80 Esmorode, Santa Comba E 29TNH16

81 Serra de Outes, Serra de Outes E 29TNHOA4

82 Quilmas, Carnota e 291MH84

83 Vilela, Muxía E 29TMH76

84 Berdoias, Vimianzo e 29TMH96

85 Nande, Laxe € 29TMH98

86 As Tarandeiras, Coristanco E 29TNH18

Fecha

12-04-87

13-04-87

14-04-87

15-04-87

16-04-87

17-04-87

18-04-87

19-04-87

20-04-87

12-05-87

27-06-87

28-06-87

29-06-87

01-07-87

02-07-87

03-07-87

04-07-87

05-07-87

06-07-87

07-07-87

08-07-87

09-10-87

10-10-87

11-10-87

12-10-87

ONDINA £7 4Z.: Distribución de la superfamilia Helicoidea en el oeste de Galicia

Tabla I. Continuación.

Table I. Continuation.

Código Localidad/Ayuntamiento

88 Rial, Trazo

89 Zas de Reis, Melide

90 Padreiro, Curtis

91 Xestal, Irixoa

92 Saa, As Pontes

93 Freires, Ortigueira

94 Regoa, Cedeira

95 Esmelle, Ferrol

96 Bouzarredonda, Neda

97 Sigrás, Cambre

98 Abellá, Frades

99 Pedrouzo, O Pino

100 Viñas, Paderne

101 Neda, Neda

102 Ponte de Mera, Ortigueira

103 Mañón, Mañón

104 Recemel, Somozas

105 Boliqueiras, As Pontes

106 Salto do Conexo, Monfero

107 Monte do Arco, Curtis

108 Rexidoira, Cesuras

109 Arzúa, Arzúa

110 Furelos, Melide

ME Vila da Igrexa, Cerceda

112 Parada, Ordes

118 Pedrafigueira, Carnota

114 Cabo Fisterra, Fisterra

MS Ozón, Muxía

116 Areosa, Vimianzo

IN Ponteceso, Ponteceso

118 Cances, Carballo

119 Bembibre, Val do Dubra

120 Sobrado dos Monxes, Sobrado

121 A Castellana, Aranga

1122 A Capela, A Capela

123 San Sadurniño, San Sadurniño

124 O Ermo, Ortigueira

125 Sismundi, Ortigueira

126 Raxón, Ferrol

127 Mabegondo, Abegondo

128 Carnoedo, Sada

129 Poulo, Ordes

130 Río, Cerceda

Provincia

DKOOOOOOoOS0OoOO0O0o0o00 0000 O0O0oO0O0o0 0000000000000 000000O00O

Coord. UTM

29TNH36

29TNH75

291NH77

291NH79

29TNJ91

29T1NJ93

29 T1NJ73

29TNJ51

29 1INJ7 1

29TNH59

29TNH57

29TNH55

29TNHÓ9

29TNJÓ 1

29T1NJ83

29TPJO3

29TNJ8 1

29TNJ90

29TNH89

291NH87

29TNH67

29TNHÓ5

29TNH85

29TNH48

29TNH46

291MH94

291MH74

291MH86

29TNHO6

29TNHO8

291NH28

29TNH26

29TNH7Ó

291NH78

29T1NJ7O

29TNJ72

29TNJ92

291NJ94

29TNJ52

29TNH58

29TNJ50

291NH5Ó

29TNH37

Fecha

13-10-87

19-10-87

20-10-87

21-10-87

22-10-87

23-10-87

24-10-87

18-01-88

19-01-88

20-01-88

21-01-88

22-01-88

23-01-88

24-01-88

26-01-88

27-01-88

28-01-88

29-01-88

25-03-88

26-03-88

27-03-88

28-03-88

29-03-88

30-03-88

31-03-88

01-04-88

Iberus, 15 (1), 1997

Tabla I. Continuación.

Table I. Continuation.

Código

132

188

134

139

136

137

138

139

140

141

142

143

144

145

146

147

148

149

150

151

152

153

154

159

156

157%

158

159

160

161

162

163

164

165

166

167

168

169

170

171

172

174

173

Localidad/Ayuntamiento

Provincia

Agualada, Coristanco

Buño, Malpica

Santa Mariña, Camariñas

Ponte do Porto, Camariñas

Fisterra, Fisterra

Logoso, Dumbría

Arceo, Boimorto

Lagoa de Sobrado, Sobrado

Aranga, Aranga

Burricios, Oza dos Ríos

Faeira, As Pontes

As Somozas, Somozas

Grañas, Mañón

Loiba, Ortigueira

Serra da Capelada, Ortigueira

Sedes, Narón

laraxe, Cabanas

ledoño, Culleredo

Leira, Ordes

Razo, Carballo

Canosa, Carballo

Corme, Ponteceso

Baio, Zas

Moraime, Muxía

lobelos, Cée

A Picota, Mazaricos

Setados, As Neves

Crecente, Crecente

Paredes, A Cañiza

Pardesoa, Forcarei

Lira, Carnota

Couto, Rodeiro

Vilafrío, Rodeiro

San Pedro de Visma, A Coruña

Sucadío, As Pontes

Punta Candelaria, Cedeira

Illa de Ons, Bueu

Momán, Xermade

Vilaverde, Ribadavia

Codesas, Forcarei

Cabo da Voutra, Muxía

Laxe, Chantada

Punta Frouxeira, Valdoviño

Illa de Ons (Sur), Bueu

UR OAOR LO O AOS LIO ADO OOoOOO.OOOOOOOOA

Coord. UTM

29TNH17

29TNH19

291MH88

291MH97

29TMH75

29TMH95

29TNHÓ66

291NH8ó

29TNH88

29TNHÓ8

291NJ80

291NJ82

29TPJO2

29TPJO4

29T1NJ84

29TNJÓ2

29TNJ6O

291NH49

291NH47

291NH29

29TNH27

29TNHO9

29TNHO7

291MH87

291MH85

29TNHO5

29TNG55

29TNG66

291NGÓ67

29TNH50

291MH83

29TNH93

291NH92

29TNJ40

29TPJO1

29T1NJ74

29TNGO9

291NH99

29TNG68

29TNHÓO

291MH77

29TNH91

29TNJÓ3

29TNGO9

Fecha

02-04-88

03-04-88

04-04-88

05-04-88

30-06-88

01-07-88

02-07-88

03-07-88

04-07-88

05-07-88

06-07-88

07-07-88

08-07-88

09-07-88

10-07-88

17-03-89

18-03-89

19-03-89

22-04-89

23-05-89

30-05-89

08-06-89

28-03-88

31-05-89

08-06-89

18-03-89

19-03-89

22-04-89

23-05-89

10-05-89

31-05-89

ONDINA £7 AL.: Distribución de la superfamilia Helicoidea en el oeste de Galicia

Distribución geográfica: Se trata de

una especie de distribución atlántico-eu-

ropea (KERNEY, CAMERON Y JUNGBLUTH,

1983). Fuera de la Península Ibérica su

área de distribución se restringe a la Bre-.

taña francesa y al País Vasco francés

(GERMAIN, 1930).

En la Península se limita a la costa sep-

tentrional. Está citada en el norte desde

Galicia hasta el País Vasco (HIDALGO, 1875;

GITTENBERGER, 1979; LARRAZ y JORDANA,

1984; CASTILLEJO, 1986; HERMIDA, OUTEIRO

Y RODRÍGUEZ, 1992; PUENTE y PRIETO, 1992;

LARRAZ y EQUISOAIN, 1993).

Familia HYGROMIIDAE Tryon, 1866

Candidula intersecta (Poiret, 1801) (Fig. 1C)

Citas previas: MACHO VELADO (1871) como H. caperata; HIDALGO (1875, 1890) como H. caperata;

SACCHI y VIOLANI (1977) como Helicella caperata; CASTILLEJO (1986).

Material examinado (93 ejemplares). 6: 1; 11: 3; 19: 1; 20: 1; 30: 1; 35: 10; 37: 1; 38: 1; 42: 3; 46: 5; 47:

2; 48: 4; 49: 6; 57: 1; 60: 6; 69: 3; 74: 3; 77: 2; 78: 5; 79: 4; 82: 1; 83: 8; 93: 6; 113: 3; 114: 4; 136: 2; 137: 1;

151: 3; 156: 2; 171: 1.

Localidades con conchas vacías: 153.

Distribución geográfica: Su área de

distribución se enmarca en el oeste de

Europa (KERNEY ET AL., 1983).

En la Península se extiende por dos

áreas, una en el sector norte oriental, el

País Vasco y Navarra, y la otra en el tercio

oeste, desde Galicia y centro de León

hasta el Algarve, con algunas citas hacia

el centro peninsular como Segovia

(HIDALGO, 1875; NOBRE, 1941; RAMOS y

APARICIO, 1985a; (CASTILLEJO, 1986;

HERMIDA ET AL., 1992). En Asturias,

desde la cita de ALTIMIRA (1969), no ha

vuelto a ser encontrada (OJEA y ANADÓN,

1983; HERMIDA ET AL., 1992), y tampoco

se tiene constancia de ella en Cantabria.

Cernuella (Cernuella) virgata (da Costa, 1778) (Fig. 1D)

Citas previas: SACCHI y VIOLANI (1977); CASTILLEJO (1986).

Material examinado (3 ejemplares). 6: 3.

Distribución geográfica: Especie

mediterránea, ampliamente distribuida

por el sur y oeste de Europa (KERNEY ET

AL., 1983).

Siendo común en el área peninsular y

en las islas Baleares, es menos frecuente

en la franja norte que comprende desde

el valle del Tajo hasta el País Vasco,

donde se convierte en una especie litoral

(NOBRE, 1941; ORTIZ DE ZÁRATE y ORTIZ

DE ZARATE, 1949; Seixas, 1976; PAUL,

1982; MANGA, 1983; LARRAZ y JORDANA,

1984; RAMOS y APARICIO, 1985b; CASTI-

LLEJO, 1986; APARICIO, 1986; ROBLES,

ISSO PEUENTESyARRIELO Ugg L9O92:

HERMIDA ET AL., 1992; LARRAZ y EQUISO-

AIN, 1993; ALTONAGA, GÓMEZ, MARTÍN,

PRIETO, PUENTE Y RALLO, 1994).

Helicella (Helicella) itala (Linneo, 1758) (Fig. 1E)

Citas previas: MACHO VELADO (1871) como H. ericetorum; HIDALGO (1875) como H. ericetorum;

CASTILLEJO (1986); OTERO y TRIGO (1989).

Material examinado (32 ejemplares). 134: 10; 151: 20; 173: 1; 174: 1.

Distribución geográfica: Se trata de

una especie distribuida por el oeste de

Europa (KERNEY ET AL., 1983).

En la Península Ibérica está presente

en la mitad norte, siendo sus citas más es-

casas en Galicia (HIDALGO, 1875; HAAS,

Maira IMA

1929; ORTIZ DE ZÁRATE y ORTIZ DE ZÁ-

RATE, 1949; MANGA, 1983, APARICIO, 1986;

CASTILLEJO, 1986; HERMIDA ET AL., 1992;

LARRAZ y EQUISOAIN, 1993; ALTONAGA ET

AL., 1994). En el área de estudio la hemos

hallado únicamente en zonas del litoral.

Xerotricha apicina (Lamarck, 1822) (Fig. 1F)

Citas previas: MACHO VELADO (1871) como H. apicina; HIDALGO (1875, 1890) como H. apicina;

ALTIMIRA (1969); SACCHI y VIOLANI (1977) como Helicella apicina; CASTILLEJO (1986).

Material examinado (3 ejemplares). 26: 3.

Distribución geográfica: Es una

especie con un área de distribución

mediterránea (KERNEY ET AL., 1983).

En la Península presenta una distri-

bución costera bastante fragmentada,

encontrándose en el norte unicamente

en algunas localidades del litoral de

Galicia (ALTIMIRA, 1969; SACCHI y

VIOLANI, 1977; CASTILLEJO, 1986), Can-

tabria y Asturias (PUENTE y PRIETO,

1992). En el litoral mediterráneo está

citada en Cataluña (Haas, 1929; ALTI-

MIRA, 1969), Valencia (ROBLES, 1990);

Málaga (ALTONAGA ET AL., 1994), conti-

nuando desde ahí, de una forma más

regular, por el cuadrante suroccidental

hasta Aveiro en Portugal (SERVAIN,

1880; LOCARD, 1899; HIDALGO, 1875;

NOBRE, 1941; RAMOS y APARICIO, 1985a)

y hacia el interior hasta Badajoz.

GASULL (1965) también la cita abun-

dantemente en las Islas Baleares.

Cochlicella acuta (Muller, 1774) (Fig. 1G)

Citas previas: MACHO VELADO (1871) como Bulimus acutus; HIDALGO (1875) como B. acutus;

HIDALGO (1890) como H. acuta; ALTIMIRA (1969); SACCHI y VIOLANI (1977); CASTILLEJO (1986);

OTERO y TRIGO (1989).

Material examinado (99 ejemplares). 4: 8; 6: 8; 11: 1; 26: 9; 49: 3; 57: 4; 60: 2; 69: 4; 82: 3; 83: 8; 85:

12; 93: 5; 113: 6; 136: 10; 151: 6; 156: 4; 168: 3; 173: 3.

Distribución geográfica: Distribuida

por la zona mediterránea y atlántica de

Europa (BOATO, BODON Y GIUSTI, 1982).

Aparece practicamente en toda la

costa ibérica e Islas Baleares (NOBRE,

1941; ORTIZ DE ZARATE y ORTIZ DE

BEcH, 1990; MARTÍNEZ-ORTÍ, MARTÍNEZ-

LÓPEZ, ROBLES Y RODRÍGUEZ BABÍO,

1990) penetrando en ocasiones hacia el

interior siguiendo los valles fluviales.

Está citada en puntos del interior como

Salamanca (HERMIDA ET AL., 1992),

ZARATE, 1949; SACCcHI, 1954; GASULL,

1965; ALTIMIRA, 1969; CASTILLEJO, 1986;

Huesca y Zaragoza (PUENTE y PRIETO,

1992).

Cochlicella barbara (Linneo, 1758) (Fig. 2A)

Citas previas: MACHO VELADO (1871) como B. ventrosus; HIDALGO (1875) como B. ventrosus; ALTI-

MIRA (1969) como C. ventricosa; SACCHI y VIOLANI (1977) como C. ventricosa; CASTILLEJO (1986)

como C. ventricosa; OTERO y TRIGO (1989) como C. ventricosa.

Material examinado (704 ejemplares). 1: 3; 2: 1; 3: 2; 4: 6; 5: 4; 6: 17; 9: 16; 11: 12; 21: 1; 24: 3; 25: 7; 26:

218; 27: 5; 29: 2; 30: 1; 37: 1; 38: 6; 43: 2; 44: 1; 47: 8; 48: 20; 49: 82; 50: 1; 51: 1; 52: 4; 57: 8; 60: 4; 61: 5; 68:

1; 69: 73; 70: 2; 73: 15; 74: 12; 76: 16; 81: 3; 82: 3; 83: 2; 85: 8; 86: 2; 93: 6; 94: 17; 95: 1; 97: 22; 113: 19; 117:

1; 118: 2; 126: 1; 128: 1; 132: 6; 133: 7; 135: 1; 136: 6; 148: 5; 149: 7; 151: 6; 156: 4; 168: 12; 174: 2.

Localidades con conchas vacías: 127.

Distribución geográfica: Especie con

un amplio rango de distribución en el área

mediterránea y a lo largo de las costas del

oeste de Europa (BACKHUYS, 1975).

112

ONDINA £7 4Z.: Distribución de la superfamilia Helicoidea en el oeste de Galicia

16 |

ana

aan

OÍ

he)

Figura 1. A: Area de estudio en la Península Ibérica. B-G: Mapas de distribución. B: Elona quimpe-

riana, C: Candidula intersecta, D: Cernuella virgata; E: Helicella itala, E: Xerotricha apicina; G:

Cochlicella acuta. Localidades citadas en este trabajo (+); procedentes de la bibliografía (*); única-

mente encontradas conchas vacías (O).

Figure 1: A: The study area in the Iberian Peninsula. B-G: Distribution maps. B: Elona quimperiana;

C: Candidula intersecta; D: Cernuella virgata; £: Helicella itala; F: Xerotricha apicina; G: Cochli-

cella acuta. Localities reported in this paper (+); bibliographic records (*); only shells found (O).

Iberus, 15 (1), 1997

En la Península presenta una distri-

bución bastante amplia, y aunque

aparece con mayor frecuencia en el

litoral, citándose en casi todas las pro-

vincias estudiadas (BOFILL y HAAS,

1920a; NOBRE, 1941; ALTIMIRA, 1969;

OJEA y ANADÓN, 1983; RAMOS y APARI-

CIO, 1985b, CASTILLEJO, 1986; MARTÍNEZ-

ORTÍ ET AL., 1990; PUENTE y PRIETO, 1991,

1992; HERMIDA ET AL., 1992; LARRAZ y

EQUISOAIN, 1993; PAREJO ET AL., 1993b).

También se ha encontrado en las islas

Baleares (GASULL, 1965; PAUL, 1982). En

la zona de estudio está ligada a la franja

litoral, aunque penetra más hacia el inte-

rior que C. acuta y C. conotdea.

Cochlicella conoidea (Draparnaud, 1801) (Fig. 2B)

Citas previas: MACHO VELADO (1871) como B. pring1; HIDALGO (1875) como B. pring1; HIDALGO

(1890) como H. conoidea; SACCHI y VIOLANI (1977); OTERO y TRIGO (1989).

Material examinado (79 ejemplares). 4: 9; 6: 8; 49: 3; 57: 4; 60: 1; 82: 4, 83: 20; 113: 4; 136: 20; 156: 6.

Distribución geográfica: Especie

común a lo largo de la costa mediterrá-

nea, desde los Pirineos orientales hasta

los Alpes marítimos (KERNEY ET AL.,

1983)

En la Península se encuentra en el li-

toral mediterráneo, aunque también está

citada en la costa atlántica portuguesa y

gallega, y aisladamente y con escasos

ejemplares en el interior (LOCARD, 1899;

HAas, 1929; NOBRE, 1941; SaccHi, 1954;

GASULL, 1965; ALTIMIRA, 1969; GASULL,

1975; SACCHI y VIOLANI, 1977; RAMOS y

APARICIO, 1985b; CASTILLEJO, 1986; MAR-

TÍNEZ-ORTÍ ET AL., 1990). La única cita

existente en el litoral cantábrico dada

por ORTIZ DE ZARATE y ORTIZ DE ZÁ-

RATE (1949) ha sido asignada por distin-

tos autores a C. barbara (PUENTE y

PRIETO, 1992).

Ashfordia granulata (Alder, 1830) (Fig. 2C)

Citas previas: MACHO VELADO (1871) como H. sericea; HIDALGO (1875) como H. sericea; CASTI-

LLEJO (1986) como Monacha (Ashfordia) granulata.

Material examinado (151 ejemplares). 6: 1; 35: 6; 53: 1; 69: 3; 79: 24; 83: 1; 90: 5; 93: 5; 94: 23; 95: 13;

96: 1; 97: 14; 102: 11; 115: 2; 117: 3; 118: 1; 120: 3; 135: 22; 143: 1; 145: 1; 149: 2; 153: 2; 155: 3; 165: 3.

Distribución geográfica: Se distri-

buye por el oeste de Europa (KERNEY ET

AL., 1983).

En la Península Ibérica su área de

distribución comprende desde Galicia

hasta el extremo occidental de Vizcaya,

penetrando hacia León (MANGA, 1983;

ANADÓN y OJEA, 1984; HOLYOAK y SED-

DON, 1985; CASTILLEJO, 1986; HERMIDA

ET AL., 1992; PUENTE y PRIETO, 1992).

Zenobiella subrufescens (Miller, 1822) (Fig. 2D)

Citas previas: CASTILLEJO (1986) como Monacha (Zenobiella) subrufescens.

Material examinado (34 ejemplares). 39: 1; 40: 1; 50: 4; 68: 1; 79: 1; 87: 5; 90: 2; 93: 3; 94: 2; 97: 1;

102: 2; 108: 1; 111: 1; 116: 1; 117: 5; 118: 1; 128: 1; 162: 1.

Localidades con conchas vacías: 13, 58.

Distribución geográfica: Se extiende

por el oeste europeo (KERNEY ET AL.,

1983). Según GERMAIN (1930) no se aleja

de las regiones de influencia marítima.

En la Península Ibérica está citada

unicamente en la franja norte que com-

prende desde Galicia hasta el País

Vasco y Navarra (OJEAa y ANADÓN,

1983; CASTILLEJO, 1986, HERMIDA ET AL.,

1992; PUENTE y PRIETO, 1992; LARRAZ y

EQUISOAIN, 1993; ALTONAGA ET AL.,

1994).

ONDINA £7 4Z.: Distribución de la superfamilia Helicoidea en el oeste de Galicia

Portugala inchoata (Morelet, 1845) (Fig. 2E)

Citas previas: MACHO VELADO (1871) como H. inchoata; HIDALGO (1875, 1890) como H. inchoata;

GITTENBERGER (1980); SAccHI (1981) como Monachoides inchoatus; CASTILLEJO (1986); OTERO y

TRIGO (1989).

Material examinado (928 ejemplares). 1: 11; 2: 17; 3:

13: 1; 14: 2; 16: 16; 17: 3; 18: 2; 19: 5; 20: 4; 21: 19; 22:

30: 3; 31: 4; 32: 3; 33: 1; 35: 10; 36: 3; 37: 1; 38: 8; 39: 16;

47: 9; 48: 17; 49: 3; 50: 12; 51: 1; 52: 8; 53: 2; do 6; 55: 5;

63: 1; 64: 6; 65: 5; 67: 8; 68: 4; 69: 6; 70: 9; 71: 2; 72: 5; 73:

89:

;4: 7; 5: 7; 6: 20; 8: 5; 9: 11; 10: 2; 11: 7; 12: 8;

: 4; 41: 14; 42: 12; 43: 5; 44: 4; 45: 7; 46: 7;

: 4; 57: 3; 58: 2; 59: 2; 60: 21; 61: 10; 62: 4;

71: 4: e 75: 17; 76: 15; 77: 7; 78: 5; 79: 9; 80:

10; 81: 2; 82: 3; 83: 10; 84: 2; 85: 5; 86: 14; 87: 1; 88: 1; 1: 2; 93: 4; 94: 5; 95: 8; 96: 2; 97: 9;

98: 1; 99: 6; 100: 5; 101: 7; 103: 1; 104: 2; 105: 1; 106: : 1; 109: 3; 110: 4; 111: 11; 112: 15;

113: 3; 114: 3; 115: 5; 116: 4; 117: 12; 118: 7; 119: 3; 121: ; 124: 1; 125: 13; 127: 1; 128: 4; 129: 1;

130: 2; 131: 4; 132: 2; 133: 4; 135: 4; 136: 21; 137: 8; 138: 2; 139: 2: 140: 1; 141: 2; 142: 4; 143: 7; 145: 6;

146: 5; 147: 2; 148: 3; 149: 4; 150: 2; 151: 8; 152: 5; 153: 13; 154: 1; 155: 8; 156: 8; 157: 4; 158: 5; 160: 1;

161: 2; 162: 4; 164: 2; 165: 2; 167: 2; 168: 3; 170: 1; 171: 8; 172: 2.

Localidades con conchas vacías: 15.

Distribución geográfica: Esta especie

es endémica del oeste de la Península

Ibérica (NOBRE, 1941; GITTENBERGER, 1980).

Está citada en el norte, centro y sur

de Portugal por diversos autores (MORE-

LET, 1877; SERVAIN, 1880; LOCARD, 1899;

NOBRE, 1941; RAMOS y APARICIO, 1985a)

y en toda la franja oeste española desde

Asturias a Huelva (MANGA, 1983; HER-

MIDA ET AL., 1992; PUENTE y PRIETO,

1992). En Galicia es uno de los elementos

más característicos de su malacofauna.

Ponentina subvirescens (Bellamy, 1839) (Fig. 2F)

Citas previas: MACHO VELADO (1871) como H. occidentalis; HIDALGO (1875) como H. occidentalis;

SACCHI y VIOLANI (1977) como Trichia occidentalis; CASTILLEJO (1986) como Ponentina ponentina.

Material examinado (321 ejemplares). 1: 2; 2: 2; 4: 1; 19: 3; 21: 5; 26: 6; 30: 1; 32: 1; 36: 3; 38: 1; 41: 4;

42: 14; 43: 10; 44: 3; 45: 2; 47: 7; 48: 3; 49: 4; 50: 10; 51: 1; 52: 8; 54: 2; 57: 5; 59: 1; 60: 6; 61: 1; 70: 1; 76:

2; 77: 2; 79: 1; 86: 1; 89: 1; 91: 2; 97: 2; 99:

115: 2; 117: 1; 119: 119; 122: 7; 132: 3; 13

Localidades con conchas vacías: 3; 6;

131; 142; 148; 151; 159; 160.

Distribución geográfica: Especie

atlántica que se distribuye por el suroeste

de Europa (KERNEY ET AL., 1983), pertene-

ciente a la fauna atlantico-lusitánica.

En la Península se distribuye a lo

largo de todo su tercio oeste (LOCARD,

5; 103: 3; 107: 2; 109: 1; 110: 1; 111: 3; 112: 1; 113: 1; 114: 9;

:2; 134: 22; 135: 1; 137: 2; 138: 1; 146: 7; 153: 8; 169: 2.

22; 23; 24; 25; 28; 29; 34; 39; 53; 73; 83; 88; 120; 121; 127;

1899; NOBRE, 1941; MANGA, 1983; Cas-

TILLEJO, 1986; HERMIDA ET AL., 1992;

PUENTE y PRIETO, 1992). Los puntos más

orientales corresponden a La Rioja,

Burgos y Alava (PUENTE y PRIETO,

1992).

Mengoana brigantina (da Silva Mengo, 1867) (Fig. 2G)

Material examinado (3 ejemplares). 9: 1; 67: 1; 128: 1.

Distribución geográfica: Esta espe-

cie es un endemismo ibérico restringido

al noroeste de la Península.

Su área de distribución se extiende

desde el extremo nororiental de Portu-

gal, hasta el oeste de Vizcaya (HIDALGO,

1875; ORTIZ DE ZARATE, 1949; RAMOS y

APARICIO, 1985a; MANGA, 1983; HER-

MIDA ET AL., 1992; PUENTE y PRIETO,

1992). Hasta este momento su límite oc-

cidental se encontraba en el este de Gali-

cia (CASTILLEJO, 1986).

Iberus, 15 (1), 1997

Observaciones: Se trata de la cita

más occidental de las conocidas y de la

primera para el área de estudio, encon-

trándose en zonas costeras. Su carácter

calcícola puede ser la causa de que

hayamos recogido un escaso número de

ejemplares en nuestra área de estudio,

predominantemente granítica.

Este taxon presenta una cierta contro-

versia dado que el holotipo de esta espe-

cie, citada como Helix brigantina (da Silva

Mengo, 1867) y recogido en Braganca

(Portugal), se ha extraviado, conserván-

dose únicamente su descripción, sin nin-

gún dibujo o representanción y sin que

haya vuelto a encontrarse ningún ejem-

plar en la localidad tipo. ROSSMASSLER

(1879) figura un ejemplar de esta especie,

aportando otros datos a su descripción y

señalando una nueva localidad en el

norte de la Península (La Liébana, Can-

tabria). NOBRE (1941) no encuentra esta

especie en ningún punto de Portugal y

presupone que la especie descrita por DA

SILVA MENGO (1867) podría tratarse de

una variedad minor de Helix inchoata

(Morelet, 1879). Este problema persiste

hoy en día y sigue siendo necesario com-

probar si el genital de los individuos en-

contrados en los alrededores de Bra-

ganca coincide con el representado por

ORTIZ DE ZARATE (1949). De no ser así, lo

que ahora se conoce como Mengoana bri-

gantina en el norte de la Península debe-

ría ser considerada como Mengoana jes-

chaul (ORTIZ DE ZARATE, 1949).

Oestophora (Oestophora) barbula (Rossmássler, 1838) (Fig. 2h)

Citas previas: GRAELLS (1846) como H. holosericea; MACHO VELADO (1871) como H. barbula;

HIDALGO (1875, 1890) como H. barbula; ORTIZ DE ZARATE (1962); SACCHI y VIOLANI (1977); CASTI-

LLEJO (1984); OTERO y TRIGO (1989).

Material examinado (661 ejemplares). 1: 10; 2: 4; 3: 7; 4: 8; 5: 2;

2; 19: 2; 20: 4; 21: 2; 22: 2; 23: 4; 24: 5; 25: 10; 26: 14; 27: 6

15; 41: 17; 42: 26; 43: 7; 45: 2; 46: 3; 47: 1; 48: 11; 49:

60: 7; 61: 1; 62: 1; 67: 2; 68: 4; 69: 33; 70: 1; 74: 2; 77:

6; 88: 2; 89: 4; 94: 2; 95: 2; 96: 1; 97: 8; 98: 2; 99: 3;

116: 2; 117: 8; 118: 3; 119: 5; 120: 6; 122: 1; 123: 4; 127: 8; 1

136: 7; 137: 4; 138: 8; 139: 1; 143: 1; 145: 2; 149: 10; 150: 1; 151:

157: 1; 158: 8; 161: 6; 168: 7; 171: 14.

Localidades con conchas vacías: 15.

Distribución geográfica: Esta espe-

cie es un endemismo ibérico pertene-

ciente a la fauna atlántico lusitánica.

Está presente en todo el oeste penin-

sular, desde Galicia y Asturias hasta el

sur de Portugal y Huelva, adentrándose

en algunos puntos del centro peninsular

; 6: 3;7: 1; 9: 2; 10: 1; 11: 1; 14: 7; 17:

; 28: 9; 29: 20; 30: 7; 32: 2; 35: 10; 37: 15; 39:

; 51: 6;52: 5;53: 1; 54: 1; 56: 2; 57: 6; 58: 8; 59: 3;

;79: de 81: 4; 82: 2; 83: 10; 84: 3; 85: 1; 86: 14; 87:

: 4; 109: 6; 110: 7; 112: 1; 113: 23; 114: 17; 115: 4;

sl 5; 129: 5; 130: 1; 132: 2; 134: 1; 135: 34;

3; 152: 8; 153: 15; 154: 2; 155: 7; 156: 8;

(NOBRE, 1941; ORTIZ DE ZÁRATE, 1962;

ALTIMIRA, 1969; MANGA, 1983; RAMOS y

APARICIO, 1985a; CASTILLEJO, 1984;

HERMIDA ET AL., 1992). Alejada de este

área se ha citado también en sureste

ibérico (HIDALGO, 1875; GASULL, 1975) y

Pirineos.

Oestophora (Oestophora) silvae Ortiz de Zárate López, 1962 (Fig. 21)

Citas previas: HIDALGO (1875, 1890) como O. lusitanica; ORTIZ DE ZARATE (1962); CASTILLEJO

(1984) como O. lusitanica var. minor; OTERO y TRIGO (1989).

Material examinado (474 ejemplares). 1: 16; 3: 1; 4: 2; 5: 5; 7: 3; 8: 2; 13: 1; 1

o ec ads 35: 4

8; 43: 10; 44: 7; 45: 2; 46: 9; 47: 9; 50: 13; 51: 3; 53: 15; 54: 2;

77: 7; 82: 1; 84: 3; 88: 1; 90: 2; 91: 2; 99: 18; 100: 10; 106: 3;

121: 16; 130: 1; 137: 4; 138: 1; 141: 6; 154: 1; 155: 1; 158: 1;

Localidades con conchas vacías: 16; 87; 93.

4: 4; 18: 2; 19: 1; 21: 8;

3: 13; ; 37:29; 39: 7; 40: 34; 41: 39; 42:

55: 2; 56: 1; 58: 5; 62: 1; 63: 3; 71: 2; 75: 1;

108: 7; 109: 7; 110: 10; 111: 1; 115: 9; 120: 1;

159: 6; 160: 3; 166: 2; 170: 4.

ONDINA £7 AZz.: Distribución de la superfamilia Helicoidea en el oeste de Galicia

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Figura 2. Mapas de distribución. A: Cochicella barbara; B: Cochlicella conoidea; C: Ashfordia granu-

lata; D: Zenobiella subrufescens, E: Portugala inchoata, E: Ponentina subvirescens, G: Mengoana bri-

gantina; H: Oestophora barbula; 1: Oestophora silvae, J: Theba pisana; K: Cepaea nemoralis, L: Helix

aspersa. Localidades citadas en este trabajo (*); procedentes de la bibliografía (*); únicamente

encontradas conchas vacías (O).

Figure 2: Distribution maps. A: Cochlicella barbara; B: Cochlicella conoidea; C: Ashfordia granu-

lata; D: Zenobiella subrufescens; E: Portugala inchoata; F: Ponentina subvirescens; G: Mengoana

brigantina; A: Oestophora barbula; /: Oestophora silvae; /: Theba pisana; K: Cepaea nemoralis; L:

Helix aspersa. Localities reported in this paper (e); bibliographic records (*); only shells found (O).

Iberus, 15 (1), 1997

Distribución geográfica: Es un en-

demismo ibérico restringido al norte de

la Península, desde Galicia hasta el

norte de Alava (CASTILLEJO, 1986; OJEA y

ANADÓN, 1983; PUENTE y PRIETO, 1992;

HERMIDA, ET AL., 1992).

Observaciones: ORTIZ DE ZARATE

(1962) al describir O. silvae expuso las dife-

rencias que existían entre esta especie y

Oestophora lusitanica (Pfeiffer, 1841). Señala

que O. silvae es más pequeña, tiene el borde

superior de la abertura más corto, un

engrosamiento mayor del peristoma en

toda su extensión, presenta una callosi-

dad blanca interna en el extremo del borde

superior de la última vuelta con la ante-

penúltima y carece del estriado espiral en

la cara inferior, cerca de la abertura, que

caracteriza a O. lusitanica. Respecto al

aparato genital destaca la distinta inserción

del músculo retractor del pene, que se

sitúa hacia la mitad de la vaina en O. lusi-

tanica y cerca del extremo en O. silvae, y la

distinta longitud de las glándulas multí-

fidas y del oviducto libre, mucho más

cortos en O. silvae.

Según ORTIZ DE ZÁRATE (1962) O.

silvae es la misma especie que fue descrita

por da Silva en 1871 como O. lusitanica

var. minor. Así mismo asigna a esta especie

todas las citas de O. lusitanica, dadas por

HIDALGO (1875), para el norte de la Penín-

sula y Galicia. Sin embargo CASTILLEJO

(1984) encontró, en diversos puntos de

Galicia, ejemplares que no coincidían com-

pletamente con la descripción de O. silvae,

siendo más similar a la de O. lusitanica var.

minor, a la que asignó esos ejemplares.

Nosotros hemos podido estudiar indi-

viduos de O. lusitanica de Portugal, y nin-

guno de nuestros ejemplares gallegos se

ajusta a las características de esta especie.

Respecto a O. silvae, hemos observado

que los datos de la descripción de ORTIZ

DE ZARATE (1962) deberían ser revisados,

ya que existen ejemplares de mayor diá-

metro que el señalado por él, y una obser-

vación detallada de la concha húmeda

pone de manifiesto, en todos los ejempla-

res, las estrías espirales en la cara inferior

que, en un principio, sólo parecía presen-

tar O. lusitanica. En lo referente al aparato

genital, las glándulas multífidas pueden

ser más largas que las descritas y el mús-

culo retractor del pene presenta cierta

variabilidad en su inserción. Observadas

estas variaciones y una vez estudiados los

ejemplares de nuestra colección y la del

Dr. Castillejo, hemos seguido a Ortiz de

Zárate considerando las citas gallegas de

O. lusitanica var. minor como O. silvae.

Familia HELICIDAE Rafinesque, 1815

Theba pisana (Muller, 1774) (Fig. 2)

Citas previas: MACHO VELADO (1871) como H. pisana; HIDALGO (1875, 1890) como H. pisana;

SACCHI y VIOLANI (1977) como Euparipha pisana; CASTILLEJO (1986); OTERO y TRIGO (1989).

Material examinado (373 ejemplares). 4: 20; 6: 3; 11: 3; 26: 10; 30: 2; 48: 1; 49: 1; 57: 14; 60: 2; 67: 5;

69: 50; 82: 1; 83: 4; 85: 12; 86: 5; 87: 10; 91: 10; 93: 2; 94: 17; 95: 1; 97: 3; 100: 3; 113: 7; 117: 1; 118: 12;

126: 9; 128: 1; 131: 33; 133: 12; 136: 15; 149: 35; 151: 25; 153: 10; 156: 3; 165: 2; 168: 5; 173: 24.

Distribución geográfica: Esta especie

presenta una distribución circunmedite-

rránea que remonta las costas atlánticas

hasta Inglaterra (KERNEY ET AL., 1983).

En la Península se extiende a lo largo

de toda la costa, aunque es capaz de colo-

nizar áreas del interior cuando se dan con-

diciones adecuadas, tales como disponi-

bilidad de sales solubles (Prieto, com. pers.).

Como citas más recientes en el interior se

pueden señalar las dadas por HERMIDA

ET AL. (1992) y PAREJO ET AL. (1993b).

Cepaea (Cepaea) nemoralis (Linneo, 1758) (Fig. 2K)

Citas previas: MACHO VELADO (1871) como H. nemoralis; HIDALGO (1875, 1890) como H.

nemoralis; SACCHI y VIOLANI (1977); SACCHI (1981); CASTILLEJO (1986); OTERO y TRIGO (1989). *

ONDINA £7 4Z.: Distribución de la superfamilia Helicoidea en el oeste de Galicia

Material examinado (1229 ejemplares). 1: 32; 2: 13; 3: 10; 4: 7; 5: 3; 6: 11;7: 2; 8: 4; 9: 4,10: 1; 11: 4; 12:

4; 13: 4; 14: 7; 16: 8; 17: 3; 18: 4; 19: 4; 20: 2; 21: 5; 22: 9; 23: 3; 24: 9; 25: 17; 26: 8; 27: 9; 28: 10; 29: 4; 30:

15; 31: 4; 32: 10; 33: 4; 35: 1; 36: 12; 37: 4; 38: 12; 39: 12; 40: 5; 41: 28; 42: 2; 43: 8; 44: 5; 45: 12; 46: 12; 47:

15; 48: 13; 49: 7; 50: 7; 51: 9; 52: 3; 53: 1; 54: 9; 55: 5; 56: 4; 57: 7; 58: 2; 59: 6; 61: 9; 62: 2; 63: 9; 64: 11; 65:

1; 66: 1; 67: 12; 68: 2; 69: 1; 70: 5; 71: 7; 72: 10; 73: 2; 74: 4; 75: 10; 76: 6; 77: 20; 79: 6; 80: 7; 81: 8; 82: 4; 83:

12; 84: 11; 85: 3; 86: 10; 87: 11; 88: 6; 89: 7; 90: 8; 91: 2; 92: 6; 93: 17; 94: 3; 95: 11; 96: 12; 97: 10; 98: 4; 99:

14; 100: 14; 101: 8; 102: 20; 103: 2; 104: 9; 105: 2; 106: 4; 107: 2; 108: 10; 109: 15; 110: 23; 111: 8; 112: 14;

113: 16; 114: 7; 115: 12; 116: 13; 117: 26; 118: 8; 119: 8; 120: 12; 121: 11; 122: 7; 123: 11; 125: 7; 126: 1;

127: 10; 128: 5; 129: 8; 130: 3; 131: 11; 132: 12; 133: 14; 134: 3; 135: 6; 136: 4; 137: 5; 138: 7; 139: 6; 140: 1;

141: 10; 142: 8; 143: 8; 144: 4; 145: 5; 146: 20; 147: 12; 148: 7; 149: 8; 150: 5; 151: 4; 152: 7; 153: 12; 154: 2;

155: 3; 156: 7; 157: 3; 158: 1; 159: 2; 160: 1; 161: 3; 162: 2; 163: 2; 164: 1; 165: 2; 166: 1; 170: 2; 171: 3.

Distribución geográfica: Presenta

una distribución centro-occidental euro-

pea (BOATO ET AL., 1982).

En la Península Ibérica se extiende

por la franja portuguesa (HIDALGO,

1875; LOCARD, 1899; NOBRE, 1941) y por

toda la mitad norte (BOFILL y HAAs,

1919, 1920a, 1920b; ORTIZ DE ZARATE y

ORTIZ DE ZARATE, 1949; MANGA, 1983;

RAMOS, 1985; APARICIO, 1986; HERMIDA

ET AL., 1992; LARRAZ y EQUISOAIN, 1993;

PAREJO ET AL., 1993b; ALTONAGA ET AL.,

1994). En la mitad sur peninsular es más

frecuente en Portugal, pero en los úl-

timos años varios autores han ampliado

su distribución en esta zona (MARTÍNEZ-

ORTI y ROBLES, 1993; PAREJO, ET AL.,

1993a).

Helix (Cornu) aspersa (Múller, 1774) (Fig. 2L)

Citas previas: MACHO VELADO (1871); HIDALGO (1875, 1890); SAccHI y VIOLANI (1977) como

Cryptomphalus aspersus; CASTILLEJO (1986); OTERO y TRIGO (1989).

Material examinado (1579 ejemplares). 1: 1; 2: 1; 3: 5; 4: 32; 5: 3; 6: 12; 8: 4; 9: 6; 10: 1; 11: 3; 12: 5; 13: 12;

14: 3; 16: 2; 17: 3; 19: 5; 20: 7; 21: 30; 22: 4; 23: 4; 24: 3; 25: 6; 26: 16; 27: 6; 28: 54; 29: 15; 30: 9; 31: 3; 32: 12;

35: 18; 36: 8; 37: 20; 38: 17; 39: 15; 40: 13; 41: 28; 42: 22; 43: 2; 44: 12; 45: 17; 46: 11; 47: 4; 48: 26; 49: 18; 50:

11; 51: 20; 52: 22; 53: 20; 54: 11; 55: 10; 56: 7; 57: 19; 58: 1; 59: 12; 60: 11; 61: 21; 62: 9; 63: 1; 64: 27; 65: 2; 67:

23; 68: 16; 69: 12; 70: 14; 71: 11; 73: 6; 75: 13; 76: 4; 77: 7; 78: 1; 79: 6; 80: 2; 81: 10; 82: 10; 83: 5; 84: 5; 85: 8;

86: 10; 87: 10; 88: 10; 90: 2; 91: 17; 92: 11; 93: 16; 94: 16; 95: 14; 96: 11; 97: 9; 98: 9; 100: 14; 101: 42; 102: 48;

103: 21; 104: 5; 106: 10; 109: 10; 110: 7; 111: 4; 112: 6; 113: 17; 114: 22; 115: 14; 116: 2; 117: 36; 118: 8; 119: 15;

120: 16; 121: 11; 122: 11; 123: 36; 124: 3; 125: 9; 126: 8; 127: 13; 128: 9; 129: 12; 130: 2; 131: 11; 132: 16; 133: 8;

134: 22; 135: 5; 136: 10; 137: 12; 138: 2; 139: 1; 141: 3; 143: 4; 144: 1; 145: 8; 146: 16; 147: 4; 148: 6; 149: 6; 150:

1; 151: 3; 152: 2; 153: 9; 154: 2; 155: 5; 156: 2; 157: 1; 158: 2; 159: 1; 160: 4; 162: 3; 165: 3; 167: 1; 168: 1; 171: 3.

Distribución geográfica: Especie

distribuida por el Mediterráneo y oeste

de Europa. (BOATO ET AL., 1982).

En la Península Ibérica se ha citado

repetidamente en todas las regiones

DISCUSIÓN

En un principio y dadas las condicio-

nes climáticas gallegas favorables para la

vida de los gasterópodos terrestres, podrí-

amos pensar que es un área de abundante

fauna malacológica. Pero comparándola

con otras zonas de la Península Ibérica

enclavadas en el área atlántica podemos

comprobar que la diversidad y abundan-

(ALTONAGA ET AL., 1994) y en las Islas

Baleares, y aún faltando zonas por estu-

diar, dado su carácter ubiquista y sinan-

trópico, puede asegurarse que está pre-

sente en todo el territorio peninsular.

cia de helícidos es especialmente escasa

en Galicia. Probablemente esto es debido,

principalmente, a la escasez de sustrato

calizo, necesario para la formación de la

concha de todos los gasterópodos en

general y, de la conchas gruesas y duras

de los helícidos en particular (BOYcorr,

1934; KERNEY y CAMERON, 1983).

Iberus, 15 (1), 1997

Con este trabajo se ha ampliado el

área de distribución conocida de la ma-

yor parte de las especies, pudiendo seña-

lar la notable presencia en la zona de es-

tudio de especies como H. aspersa, C. ne-

moralis y P. inchoata, que aparecen en más

del 80% de las cuadrículas visitadas. A

diferencia de ésto, existen otras especies

que han presentado distribuciones res-

tringidas, especialmente aquellas origi-

nariamente mediterráneas, intimamente

ligadas al litoral como T. pisana, C. acuta,

C. barbara, y C. conoidea, destacando es-

pecialmente esta última, que al igual que

sucede en el resto de la Península, no as-

ciende por la costa cantábrica. En contra-

posición a ésto, H. itala, también con un

comportamiento litoral en el área de es-

tudio, no desciende del norte de Galicia,

siendo su cita más al sur, en todo este te-

rritorio, la Sierra de O Courel, zona ca-

liza con una altitud superior a los 900 m.

También hemos de señalar que la

mayor parte de las especies han sido

capturadas en gran variedad de hábitats

(bajo troncos, muros, linderas de culti-

vos...), pudiendo destacar únicamente

la preferencia de E. quimperiana por los

diferentes tipos de arbolados, en mayor

medida pinares y robledales, y de P.

inchoata y P. subvirescens en los prados y

zonas carentes de vegetación arborea.

BIBLIOGRAFÍA

ALTIMIRA, C., 1969. Notas Malacológicas. VIH

Moluscos del Delta del Llobregat. IX Nuevas

aportaciones y datos a la fauna malacoló-

gica catalana. XI Moluscos terrestres y de

agua dulce recogidos en la provincia de Lugo

y en Asturias. Publicaciones del Instituto de

Biología Aplicada, 46: 91-113.

ALTONAGA, K.; GÓMEZ, B.; MARTÍN, R.; PRIETO,

C.; PUENTE, A. y RALLO, A., 1994. Estudio fau-

nístico y biogeográfico de los moluscos terrestres

del norte de la Península Ibérica. 505 pp. Ed.

Eusko Legebiltzarra, Parlamento Vasco.

ANADÓN, N. y OJEA, M., 1984. Gasterópodos te-

rrestres del Monte Naranco (Oviedo, Astu-

rias). Distribución, diversidad y afinidades

faunísticas. Revista de Biología de la Universi-

dad de Oviedo, 2: 121-129.

APARICIO, M. T., 1986. The geographic distri-

bution of the Family Helicidae in Central

Spain. Proceedings of the 8* International Ma-

lacological Congress, Budapest.

Por último queremos mencionar que

hemos encontrado la mayoría de las

especies citadas anteriormente a noso-

tros en el área de estudio, y las que no lo

han sido podemos atribuirlo, no tanto a

errores de muestreo, como a la posibili-

dad de que se encuentren poco repre-

sentadas. Este es el caso de Helicigona

lapicida (Linneo, 1758) citada por ROLÁN

y OTERO (1988) en una única localidad

de la provincia de A Coruña. Así mismo,

hay que señalar Otala punctata (Múller,

1771) citada como Otala lactea (Múller,

1771) por CASTILLEJO (1986) en la esta-

ción biológica de Lourizán (Ponteve-

dra). Esta especie de ámbito mediterrá-

neo fue, con seguridad, introducida con

un cultivo experimental en la estación

agrícola, y la población, muy abundante

en el momento de su cita (el autor

recoge unos 300 ejemplares) probable-

mente no prosperó, puesto que hemos

realizado repetidas visitas a esa zona,

sin encontrar ningún ejemplar.

AGRADECIMIENTOS

Este trabajo forma parte del Proyecto

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O Sociedad Española de Malacología

Iberus, 15 (1): 23-29, 1997

Una nueva especie de Lirularía (Gastropoda: Trochidae) de

las islas de Sáo Tomé y Príncipe, África Occidental

A new species of Lirularia (Gastropoda: Trochidae) from Sao Tomé

y Príncipe Islands, West Africa

Federico RUBIO* y Emilio ROLÁN**

Recibido el 18-111-1996. Aceptado el 8-X-1996

RESUMEN

El estudio del material obtenido durante dos expediciones realizadas a las Islas de Sáo

Tomé y Príncipe en los años 1989 y 1990, ha proporcionado una nueva especie de tró-

quido con apariencia de Solariella, que por sus características morfológicas y radulares

pertenece a la subfamilia Lirulariinae. Se describe esta especie, nueva para la ciencia,

comentándose las características morfológicas de la concha, partes blandas y rádula, e

incluyéndola en el género Lirularia. Se discute la inclusión en dicho género de otras dos

especies de la costa occidental africana.

ABSTRACT

The study of the material obtained in Sáo Tomé and Principe Islands during two expedi-

tions in the years 1989 and 1990, have yielded a new trochid species with Solariella,

which for its morphological and radular characters belongs to the subfamily Lirulariinae.

This species is described as new for science within the genus Lirularia and its morphologi-

cal characteristics of shell, soft parts and radula are commented on. The assignment to this

genus of other two species of the West African coast is discussed.

PALABRAS CLAVE: Gastropoda, Trochidae, Lirularia, nueva especie, Islas de Siío Tomé y Príncipe.

KEY WORDS: Gastropoda, Trochidae, Lirularia, new species, Sio Tomé and Príncipe Islands.

INTRODUCCIÓN

Se han citado varias especies de Sola-

riella s. 1. de la costa africana en diversos

trabajos. SMITH (1871) describe la primera,

Solariella canaliculata Smith, 1871, de

Whydah (Dahomey), actualmente Repú-

blica de Benin. Esta misma especie y Sola-

riella dereimsí Dollfus, 1911 son citadas para

Angola por GOFAs, PINTO AFONSO Y

BRANDAO (1985); NICKLÉS (1950) menciona

Solariella monodi Fischer y Nicklés, 1946

para Guinea Francesa (Guinea Conakry)

y S. dereimsi para Mauritania y Senegal.

Alguna otra especie, como S. valida Daut-

zenberg y Fischer, 1906, descrita para el

archipiélago de Cabo Verde, procede de

aguas profundas.

En BERNARD (1984) aparece repre-

sentada una concha con la denomina-

*Dpto. de Zoología. Facultad de Ciencias Biológicas, Universidad de Valencia. Dr. Moliner, 50, 46100 Burjasot

(Valencia).

**Cánovas del Castillo, 22-59 F, 36202 Vigo.

Iberus, 15 (1), 1997

ción Solariella sp., pero en realidad se

trata de Cyclostremiscus calameli (Jousse-

aume, 1872) que es en realidad un Torni-

dae (Gofas, com. pers.).

No hay citas de especies de Solariella

para el archipiélago de Sáo Tomé y Prín-

cipe, aunque en el último listado de es-

pecies de FERNANDES Y ROLÁN (1993), se

menciona una única especie de este gé-

nero, como Solariella sp.

HERBERT (1987) demostró que muchas

de las especies de África del Sur, consi-

deradas tradicionalmente del género Sola-

riella no lo son, sino que pertenecen a la

subfamilia Umboniinae en lugar de Sola-

riellinae. HICKMAN Y MCLEAN (1990) pu-

blicaron una revisión de la superfamilia

Trochoidea y, atendiendo a la morfología

de sus partes blandas y rádula, agrupa-

ron las “Solariellas” en un grupo informal

formado por las subfamilias Trochinae Ra-

finesque, 1815, Stomatellidae Gray, 1840,

Calliostomatinae Thiele, 1924 y Solarielli-

nae Powell, 1951; agrupando las especies

de Umboniinae en otro grupo informal

formado por Lirulariinae Hickman y

McLean, 1990, Halistylinae Keen, 1958 y

Umboniinae Adams y Adams, 1854. Wa-

RÉN (1993), siguiendo dicha ordenación,

considera que “Solariella” canaliculata y

“Solariella” dereimsi pertenecen a la subfa-

milia Umboniinae.

Durante las campañas de recolección

de moluscos efectuadas por el segundo

RESULTADOS

autor en los años 1989 y 1990 en el archi-

piélago de Sáo Tomé y Príncipe, Golfo

de Guinea, se recolectaron ejemplares y

conchas de una “Solariella” aparente-

mente diferente de las especies previa-

mente conocidas. Esta misma especie,

referida como Solariella sp. por FERNAN-

DES Y ROLÁN (1993) y WARÉN (1993), es

ahora descrita, aunque en un género

diferente.

Al mismo tiempo, las especies “Sola-

riella” canaliculata y “Solariella” dereimsi,

recolectadas y observadas en las expedi-

ciones a Angola y Mauritania de 1989 y

1996 respectivamente, son transferidas a

la subfamilia Lirulariinae, género Lirula-

ria, basándonos en la similitud morfoló-

gica de sus partes blandas y rádula.

MATERIAL Y MÉTODOS

Se han estudiado 8 ejemplares y 82

conchas, procedentes de sedimentos

obtenidos en distintas localidades del

archipiélago, mediante buceo a pulmón

libre, a profundidades comprendidas

entre 5 y 15 metros. Tras la observación

de su comportamiento, algunos indivi-

duos se relajaron y posteriormente se

fijaron en solución tamponada de for-

maldehído al 5%. Para la observación

conquiológica y radular se ha utilizado

la microscopía electrónica de barrido.

Superfamilia TROCHACEA Rafinesque, 1815

Familia TROCHIDAE Rafinesque, 1815

Subfamilia LIRULARINAE Hickman y McLean, 1990

Género Lirularia Dall, 1909

Lirularia antoniae spec. nov. (Figs. 1A-B, 2 - 6)

Material estudiado: Isla de Sáo Tomé: Praia das conchas: 8 ejemplares y 16 conchas a -15 m;

Ciudad Sáo Tomé: 36 conchas. Isla de Príncipe: Bahía de Santo Antonio: 24 conchas a -10 my;

Bahía das Agulhas: 6 conchas.

Material tipo: Holotipo (Fig. 1B) y dos paratipos procedentes de la localidad tipo, depositados en

el Museo Nacional de Ciencias Naturales de Madrid, con el n* 15.05/23749, dos paratipos proce-

dentes de Praia das Conchas (Sáo Tomé) en el Muséum National d Histoire Naturelle de París, dos

paratipos procedentes de Ciudad de Sáo Tomé en el American Museum of Natural History de Nueva

York y en The Natural History Museum de Londres y 25 en cada una de las colecciones de los autores.

Localidad tipo: Praia das Conchas, Sáo Tomé.

Etimología: La especie está dedicada a Antonia Hueso, esposa del primer autor.

RUBIO Y ROLÁN: Nueva especie de Lirularia de Sáo “Tomé y Príncipe

Figura 1. Lirularia antoniae spec. nov. Holotipo. Praia das Conchas, Sáo Tomé. A: animal en movi-

miento. B: concha (MNCN). Escalas 1 mm.

Abreviaturas. o: ojo; op: opérculo; p: pie pd: proceso digitiforme sobre el extremo del morro; s:

sifón; tc: tentáculo cefálico; te: tentáculo epipodial.

Figure 1. Lirularia antoniae spec. nov. Holotype. Praia das Conchas, Sáo Tomé. A: crawling animal.

B: shell (MNCN). Scale bars 1 mm.

Abbreviations. o: eye; op: operculum; p: foot; pd: digitiform processes on the tip of'snout; s: siphon; tc: cep-

halic tentacle; te: epipodial tentacle.

Descripción: Concha sólida, bri-

llante, nacarada, de perfil cónico y

espira algo elevada, compuesta por

unas 5 vueltas convexas, que están sepa-

radas por una sutura ancha y acanalada.

Protoconcha (Fig. 5) con apenas una

vuelta de espira, lisa, con el núcleo de-

formado y de unas 200 ym. Ornamenta-

ción formada por cordones espirales y

costillas transversales muy numerosas,

que al entrecruzarse forman pequeños

nódulos; se observan, además, sutiles

líneas de crecimiento que se extienden

paralelas a las costillas. Última vuelta

con 10 cordones espirales, de los que el

primero, subsutural, y el décimo,

periumbilical, son los más prominentes

por ser nodulosos y angulan la concha.

Los restantes cordones son poco marca-

dos, sobre todo en la parte media de la

periferia, donde apenas son percepti-

bles. Las costillas transversales están

menos marcadas en la primera y última

vuelta de la teloconcha, aunque en ésta

última son numerosísimas; su curso es

prosoclino y atraviesan la totalidad de la

vuelta. Ombligo ancho y profundo, bor-

deado por el décimo cordón espiral; en

su interior se observan otros cuatro cor-

dones espirales más. Abertura subcircu-

lar, prosoclina; labio externo fino, angu-

lado por la presencia de los cordones

espirales; labio interno, ligeramente

arqueado, reflejado hacia el exterior,

pero sin llegar a ocluir el ombligo.

Coloración muy variable, de blanco-

amarillento a rosa pálido, con manchas

pardo rojizas o pardo oscuras y cierta

iridiscencia.

Respecto a sus dimensiones, el holo-

tipo (Fig. 1B) mide 1,78 mm de altura y

1,98 mm de anchura.

El animal (Fig. 1A) es de color blan-

quecino excepto el sifón y una franja de

color negro situada en la parte distal del

morro. La cabeza tiene un par de tentá-

culos cefálicos muy largos y con micro-

papilas, ojos negros situados sobre

cortos pedúnculos y carece de membra-

nas cefálicas. El morro está muy depri-

23)

Tberus, 15 (1), 1997

mido distalmente y sus extremos se pro-

longan transversalmente; hay un

proceso digitiforme con forma de peine

sobre su extremo. A cada lado de la

cabeza se observa un lóbulo cervical

modificado; el izquierdo, subdividido

en apéndices tentaculiformes y, el

derecho, que es plano, en el animal vivo,

se enrolla para formar una estructura

tubular con aspecto de sifón, moteado

con manchas negro-opacas y de un

tamaño similar a los tentáculos cefálicos.

Epipodio con cuatro pares de tentácu-

los, carentes de macropapilas sensoria-

les en su base, el primer par en posición

anterior y los tres pares restantes alrede-

dor del lóbulo opercular. El pie es muy

móvil, su extremo anterior es bilobulado

y se prolonga lateralmente y luego se

afila progresivamente hacia su extremo

posterior, para acabar en punta.

Rádula (Fig. 6) formula N. 4. 1. 4. N.

El diente central y los laterales son muy

similares y están reducidos a láminas

basales con una pequeña cúspide cada

uno. El diente central está cubierto en

parte por las láminas laterales más inter-

nas. Dientes marginales con cúspides

relativamente largas y anchas, con den-

tículos romos y aserrados.

Distribución: Sólo conocida del

archipiélago de Sáo Tomé y Príncipe

(Golfo de Guinea). No se han encon-

trado ejemplares de esta especie en

zonas continentales próximas al archi-

piélago, por lo que, probablemente, se

trata de un endemismo insular.

Hábitat: Especie infralitoral que vive

sobre fondos de arena en zonas de

aguas claras, entre -5 y -15 metros.

Discusión: Tanto la estructura plana,

enrollada en forma de sifón, como los

procesos digitiformes visibles a cada

lado de la cabeza en Lirularia antoniae, al

igual que en otras especies de África oc-

cidental (“Solariella” canaliculata y “Sola-

riella” dereimsi1) (Gofás com. pers.), son

probablemente lóbulos cervicales modi-

ficados, homólogos a los de otros Trocoi-

deos. Su función, para el lado inhalante,

se puede suponer que es la de actuar

como un filtro para evitar la entrada de

las partículas en la cavidad paleal. El ló-

bulo derecho tiene una función exha-

lante. Dichas estructuras surgen como

consecuencia de la adaptación de las es-

pecies de Solariellinae, Umboniinae y

Lirulariinae de África occidental a los

fondos blandos sobre los que habitan, a

diferencia de otros trocoideos (Clancu-

lus, Collonia, Gibbula, Tricolia, etc.) cuyo

hábitat está limitado a fondos rocosos.

Aspectos de esta adaptación coinciden-

tes con estas características anatómicas

pueden verse en FRETTER (1975) y HICk-

MAN (1985).

Siguiendo la clasificación de la fami-

lia Trochidae propuesta por HICKMAN Y

MCLEAN (1990), y atendiendo a los ca-

racteres morfológicos y radulares que

diferencian las tres subfamilias que per-

tenecen al grupo informal Halistylinae +

Umboniinae + Lirulariinae, hemos si-

tuado la nueva especie en la subfamilia

Lirulariinae, género Lirularia. Las espe-

cies “Solariella” canaliculata (Fig. 7) y

“Solariella” dereimsil (Figs. 8-9) son con-

genéricas entre sí, y se diferencian ana-

tómicamente de Lirularia antontae tan

solo por la distribución de las papillas

del morro; sin embargo, comparten ca-

racteres comunes como:

e protoconchas de pequeño tamaño,

no superior a 200 um, con el núcleo

deformado;

e ombligo amplio, no ocluido ni

total ni parcialmente;

e radulas similares, muy parecidas a

su vez a las de Umbonium;

e lóbulo cervical izquierdo subdivi-

dido en un proceso tentaculiforme y

lóbulo cervical derecho plano, enrollado

para formar una especie de sifón.

Todo esto nos hace considerarlas

pertenecientes a la subfamilia Lirularii-

nae, género Lirularia, en lugar de Sola-

riellinae o Umboniinae.

Eirularia antoniae, L. canaliculata y L.

dereímsii, se diferencian de las especies

pertenecientes a Solariellinae por tener

protoconcha pequeña con el núcleo de-

formado, la rádula con su zona central

simplificada, el diente central y los dien-

RUBIO Y ROLÁN: Nueva especie de Lirularia de Sao “Tomé y Príncipe

Figuras 2-6. Lirularia antoniae spec. nov., Praia das Conchas, Sáo Tomé. 2: individuo juvenil, vista

apical (col. E Rubio); 3: individuo juvenil, vista dorsal; 4: paratipo (col. E. Rubio); 5: protoconcha;

6: rádula. Escalas, 2-4: 0,5 mm; 5: 100 um; 6: 12 pm.

Figures 2-6. Lirularia antoniae spec. nov., Praia das Conchas, Sáo Tomé. 2: young specimen, apical view

(E Rubio coll.). 3: young specimen, dorsal view. 4: paratype (E Rubio coll.); 5: protoconch; 6: radula.

Scale bars, 2-4: 0,5 mm; 5: 100 ym; 6: 12 um.

Iberus, 15 (1), 1997

Figuras 7-9. Animal de otras especies de Lirularia de Africa. 7: L. dereimsiz (tomado de GOFAS,

PINTO AFONSO Y BRANDAO, 1985); 8-9: L. canaliculata (dibujo de S. Gofas).

Figures 7-9. Animal of other species of Lirularia from Africa. 7: L. dereimsii (after GOFAS, PINTO

AÁFONSO AND BRANDAO, 1985); 8-9: L. canaliculata (drawing from S. Gofas).

tes laterales reducidos, los tentáculos epi-

podiales carentes de macropapilas senso-

riales en su base y las papilas situadas en

el extremo anterior del morro y no alre-

dedor del disco oral. De las especies per-

tenecientes a Umboniinae, tribu Um-

boniini se diferencian porque estas últi-

mas poseen tentáculos cefálicos dimórfi-

cos y el lóbulo cervical izquierdo en-

vuelve el tentáculo cefálico izquierdo y

RUBIO Y ROLÁN: Nueva especie de Lirularia de Sao Tomé y Príncipe

pedúnculo ocular. De las especies perte-

necientes a la tribu Monileini se diferen-

cian porque presentan un ombligo cerrado,

total o parcialmente, por un callo; por po-

seer pedúnculos oculares prominentes,

AGRADECIMIENTOS

Al Servicio de Microscopía Electró-

nica de la Universidad de Valencia, por la

ayuda prestada en la realización de foto-

grafías al Microscopio Electrónico de Ba-

rrido. A Francisco Fernandes por su cola-

BIBLIOGRAFÍA

BERNARD, P. A., 1984. Coquillages du Gabon. Li-

breville. 140 pp., 75 láms.

FERNANDES, F. Y ROLÁN, E., 1993. Moluscos ma-

rinos de Sao Tomé y Principe: actualización

bibliográfica y nuevas aportaciones. Iberus,

11 (1): 31-47.

FRETTER, V., 1975. Umbonium vestiatium, a filter-

feeding trochid. Journal of Conchology, 177:

541-552.

GOFAS, S., PINTO AFONSO, J. Y BRANDAO, M.,

1985. Conchas e moluscos de Angola. Universi-

dad de Agostinho Neto / Elf Aquitaine. An-

gola. 139 pp.

HERBERT, D. G., 1987. Revision of the Solarie-

llinae (Mollusca: Prosobranchia: Trochidae)

in South Africa. Annals of the Natal Museum,

28: 283-382.

HICKMAN, C. S., 1985. Comparative morphology

and ecology of free living suspension-fee-

ding gastropods from Hong Kong. En Mor-

ton, B. y Dudgeon D. (Eds.): Proceedings of the

second International Workshop on the Malaco-

fauna of Hong kong and Southern China, Hong

kong University Press.: 217-234.

con anchos ojos y bases desarrolladas; y

porque su lóbulo cervical izquierdo (in-

halante) está subdividido terminalmente

en un proceso tentaculiforme dimórfico,

orientado regular y alternativamente.

boración en las expediciones a Sáo Tomé

y Príncipe. A Serge Gofas por la lectura

crítica del manuscrito, por su informa-

ción y aportación de dibujos, y por sus

sugerencias.

HICKMAN, C. S. Y MCLEAN, J. H., 1990. Syste-

matic revision and suprageneric classification of

trochacean gastropods. Natural History Mu-

seum of Los Angeles County, Science Se-

ries, 35. 169 pp.

NICKLÉS, M., 1950. Mollusques testacés marins de

la cóte occidentale d'Afrique. Lechevalier, Pa-

ris, 269 pp., 434 figs.

SmITH, E. A., 1871. A list of species of shells from

West Africa, with descriptions of those hit-

herto undescribed. Proceedings of the Zoological

Society of London, 1871: 727-739.

WARÉN, A. S., 1993. New and little know Mo-

llusca from Iceland and Scandinavia. Part 2.

Sarsia, 78: 159-201.

2D,

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E e bl .

O Sociedad Española de Malacología

Iberus, 15 (1): 31-34, 1997

Una nueva especie de Anticlimax (Gastropoda: Vitrinelli-

dae) de Cuba

A new species of Anticlimax (Gastropoda: Vitrinellidae) from Cuba

Emilio ROLÁN” Raúl FERNÁNDEZ-GARCÉS” y Federico RUBIO”

Recibido el 18-111-1996. Aceptado el 8-X-1996

RESUMEN

Se describe una nueva especie del género Anticlimax (Gastropoda: Vitrinellidae). Se dis-

cute su asignación genérica y se compara con las especies más próximas.

ABSTRACT

A new species of the genus Anticlimax (Gastropoda: Vitrinellidae) is described. lts generic

assignment is discussed and a comparison is made with allied species.

PALABRAS CLAVE: Gastropoda, Vitrinellidae, Anticlimax, especie nueva, Cuba.

KEY WORDS: Gastropoda, Vitrinellidae, Anticlimax, new species, Cuba.

INTRODUCCIÓN

DALL (1903) menciona por primera

vez el subgénero Climacia en la descrip-

ción de Teinostoma (Climacia) calliglyp-

tum, sin hacer descripción alguna de las

características del subgénero ni de las de

la especie típica, siendo ambos definidos

por figuras (PILSBRY Y MCGINTY, 1946a).

PILSBRY Y MCGINTY (1946a) dan a

Climacia categoría genérica y describen y

comentan las características de T.

calliglyptum como las de la especie típica

(por monotipia); al tiempo describen

varias especies nuevas que incluyen en

Climacia.

AGUAYO Y BORRO (1946) sustituyen

por Climacina el previamente ocupado

nombre genérico de Climacia Dall, 1903

non M'Lachlan, 1869 (Neuroptera). Poco

" Cánovas del Castillo 22, 36202 Vigo, España.

“Calle 41, 6607, Cienfuegos 55100, Cuba.

tiempo después, PiLsBRY Y MCGINTY

(1946b) sustituyen Climacina Aguayo y

Borro, 1946 non Gemmellaro, 1878

(Mollusca), por un nuevo nombre: Anti-

climax.

PILSBRY Y OLSSON (1950) hacen una

revisión del género describiendo

algunas especies nuevas del Terciario

americano, y lo subdividen en dos sub-

géneros Anticlimax y Subclimax subgen.

nov.

En el material recolectado en Cuba

durante los muestreos realizados por los

dos primeros autores en los últimos

años, se han encontrado conchas de una

especie que parece pertenecer a este

género, y que es descrita en el presente

trabajo.

Departamento de Zoología, Universidad de Valencia, Dr. Moliner 50, 41600 Burjasot, España.

Iberus, 15 (1), 1997

RESULTADOS

Género Anticlimax Pilsbry y McGinty, 1946

Especie tipo, por monotipia: Teinostoma (Climacia) calliglyptum Dall, 1903

Anticlimax decorata spec. nov.

Material tipo: Holotipo, de Rancho Luna, Cienfuegos, Cuba (Fig. 2), con 1,3 mm de dimensión

máxima, Museo Nacional de Ciencias Naturales de Madrid n* 15.05/27420. De la misma locali-

dad, un paratipo en las colecciones del American Museum of Natural History de Nueva York

(Fig. 1) y The Natural History Museum de Londres y dos en la de R. Fernández Garcés. Zona del

Hotel Comodoro, La Habana, Cuba: un paratipo en la colección del Instituto de Ecología y Siste-

mática de La Habana. Jibacoa, Distrito de La Habana, Cuba: un paratipo en la colección de F.

Rubio y otro en la de E. Rolán.

Localidad tipo: Rancho Luna, Cienfuegos, Cuba.

Etimología: El nombre específico deriva de su elaborada escultura.

Descripción: Concha (Figs. 1 y 2)

discoidal algo ovalada, aplanada en la

base y con el ápice apenas saliente,

espira con elevación uniforme, suave-

mente convexa y no muy pronunciada.

Color blanquecino.

Protoconcha lisa con 1*/4 vueltas de

espira. Termina bruscamente y de forma

bien delimitada con la teloconcha.

Teloconcha formada por 1*/4 vuel-

tas. La microescultura es compleja: al

final de la última vuelta hay tres cordo-

nes espirales en posición subsutural, de

los cuales, los dos primeros se inician de

forma progresiva, mientras el tercero se

origina por una división del segundo. El

resto de la superficie presenta surcos

bastante uniformes constituidos, al prin-

cipio, por perforaciones y, posterior-

mente, por hundimientos de forma más

alargada. Esta escultura desaparece casi

por completo en el último cuarto de

vuelta de forma muy constante. Hacia la

parte inferior de la última vuelta la

concha se angula y se hace prominente

por medio de un grueso cordón que

sobresale del perfil de la concha a modo

de quilla y sobre el cual aparecen varios

cordoncillos en zigzag. Este cordón,

visto desde la parte superior de la

concha, es más visible por un lado que

por el otro, lo que aumenta el perfil

ovoide de la concha. La parte de la

última vuelta que está por debajo de la

angulación apenas sobrepasa al cordón

periférico y es cóncava en la proximidad

del mismo. En su parte central hay un

ombligo profundo. Toda la base pre-

senta cordoncillos espirales; los que

bordean el ombligo son gruesos y zigza-

gueantes, y lo mismo ocurre con los que

se sitúan sobre la quilla, siendo los res-

tantes algo más finos y uniformes.

Abertura circular, con borde fino,

engrosado en su parte externa por el

final del cordón espiral sobresaliente.

El animal es desconocido.

Discusión: La especie ahora descrita

es incluida en el género Anticlimax como

una aproximación, ya que presenta

algunas diferencias con la especie tipo.

Estas diferencias son, principalmente, la

carencia de escultura axial en la base y

la ausencia de una prolongación de la

abertura en la finalización del cordón

espiral. Sin embargo, la subespecie A.

tholus prodromus Pilsbry y Olsson, 1950,

apenas presenta escultura axial en la

base y también carece de la prolonga-

ción de la abertura, habiendo sido no

obstante incluida en este género. A. deco-

rata spec. nov., además de parecerse a

esta última especie, tiene muchas de sus

características coincidentes con especies

que han sido consideradas pertene-

cientes al género Anticlimax, como la

base aplanada o cóncava, el cordón peri-

férico, la superficie convexa por encima

del mismo, la existencia de cordones en

ROLÁN E7 4£.: Una nueva especie de Anticlimax de Cuba

Figuras 1, 2. Anticlimax decorata spec. nov. 1: paratipo, Rancho Luna, Cienfuegos, Cuba, American

Museum of Natural History de Nueva York; 2: holotipo, Rancho Luna, Cienfuegos, Cuba, Museo

Nacional de Ciencias Naturales de Madrid. Escala 0,5 mm.

Figures 1, 2. Anticlimax decorata 2. sp. 1: paratype, Rancho Luna, Cienfuegos, Cuba, American

Museum of Natural History New York; 2: holotype, Rancho Luna, Cienfuegos, Cuba, Museo Nacional

de Ciencias Naturales, Madrid. Scale bar 0.5 mm.

el dorso formados por perforaciones, y

los cordones en zigzag de la base.

Su parecido con A. athleenae (Pilsbry

y McGinty, 1946) es bastante notable,

aunque esta especie se diferencia de A.

decorata porque presenta ondulaciones

en la base, y carece, tanto en la base

como en el cordón periférico, de cordon-

cillos en zigzag. Tambien con A. tholus

(Pilsbry y McGinty, 1946) tiene un cierto”

parecido, pero el ombligo de esta última

está ocluido por un fuerte callo.

Las restantes especies incluidas en

este género, o tienen escultura axial, o el

Iberus, 15 (1), 1997

ombligo está ocluido por un callo, o

tienen una prolongación triangular en la

abertura como continuación de la quilla

periférica, por lo que son claramente

diferentes de A. decorata.

Según ESPINOSA, FERNÁNDEZ-GAR-

CÉs Y ROLÁN (1995), para la fauna de

Cuba, únicamente era conocida una

especie en el género Anticlimax: A. pro-

boscidea (Aguayo, 1949), la cual es más

elevada y tiene una gran prolongación

de la abertura.

La principal diferencia entre los dos

subgéneros, Anticlimax y Subclimax, en

los que PILSBRY Y OLSSON (1950) dividie-

ron las especies del género Anticlimax es

que, en el primero, el ombligo es evi-

dente mientras que, en el segundo,

existe un callo columelar que cubre par-

AGRADECIMIENTOS

A Bernardo Fernández Souto del

Servicio General de Apoyo a la Inves-

tigación de la Universidad de A Co-

BIBLIOGRAFÍA

AGUAYO, C. G. Y BORRO, P., 1946. Nuevos mo-

luscos del Terciario superior de Cuba. Revista

de la Sociedad Malacológica “Carlos de la Torre”,

4 (1): 9-12, 1 lám.

DaLL, W. H., 1903. Two new mollusks from

the West Coast of America. The Nautilus, 17

(4): 37-38.

ESPINOSA, J., FERNÁNDEZ GARCÉS, R. Y ROLÁN,

E., 1995. Catálogo actualizado de los molus-

cos marinos de Cuba. Reseñas Malacológicas,

9: 1-90.

cial o totalmente el ombligo. Estos

autores mencionan que no conocen

especies de estructura intermedia entre

ambos subgéneros. Anticlimax decorata,

por su ombligo abierto y carencia de

callo columelar, no estaría incluida en el

subgénero Subclimax; pero por la caren-

cia de cualquier tipo de prolongación

triangular en la abertura, tampoco pre-

sentaría perfectamente las características

del subgénero Anticlimax. Por tanto,

Anticlimax decorata representa una forma

intermedia entre ambos subgéneros.

En cualquier caso, en ausencia de

información sobre las partes blandas, es

preferible no asumir su posición taxonó-

mica genérica más que como probable y

no hacer uso de asignación subgenérica

alguna.

ruña, por la realización de las fotogra-

fías en el microscopio electrónico de

barrido.

PiLsBRY, H. A. Y MCGINTY, T. L., 1946a. “Cy-

clostrematidae” and Vitrinellidae of Flo-

rida and Bahamas, III. The Nautilus, 59: 77-

83, pl. 8.

PiLSBRY, H. A. Y MCGINTY, T. L., 1946b. Vitri-

nellidae of Florida, part. 4. The Nautilus, 60

(1) 12-18.

PiLsBRY, H. A. Y OLSSON, A. A., 1950. Review

of Anticlimax, with new tertiary species (Gas-

tropoda, Vitrinellidae). Bulletin of American Pa-

leontology, 33 (135): 105-116, lám. 17-20.

O Sociedad Española de Malacología

Iberus, 15 (1): 35-40, 1997

La familia Pyramidellidae Gray, 1840 (Mollusca, Gastro-

poda) en África occidental. 1. El género Sayella Dall, 1885

The family Pyramidellidae Gray, 1840 (Mollusca, Castiopodaa in

West Africa. 1. The genus Sayella Dall, 1885

Anselmo PEÑAS* y Emilio ROLÁN**

Recibido el 8-VIIL-1996. Aceptado el 8-X-1996

RESUMEN

Se describen dos especies nuevas del género Sayella Dall, 1885 de África Occidental,

una de ellas encontrada en los Archipiélagos de Cabo Verde y de Sáo Tomé y Príncipe, y

otra, sólo en el segundo de ellos.

ABSTRACT

Two new species of the genus Sayella Dall, 1885 from West Africa are described; one of

them found in the archipelagos of Cape Verde, and Sáo Tomé and Principe, the other one

only in the latter.

PALABRAS CLAVE: Pyramidellidae, Sayella, África occidental, especies nuevas.

KEY WORDS: Pyramidellidae, Sayella, West Africa, new species.

INTRODUCCIÓN

Después de realizada la revisión de

los piramidélidos del Mediterráneo

ibérico (PEÑAS, TEMPLADO Y MARTINEZ,

1996) y animados por los recientes tra-

bajos de NOFRONI Y SCHANDER (1994) y

SCHANDER (1994), en los que se descri-

ben 31 especies nuevas de este grupo en

África occidental, unido a la gran canti-

dad de material que poseemos, nos

decidió a abordar el estudio de esta

familia en las costas atlánticas africanas.

Comenzamos esta revisión con el género

Sayella Dall, 1885, hasta ahora no men-

cionado en el área geográfica de estudio,

y continuaremos de forma inmediata

con la de los géneros Turbonilla, Eulime-

lla, Chrysallida y otros.

El género Sayella fue descrito por

DaLL (1885) para una especie fósil del

Mioceno y bajo Plioceno de Louisiana.

Inicialmente, la especie tipo, Leuconia

hemphillii Dall, 1884, fue incluida por su

autor en la familia Ellobiidae (Pulmo-

nata). Sayella no aparece mencionado en

las revisiones genéricas de THIELE (1931-

35) o de WENZ (1938). La posición de

este género y sus relaciones con otros

Pyramidellidae son discutidas por ODE

(1994), que comenta sus diferencias con

Odostomia y la dificultad para la separa-

ción de especies, revisando también los

taxones del Atlántico occidental. Las

características del género son referidas

por ABBOTT (1974) como “concha pu-

* Carrer Olérdola, 39; 08800 Vilanova i la Geltrú. Barcelona.

** Cánovas del Castillo, 22; 36202 Vigo.

ber ISO

poide-alargada, frágil, lisa, con vueltas

convexas. Animal con tentáculos apla-

nados, triangulares. Periostraco amari-

llento”. Se ha señalado en aguas

someras y fondos fangosos (ABBOTT,

1974).

Recientemente, WISE (1996), basán-

dose en datos anatómicos, crea la subfa-

milia Sayellinae para incluir a las espe-

cies de este género. Asimismo, describe

para la especie S. crosseana (Dall, 1885)

un nuevo género, Petitiella, por sus dife-

rencias anatómicas, al que también

incluye en esta subfamilia.

En el presente trabajo abordamos la

descripción de dos especies que consi-

deramos nuevas y que asignamos provi-

sionalmente a Sayella, pués carecemos

de los datos anatómicos necesarios para

confirmar su pertenencia a este género.

Dado el escaso número de vueltas de su

protoconcha, cabe pensar que estas

especies carecen de un desarrollo larva-

rio plantotrófico, y que, por tanto, es

poco probable una relación estrecha con

especies del Atlántico occidental. Dicho

género no ha sido mencionado con

RESULTADOS

anterioridad en las costas occidentales

de África.

MATERIAL Y MÉTODOS

El material estudiado en el presente

trabajo procede de las expediciones efec-

tuadas por el segundo de los autores al

archipiélago de Cabo Verde entre los

años 1978 y 1988, y a Sáo Tomé y Prín-

cipe en los años 1989 y 1990. Dicho

material fue separado de los detritos

recogidos mediante buceo y dragado.

Abreviaturas empleadas:

AMNH: American Museum of Natural

History, New York

BMNH: The Natural History Museum,

Londres

CER: Colección de E. Rolán

CPM: Colección de P. Micali

MNCN: Museo Nacional de Ciencias

Naturales, Madrid

MNHN: Museum National d'Histoire

Naturelle, Paris

Familia PYRAMIDELLIDAE Gray, 1840

Subfamilia SAYELLINAE Wise, 1996

Género Sayella Dall, 1885

Las características del género Sayella

son: concha pequeña, pupoide alargada,

vueltas lisas, sutura poco profunda,

bandas espirales de color, protoconcha

corta con núcleo oculto, y periostraco

acastañado.

Sayella micalii spec. nov. (Figs 1-5)

Material tipo: Holotipo (Fig. 1) de 2,86 x 1,35 mm depositado en MNCN (n* 15.05/23757) y 1

paratipo, ambos de la Bahía de Santo Antonio, Príncipe, República de Sáo Tomé y Príncipe.

Otros paratipos en las siguientes colecciones: MNHN, 1 concha y dos juveniles de Bahía das

Agulhas, Príncipe; BMNH, 1 concha, y AMNH, 1 concha, ambas a -10 m, de Regona, Isla de Sal,

Archipiélago de Cabo Verde; CPM, 1 concha, -8 m, de Furna, Isla de Brava, Archipiélago de

Cabo Verde; CER, 1 concha, -2 m, de la ciudad de Sáo Tomé, República de Sao Tomé y Príncipe y

otra, -10 m, de Furna, Isla de Brava, Cabo Verde.

Localidad tipo: Bahía de Santo Antonio, en la isla de Príncipe, Archipiélago de Sáo Tomé y Prín-

cipe.

Etimología: El nombre específico está dedicado al malacólogo Pasquale Micali, experto en Pyra-

midellidae del Mediterráneo, por su habitual cooperación en el estudio de estos moluscos.

PEÑAS Y ROLÁN : El género Sayella (Pyramidellidae) en África occidental

Figuras 1-4. Sayella micalii spec. nov. 1: holotipo (MNCN), Bahía de Santo Antonio, Príncipe, Archi-

piélago de Sáo Tomé y Príncipe; 2, 3: protoconcha; 4: detalle de la columela y de la microescultura.

Figures 1-4. Sayella micalii 2. sp. 1: holotype (MNCN), Santo Antonio Bay, Principe, Archipelago of

Sáo Tomé and Principe; 2, 3: protoconch; 4: detail of the columela and microesculpture.

Descripción: Concha (Figs. 1 y 5)

oval-cónica, muy frágil.

Protoconcha (Figs. 2 y 3) obtusa,

muy grande, de tipo B (según Aartsen,

1981), con un diámetro de 490 yum. Eje

con un ángulo de unos 100” en relación

al de la teloconcha, y núcleo parcial-

mente oculto. Coloración castaña clara.

Teloconcha con vueltas convexas, algo

escalonadas, que crecen deprisa; la última

vuelta es muy grande, representando un

60% del total de su altura. Sutura pro-

funda con una débil repisa subsutural.

Vueltas de espira lisas, excepto débiles

líneas de crecimiento ligeramente proso-

clinas y estrías microscópicas espirales,

más visibles en la zona umbilical (Fig. 4).

Abertura grande, oval alargada, con

un labio externo simple, cortante. Colu-

mela ligeramente arqueada, delgada, pero

muy replegada hacia la zona umbilical;

tiene dos dientes, el mayor muy saliente

Iberus, 15 (1), 1997

y situado en la parte superior de la colu-

mela; el segundo en la parte central, en

forma de cordoncillo oblícuo. Fisura umbi-

lical profunda. Color vítreo, semitranspa-

rente, con una banda suprasutural castaña

y otra en la base de la última vuelta, en la

que también se aprecia otra banda subsu-

tural (Fig. 5). Periostraco amarillento.

Distribución: Conocida únicamente

de los Archipiélagos de Cabo Verde y

Sáo Tomé y Príncipe.

Hábitat: En sedimentos de arena y

fango, entre 4 y 10 m de profundidad.

Discusión: No hay ninguna especie

similar a Sayella micalíí spec. nov. en el

Mediterráneo ni en las costas de África

occidental. Todas las especies del Caribe

asignadas a este género (ver ABBOTT,

1974) son diferentes porque tienen más

vueltas de teleoconcha, con un incre-

mento más lento de la espira y una

forma más lanceolada que cónica.

Sayella mercedordae spec. nov. (Figs. 6-8)

Material tipo: Holotipo (Fig. 7) de 2,10 x 0,90 mm depositado en MNCN (n* 15.05/23758); 1

paratipo (Fig. 6) de Rife de Chaves, Boavista, en CER.

Localidad tipo: Bahía de Mordeira, en la isla de Sal, Archipiélago de Cabo Verde.

Etimología: El nombre específico está dedicado a Mercedes Dorda, esposa del malacólogo

Esteban Calderón, por su colaboración a lo largo de toda su vida en la creación y mantenimiento

de su colección.

Descripción: Concha (Figs. 6 y 7)

oval-cónica, pequeña, y frágil.

Protoconcha (Figs. 8) pequeña, de

tipo B, tendente a € (de acuerdo con

AARTSEN, 1981). Su eje forma un ángulo

de unos 100% en relación al de la telocon- :

cha. Núcleo oculto.

Teloconcha con vueltas convexas,

que crecen deprisa en altura y anchura.

La última vuelta es muy grande, repre-

sentando un 70% del total de la altura

de la concha. Sutura muy marcada, algo

profunda. Vueltas de espira lisas, excep-

tuando líneas de crecimiento ligera-

mente prosoclinas y estrías microscópi-

cas espirales en la base.

Abertura oval-piriforme, estrechada

hacia arriba. Columela opistoclina, algo

arqueada, sin un diente claro, pero con

un pliegue columelar en su parte cen-

tral. No hay ombligo. Color blanquecino

con dos estrechas bandas de color pardo

por vuelta, y otra más, del mismo color,

próxima a la zona columelar (Fig. 8).

AGRADECIMIENTOS

A José Bedoya por las fotografías al

MEB realizadas en el MNCN de Madrid.

Distribución: Conocida únicamente

del Archipiélago de Cabo Verde.

Hábitat: Fondos arenosos entre 3 y 5

metros de profundidad, con zonas de

fango próximas.

Discusión: Sayella micalii spec. nov.,

es mucho grande, ancha, con forma más

piramidal y con protoconcha mucho

más gruesa.

Las especies del Caribe son todas de

mayor tamaño; Sayella fusca (C. B.

Adams, 1839) tiene la sutura menos pro-

funda y una banda clara subsutural; S.

livida Rehder, 1935, tiene también una

banda clara subsutural y un mayor

número de vueltas de espira; S. crosseana

(Dall, 1885) es mucho más alargada y

proporcionalmente más estrecha; S.

hemphilli (Dall, 1889) y S. chesapeakea

Morrison, 1939, tienen la sutura apenas

marcada y un número mayor de vueltas

de espira.

A Pasquale Micali por la lectura crítica del

manuscrito.

PEÑAS Y ROLÁN : El género Sayella (Pyramidellidae) en África occidental

Figura 5. Sayella micalii spec. nov. Dibujo del paratipo (CER) mostrando la distribución de las ban-

das de color. Figuras 6-8. S. mercedordae spec. nov. 6: dibujo de un paratipo (CER) mostrando la

distribución de las bandas de color; 7: holotipo (MNCN), Bahía de Mordeira, Isla de Sal,

Archipiélago de Cabo Verde; 8: protoconcha.

Figure 5. Sayella micalii 2. sp. Drawing of paratype (CER) showing the distribution of the colour bands.

Figures 6-8. S. mercedordae 7. sp. 6: drawing of a paratype (CER) showing the distribution of the colour

bands; 7: holotype (MNCN), Mordeira Bay, Sal Island, Archipelago of Cape Verde; 8: protoconch.

BIBLIOGRAFÍA

AARTSEN, J. J. VAN, 1981. European Pyramide-

llidae: HU. Turbonilla. Bollettino Malacologico, 17

(5-6): 61-88.

ABBOTT, R. T., 1974. American seashells. Van Nos-

trand Reinhold Co. New York. 663 pp., 24

láms.

NOFRONI, I. Y SCHANDER, C., 1994. Description

of three new species of Pyramidellidae (Gas-

tropoda, Heterobranchia) from West Africa.

Notiziario CISMA, 15: 1-10.

ODE, H., 1994. Monograph. Distribution and Re-

cords of the marine Mollusca in the Northwest

Gulf of Mexico. Texas Conchologist, 31 (1): 9-32.

Iberus, 15 (1), 1997

Prxas, A., TEMPLADO, J. Y MARTINEZ, J. L., 1996.

Contribución al conocimiento de los Pyra-

midelloidea (Gastropoda: Heterostropha)

del Mediterráneo español. Iberus, 14 (1): 1-82.

SCHANDER, C., 1994. Twenty-eight new species

of Pyramidellidae (Gastropoda, Hetero-

branchia) from West Africa. Notiziario CISMA,

17: 11-78.

THIELE, J., 1931-35. Handbook of systematic mala-

cology. Gustav Fischer. 189 pp.

VAUGHT, K. C., 1989. A classification of the living

Mollusca. American malacologists. Mel-

bourne, Florida. 189 pp.

WENZ, W., 1938. Handbuch der Paláozoologie.

Von Gebrúder Borntraeger. Berlin. 1638 pp.

WISE, J. B., 1996. Morphology and philogene-

tic relationship of certain pyramidelid taxa

(Heterobranchia). Malacologia, 37(2): 443-551.

O Sociedad Española de Malacología —_—_—_—_—_—— Iberus, 15 (1): 41-93, 1997

Fauna malacológica del litoral del Garraf (NE de la Penín-

sula Ibérica)

Malacological marine fauna from Garraf coast (NE Iberian Penin-

sula)

Gonzalo GIRIBET* y Anselmo PEÑAS**

Recibido el 22-VII-1996. Aceptado el 11-X-1996

RESUMEN

Se presenta una lista de 622 especies de moluscos marinos (7 Poliplacóforos, 417 Gaste-

rópodos, 190 Bivalvos y 8 Escafópodos) recolectados en el litoral del Garraf (Barcelona,

NE de la Península Ibérica). De estas especies, 53 se citan por primera vez en el Medite-

rráneo español, siendo dos de ellas primera cita para todo el Mediterráneo, Trophon bar-

vicensis y Pleurotomella coeloraphe. De particular interés ha resultado el estudio de un

nuevo yacimiento de sedimentos Wúrmienses, asociado a una biocenosis de corales blan-

cos, entre 250 y 350 m de profundidad, y el análisis del contenido gástrico de unos tres

mil ejemplares de estrellas de mar del género Astropecten, recolectadas entre 40 y 350 m

de profundidad. Se incluyen, asimismo, comentarios sobre algunos de los taxones mencio-

nados y se ilustran al MEB muchos de ellos, con especial atención a los de las familias

Cerithiopsidae, Turridae de profundidad, Yoldiidae y Thyasiridae.

ABSTRACT

We report a checklist of 622 marine molluses (7 Poliplacophors, 417 Gastropods, 190

Bivalves and 8 Scaphopods) from “El Garraf” coast (Barcelona, NE Iberian Penninsula).

From these species, 53 are new findings for the Spanish Mediterranean, and two of them,

Trophon barvicensis and Pleurotomella coeloraphe, are reported for the first time for the

whole Mediterranean. A new Wúrm bed associated with a white coral biocenosis has

been found off Vallcarca at depths between 250 and 350 m, and is described here. Data

about molluscs identified from the gut contents of about 3000 specimens of Astropecten

sea stars found between 40 and 350 m depth are also reported. Also, we include com-

ments about some of the listed taxa and a special SEM image collections, particularly of

such groups as Cerithiopsidae, deep-sea Turridae, Yoldiidae and Thyasiridae.

PALABRAS CLAVE: Moluscos marinos, Garraf, NE Península Ibérica, Mar Mediterráneo, tanatocenosis

Wiiúrmiense, biocenosis de coral blanco, contenidos estomacales de Astropecten.

KEY WORDS: Marine molluscs, Garraf, NE Iberian Penninsula, Mediterranean Sea, Wiirm tanatocenosis,

white coral biocenosis, Astropecten gut contents.

* Departament de Biologia Animal, Facultat de Biologia, Universitat de Barcelona, Av. Diagonal 645, 08071

Barcelona.

** Carrer Olérdola 39, 08800 Vilanova i la Geltrú, Barcelona.

Iberus, 15 (1), 1997

INTRODUCCIÓN

El presente trabajo trata sobre los

moluscos marinos (exceptuando la

Clase Cephalopoda) que se han encon-

trado en el litoral de la comarca del

Garraf (Barcelona) durante más de una

década de recolección y estudio de

material. Aunque, aparentemente, la

región objeto de estudio no presente

ninguna particularidad biogeográfica o

física que la delimite desde un punto de

vista biológico, la consideramos de

especial interés, puesto que en un área

relativamente pequeña se encuentran

representados buena parte de los ecosis-

temas marinos del Mediterráneo. Ésto se

refleja en la gran diversidad de especies

encontrada en esta zona. Otro factor

importante que determina la riqueza

faunística del Garraf es la existencia de

diferentes tipos de fondos, con numero-

sos cañones submarinos, por lo que a

sólo 14 km del puerto principal (Vila-

nova i la Geltrú) se alcanzan profundi-

dades de unos 530 m, mientras que en

otras zonas próximas la distancia se tri-

plica para llegar a profundidades seme-

jantes.

Varios autores han estudiado la

fauna malacológica marina de esta

comarca (SAMA, 1916; HIDALGO, 1917;

VILELLA, 1968; Ros, 1975; BALLESTEROS

1977, 1978 Y 1984; ASENSI, 1984), que ha

ofrecido una gran riqueza en cuanto a

número de especies, pero en ninguno de

los casos anteriores se habían muestre-

ado todos los hábitats encontrados en

este litoral, o al menos no se había hecho

de una forma tan extensiva. Ya HIDALGO

(1917) citaba para la zona de estudio 307

especies de moluscos (191 Gasterópo-

dos, 108 Bivalvos, 4 Escafópodos y 1

Cefalópodo), y SAMA (1916) citaba 319

especies y 81 variedades.

ZONA DE ESTUDIO

El Garraf es una pequeña comarca

litoral situada al sur de Barcelona (Fig.

1), en cuyo interior se encuentra el

Parque Natural del Garraf. Los aproxi-

madamente 25 km de costa que presenta

esta comarca, están comprendidos entre

la desembocadura del río Foix (41* 12'

N, 1* 40” E) y punta Ginesta (41* 16' N,

1” 57' E). Tres son los municipios litora-

les que se encuentran: Sitges, Vilanova i

la Geltrú, y Cubelles, incluyendo el

primero de ellos las pedanías de Garraf

y Vallcarca. La existencia del importante

puerto pesquero de Vilanova i la Geltrú,

así como la colaboración de algunos de

los pescadores de su cofradía, han sido

factores decisivos para proporcionarnos

gran parte del material en el que está

basado este estudio.

Las playas de arena fina con pen-

dientes poco pronunciadas son domi-

nantes en la zona, aunque antes de la

construcción generalizada de espigones,

había algunas calas de arenas gruesas

con pendientes más pronunciadas,

como Cala Morisca (en Vallcarca) o

Aiguadolc (en Sitges). Los fondos de

arena fina son el hábitat típico de

algunos Nassariidae, Naticidae y varios

bivalvos, sobre todo de las especies de

aguas someras de las familias Tellinidae,

Pharidae, Donacidae, Veneridae, Mactri-

dae, Pandoridae y Thraciidae. También

se encuentran gran cantidad de escolle-

ras O espigones, que están proliferando

por toda la zona, tanto para la creación

de puertos deportivos, como para

formar playas artificiales.

Además son importantes las paredes

rocosas de los acantilados calcáreos

típicos del macizo del Garraf, que

emergen casi verticalmente de fondos

arenosos desde profundidades com-

prendidas entre 0, 5 y 4 m. Estas rocas

son poco ricas desde el punto de vista

malacológico, aunque presentan

grandes bancos de Mytilus galloprovin-

cialis (Linnaeus, 1758), y pueden llegar a

abundar especies como Thais haemastoma

(Linnaeus, 1767) y, sobre todo por

encima de la línea de marea, algunas

especies de los géneros Patella, Gibbula o

Littorina.

La desembocadura de algunas rieras

de cauce intermitente son muy intere-

santes desde el punto de vista faunís-

tico. Este es el caso de la riera de Vila-

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

41%30'

410

030' 12

Figura 1. Mapa de la zona estudiada.

Figure 1. Map of studied area.

franca, que desemboca al sur de Sitges,

y especialmente la desembocadura del

río Foix en Cubelles. Esta última zona

está formada por una base de arena fina,

casi fangosa, rica en sedimentos orgáni-

cos, cubierta casi completamente por

cantos rodados y piedras aportadas

esporádicamente por el río (suele estar

seco por la presencia de una presa de

contención situada unos kilómetros más

arriba), que sólo lleva agua en forma de

grandes avenidas. La profundidad

máxima en esta zona de cantos rodados

es de aproximadamente 1 m, y a partir

de aquí ya se encuentra el típico fondo

arenoso que caracteriza a las playas cir-

cundantes. La base pedregosa presenta

una gran riqueza malacológica, princi-

palmente de Opistobranquios. Esta zona

concreta de Cubelles ha sido previa-

mente estudiada por Ros (1975) y, prin-

cipalmente, por BALLESTEROS (1977, 1978

y 1984), y constituye la localidad tipo de

Río Foix

Barcelona '

1%30' 2

Taringa faba (Ballesteros, Llera y Ortea,

1984). Una buena descripción del recu-

brimiento algal así como de la fauna de

Invertebrados acompañante se puede

encontrar en BALLESTEROS (1984).

Situada paralelamente a la costa y a

una distancia media de ésta de aproxi-

madamente 2, 5 km, se encuentra una

pradera de Posidonia oceanica (L.) Dellile.

Hace unos 25 años, esta pradera de

fanerógamas era muy densa y extensa,

encontrándose desde los 8 m de profun-

didad (frente a Terramar, en Sitges)

hasta 22 m en algunos puntos, y su lon-

gitud era de unos 10 km de largo por

casi 2 km de anchura. Esta pradera ha

sufrido una regresión considerable

durante las dos últimas décadas, y con

ella muchas de las especies típicas de

estos hábitats, como Pinna nobilis Linna-

eus, 1758, muy común anteriormente, y

que ya prácticamente no se encuentra

viva en esta comarca litoral.

Iberus, 15 (1), 1997

Los fondos de aguas profundas se

dividen en “La Mar de Terra” y “La Mar

de Fora”, que están separadas por una

serie de rocas dispuestas paralelamente

a la costa. “La Mar de Terra” es una pla-

nicie fangosa con una profundidad

máxima que oscila entre los 74 y los 105

m, y que se halla a una distancia media

de la costa de 9,5 km. Aquí destacan

algunas zonas rocosas aisladas, varias

zonas con gorgonias (Eunicella singularis

(Esper) y Leptogorgia sarmentosa (Esper)),

en las que abunda el bivalvo Pteria

hirundo (Linnaeus, 1758). También es

importante en esta zona una amplia

extensión de concreciones calcáreas y

coralinas situadas una entre Sitges y

Vilanova, y otra frente a Vallcarca, for-

madas por típicos fondos de maérl, en

los que abundan las algas calcáreas Lit-

hothamnion calcareum (Pallas) Areschoug

y Lithothamniom corallioides Crouan,

principalmente.

“La Mar de Fora” es más variada y

generalmente escarpada. Su profundi-

dad va desde los 105 hasta los 1600 m en

el canal de Foix. Destacan entre el S y SO

de Vilanova una zona de barrancos sub-

marinos que convergen hacia una pro-

fundidad de unos 500 m, un caladero

muy rico en el centro, y unas planicies en

cuyos límites, frente a Vallcarca, se en-

cuentran los fondos de “El Parrusset”.

Éste es un cañón submarino profundo

(los pescadores faenan a profundidades

entre 200 y 450 m), de fondo rico en nó-

dulos de ferromanganeso y que alberga

una biocenosis de coral blanco (sensu

PÉRES Y PICARD, 1964), asociada a una ta-

natocenosis de fauna Wúrmiense de

gran interés malacológico.

MATERIAL Y MÉTODOS

El presente trabajo-está basado en el

material recolectado por los autores o

proporcionado por pescadores durante

más de una década. También se ha revi-

sado material de colecciones malacoló-

gicas privadas de la zona de estudio.

El material ha sido recolectado en

fondos arenosos, escolleras, rocas o

acantilados mediante inmersión, y si-

multaneando con visitas a las diferentes

playas de la zona, especialmente des-

pués de los temporales. Hay que tener

en cuenta que las especies de aguas pro-

fundas que aparecen ocasionalmente en

las playas o en aguas someras, suelen

proceder de los restos arrojados al mar

por los barcos pesqueros, o de la lim-

pieza de las redes de pesca antes de en-

trar en el puerto.

La zona de la desembocadura del río

Foix en Cubelles se ha muestreado quin-

cenalmente durante más de un año

(junio de 1992 a agosto de 1993) de una

forma exhaustiva, volteando todas las

piedras en dos transectos, uno de 15 x 1

m paralelo a la línea de costa y a 0,5 m

de profundidad, y otro de 25 x 1 m, per-

pendicular a la costa y a una profundi-

dad comprendida entre 0 y 1 m.

El material de aguas profundas ha

sido proporcionado por pescadores de

arrastre. Además, en más de 20 ocasio-

nes durante el período de muestreo, se

ha acompañado a los pescadores con

objeto de separar el material por hábi-

tats y profundidades in situ, para

obtener una información más detallada

sobre el mismo.

Se ha analizado el contenido estoma-

cal de unos 3000 ejemplares de los aste-

roideos Astropecten aranciacus (L.) y As-

tropecten irregularis (Linck), procedentes

de más de 50 arrastres, principalmente

de fondos fangosos entre 40 y 350 m de

profundidad. Se han estudiado también

seis muestras de detrito fangoso (aproxi-

madamente 20 kg en total) procedente

de “El Parrusset”, entre 200 y 350 m de

profundidad, recolectadas entre 1994 y

1996. El detrito ha sido lavado y pasado

por una serie de tamices, siendo el más

fino de 0,4 mm de luz de malla. Además

se han analizado unos 30 kg de sedi-

mentos obtenidos en playas de la zona;

unos 5 kg de sedimentos arenoso de 2 m

de profundidad obtenidos en Sitges; y 1

kg de sedimento arenoso de 2 m de pro-

fundidad obtenido en el interior del

puerto de Vallcarca. Una gran cantidad

de micromoluscos se ha obtenido del es-

tudio de estos sedimentos, lo que per-

mite obtener un elevado número de es-

pecies, aunque de la mayor parte de

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

ellas se hallen sólo conchas (tanatoceno-

sis), por lo que no es posible precisar sus

hábitats.

La mayor parte del trabajo de Opis-

tobranquios se ha basado en las publica-

ciones de BALLESTEROS (1977, 1978 y

1984), Ros (1975) y ASENSI (1984), y

siempre que el material no haya sido

recolectado por los autores, se indica la

cita bibliográfica de la cual proviene.

Además, se han revisado las colec-

ciones de A. Tubau, M. Roca y P. Ortoll,

malacólogos aficionados o pescadores

de Vilanova i la Geltrú. No ha sido

posible, sin embargo localizar la colec-

ción de SAMA (1916), compuesta por 400

especies de moluscos procedentes del

litoral entre Vilanova i la Geltrú (Barce-

lona) y Calafell (Tarragona).

El material fotografiado al microsco-

pio electrónico de barrido (M.E.B.), ha

sido previamente hervido en agua desti-

lada y tratado con ultrasonidos, con el

objeto de eliminar las impurezas deposi-

tadas en las conchas, aunque en algunos

casos, cuando las conchas eran dema-

siado finas, no se ha realizado el trata-

miento con ultrasonidos. Las muestras

han sido fotografiadas en un MEB

Hitachi S-2300 a 15KV. En algunos casos

se han seleccionado para fotografiar

ejemplares procedentes de otras zonas,

con el objeto de ilustrar los ejemplares

mejor conservados.

El listado de especies ha sido confec-

cionado siguiendo a SABELLI, GIANUZZI-

SAVELLI Y BEDULLI (1990), excepto para

algunos taxones, para los que se han

empleado revisiones taxonómicas más

recientes.

Parte del material aquí tratado ha

sido cedido al Museu del Mar de Vila-

nova 1 la Geltrú.

RESULTADOS

El número total de especies de

moluscos marinos recogidos en este

trabajo es de 622 (7 Poliplacóforos, 417

Gasterópodos, 190 Bivalvos y 8 Escafó-

podos), correspondiendo aproximada-

mente un 2,7% a especies que se han

encontrado exclusivamente en forma

subfósil en el Garraf, algunas de las

cuales no viven actualmente en el Medi-

terráneo, O habitan en zonas más pro-

fundas.

Lista de especies (Tabla I): A la

izquierda aparece el nombre de cada

especie, que irá en negrita en el caso de

que sea objeto de comentarios en la dis-

cusión, irá precedida de un asterisco (*)

cuando constituya primera cita en el

Mediterráneo español, y de dos (**)

cuando constituya primera cita en el

Mediterráneo en general. A continua-

ción se describe brevemente el tipo de

hábitat donde se ha encontrado la

especie (lo cual algunas veces no refleja

su hábitat real) y el rango batimétrico:

“s” (supralitoral), “m” (mesolitoral), “1”

(infralitoral: de O a 30 m), “c” (circalito-

ral: de 30 a 200 m), y “b” (batial: más de

200 m). En algunos casos no se disponía

de estos datos, por lo que no se regis-

tran. En el caso de las especies fósiles,

tampoco se indica el hábitat. En la

siguiente columna se señalan, con un

número, las especies ilustradas, indi-

cando dicho número el de la figura

correspondiente. A continuación se

señala la abundancia (+: 1-2 ejemplares,

++: 3-10, +++: 11-100, ++++: más de

100), y se identifican con una “p” a las

especies procedentes del detrito de “El

Parrusset” y con una “f” a aquellas que

han sido halladas fósiles. En el caso de

que las letras “p” y “f” aparezcan entre

paréntesis, significa que la especie en

cuestión no ha sido hallada exclusiva-

mente en “El Parrusset” o no ha sido

encontrada exclusivamente fósil, respec-

tivamente. También se indica con una

“y” si la especie ha sido hallada viva en

el área de estudio, y con “Aa” o “Ai” se

señalan las especies obtenidas en conte-

nidos estomacales de Astropecten aran-

ciacus O A. irregularis, respectivamente.

Evidentemente no ha sido posible espe-

cificar todos los ambientes donde se han

recolectado las muestras, y es por esto

que nos hemos limitado a mencionar la

procedencia de aquellas muestras obte-

nidas de estas formas particulares.

Iberus, 15 (1), 1997

Tabla I. Listado de especies encontradas en el área de estudio, hábitat donde se han encontrado,

rango batimétrico, figuras en las que están representadas, abundancia y procedencia.

Las especies en negrita están comentadas en el texto. No se incluyen los datos no disponibles de bari-

metría, ni de hábitat en las especies fósiles. En las especies de opistobranquios que no han sido reco-

lectadas por los autores se incluye la referencia bibliográfica de donde procede la cita.

Códigos. *: primera cita en el Mediterráneo español; **: primera cita en el Mediterráneo; s: suprali-

toral; m: mesolitoral; i infralitoral (0-30 m); c: circalitoral (30-200 m); b: batial (>200 m); +: 1-2 ejem-

plares; ++: 3-10 ejemplares; +++: 11-100 ejemplares; ++++: más de 100 ejemplares; p: especie proce-

dente del detrito de El Parrusset; f: especies halladas fósiles; (p): especie hallada no sólo en El Parrus-

set; (£): especie hallada no exclusivamente fósil; v: especie encontrada viva en el área de estudio; Aa:

especie obtenida en contenido estomacal de Astropecten aranciacus, Ai: idem de Astropecten irregularis.

Table I. List of species found in the study area, habitat where they have been collected, bathymetric range,

figures when included, abundance and other data related with their collection.

Species in bold are discussed in the text. Data on bathymetric range are included only when known, habi-

tat of fossil species always excluded. A bibliographic reference is given for the opisthobranch species not

collected by the authors.

Codes. *: first record in the Spanish Mediterranean; **: first record in the Mediterranean Sea; s: upper

littoral; m: midlittoral; i lower littoral (0-30 m); c: circa littoral (30-200 m); b: bathyal (>200 m); +:

1-2 specimens; ++: 3-10 specimens; +++: 11-100 specimens; ++++: more than 100 specimens; p: spe-

cies found in El Parrusset detritus; f. species found fossil; (p): species found not only in El Parrusset; (f):

species found not only fossil; v: species found alive in the study area; Aa: species collected in Astropecten

aranciacus gut contents; Ál: idem in Asttopecten irregularis gut contents.

Close POLYPLACOPHORA

Familia LEPTOCHITONIDAE

Lepidopleurus cajetanus (Poli, 1791): piedras, ¡ + ov

Familia ISCHNOCHITONIDAE

Callochiton septemvalvis euplaeae (0. G. Costa, 1829): algas (Payssonnelia) y conchas muertas, ic Hov

Lepidochitona cinerea (Linnaeus, 1767): piedras y espigones, mi H+ oV

Lepidochitona corrugata (Reeve, 1848): piedras y espigones, mi H+ oV

Familia CHITONIDAE

Chiton olivaceus Spengler, 1797: piedras y espigones, m:i HE oV

Familia ACANTHOCHITONIDAE

Acanthochitona crinita (Pennant, 1777): piedras, ¡ ++ v A

Acanthochitona fascicularis (Linnaeus, 1767): piedras y conchas muertas, ic Hov

Clase GASTROPODA

Familia PATELLIDAE

Potella caerulea Linnaeus, 1758: rocas y espigones, mii HH v

Patella rustica Linnaeus, 1758: rocas y espigones, s HH V

Patella ulyssiponensis Gmelin, 1791: rocas y espigones, ¡ H+-ov

Familia ACMAEIDAE

Acmaea virginea (0. E. Múller, 1776): sedimentos en zonas rocosas, i ++

Familia LEPETIDAE

lothia fulva (O. F. Múller, 1776): b + pÍ

Familia COCCULINIDAE

Coccopigya sp.: b + Pp

Familia LEPETELLIDAE

Lepetella cfr. espinosae Dantart y Luque, 1994: b H+ op

Familia ADDISONIIDAE

Addisonia excentrica Tiberi, 1857: en capsula ovígeras de Scyliorrhinus, ca Fig.3 ++ (p),v

Familia NERITIDAE

Smaragdia viridis (Linnaeus, 1758): fango, ic ++ Mo

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Familia FISSURELLIDAE

Fissurella nubecula (Linnaeus, 1758): en rocas y piedras, mi

Diodora gibberula (Lamarck, 1822): en piedras, ¡

Diodora graeca (Linnaeus, 1758): en rocas y piedras, ¡

Emarginula fissura (Linnaeus, 1758): b

Emarginula octaviana Coen, 1939: ic

*Emarginula pustula Thiele in Kuester, 1913: b

Emarginula rosea T. Bell, 1824: cb

Familia SCISSURELLIDAE

Anatoma aspera (Philippi, 1844): b

Familia HALIOTIDAE

Haliotis tuberculata lamellosa Lamarck, 1822: ¡

Familia TROCHIDAE

Clanculus cruciatus (Linnaeus, 1758): piedras, ¡

Clanculus jussievi (Payraudeau, 1826): piedras, ¡

Jujubinus exasperatus (Pennant, 1777): ic

Jujubinus montagui (W. Wood, 1828): fango, c

Jujubinus striatus (Linnaeus, 1758): fango, c

Gibbula albida (Gmelin, 1791)

Gibbula magus (Linnaeus, 1758): cascajo y múerl, c

Gibbula racketti (Payraudeau, 1826): piedras, ¡

Gibbula fanulum (Gmelin, 1791)

Gibbula guttadauri (Philippi, 1836)

Gibbula leucophaea (Philippi, 1836)

Gibbula philberti (Récluz, 1843): piedras y espigones, m

Gibbula richardi (Payraudeau, 1826): piedras, m

Gibbula varia (Linnaeus, 1758): piedras, m

Gibbula divaricata (Linnaeus, 1758): piedras, m

Osilinus articulatus Lamarck, 1822: piedras, m

Osilinus turbinatus (Bor, 1778): piedras, m

Calliostoma conulus (Linnaeus, 1758): rocas y múers, ic

Calliostoma dubium (Philippi, 1844)

Calliostoma laugieri laugieri (Payraudeau, 1826): i

Calliostoma zizyphinum (Linnaeus, 1758)

Calliostoma granulatum (Born, 1778): fango, cb

Danilia otaviana (Cantraine, 1835): b

Familia SKENEIDAE

Dikoleps pusilla (Jeffreys, 1847): b

*Lissotesta gittenbergeri (van Aortsen y Bogi, 1988): b

Familia TURBINIDAE

Bolma rugosa (Linnaeus, 1767): rocas y fango, c-b

Familia COLLONIIDAE

Homalopoma sanguineum (Linnaeus, 1758)

Familia TRICOLIIDAE

Tricolia pullus (Linnaeus, 1758): ¡

Tricolia speciosa (von Múhlfeldt, 1824): ¡

Tricolia tenvis (Michaud, 1829):

Familia CERITHIIDAE

Cerithium alucaster (Brocchi, 1814): fango y múerl, i-c

Cerithium lividulum Risso, 1826

Cerithium vulgatum Bruguiére, 1792: fango y múerl, i-c

Bittium latreillei (Payraudeau, 1826): ix

Bittium reticulatum (da Costa, 1778): b

Bittium submamillatum (Rayneval y Ponzi, 1854): fango y fondos detríticos, c

Fig. 4 +

+++

Iberus, 15 (1), 1997

Familia FOSSARIDAE

Fossarus ambiguus (Linnaeus, 1758): ¡

“Familia SILIQUARIIDAE

Tenagodus obtusus (Schumacher, 1817)

Familia TURRITELLIDAE

Turritella communis Risso, 1826: fango, c

Turritella monterosatoi Kobelt, 1888: fango, c

Familia LITTORINIDAE

Littorina neritoides (Linnaeus, 1758): rocas y espigones, s

Littorina punctata (Gmelin, 1791): rocas y espigones, s

Familia SKENEOPSIDAE

Skeneopsis planorbis (Fabricius, 1780): arena, ¡

Familia RISSOIDAE

Rissoa auriscalpium (Linnaeus, 1758): i

Rissoa decorata Philippi, 1846: ¡

*Rissoa gemmula (Fischer in de Folin, 1871): i Figs. 2, 20-21

Rissoa querinii Récluz, 1843: ¡

Rissoa labiosa (Montagu, 1803): ¡

Rissoa lia (Monterosato, 1884 ex Benoit ms.): i

Rissoa monodonta Philippi, 1836: ¡

Rissoa similis Scacchi, 1836: ¡

Rissoa ventricosa Desmarest, 1814: ¡

Rissoa violacea Desmarest, 1814: ¡

Alvania beani (Hanley in Thorpe, 1844): ¡

Alvania cancellata (da Costa, 1778): ¡

Alvania cimex (Linnaeus, 1758): i

Alvania cimicoides (Forbes, 1844): fango y detrito coralígeno, c-b

Alvania discors (Allan, 1818): ¡

Alvania geryonia (Nardo, 1847 ex Chiereghini ms.): ¡

Alvania lactea (Michaud, 1832): ¡

Alvania lineata Risso, 1826: ¡

Alvania punctura (Montagu, 1803): fango y detrito coralígeno, cb

Alvania rudis (Philippi, 1844): i

Alvania subcrenulata (B. D. D., 1884): ¡

* Alvania subsoluta (Aradas, 1847): b Figs. 8, 11, 12

Alvania testae (Aradas y Maggiore, 1844): fango y detrito coralígeno, c-b Figs. 7, 9, 10

*Alvania zylensis Gotas y Warén, 1982: b Figs. 13, 14

Alvania semistriata (Montagu, 1808): ib

Alvania carinata (da Costa, 1778): i

*Benthonella tenella (Jeffreys, 1869): fango, c

Manzonia crassa (Kanmacher, 1798): i

Manzonia zetlandica (Montagu, 1815): b

Obtusella intersecta (S. W. Wood, 1857): b

Obtusella macilenta (Monterosato, 1880): fango y detrito coralígeno, cb Fig. 6

Pusillina inconspicua (Alder, 1844): ib Figs. 17, 18, 19

Pusillina philippi (Aradas y Maggiore, 1844): +b Figs. 15, 16

Pusillina radiata (Philippi, 1836): ¡

Setia maculata (Monterosato, 1869): ¡

Rissoina bruguierei (Poyraudeau, 1826): ¡

Familia ADEORBIDAE

Circulus striatus (Philippi, 1836): ¡

Familia ASSIMINEIDAE

Paludinella sicana (Brugnone, 1876): ¡

+++

(p), v, Ai

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Familia CAECIDAE

Caecum auriculatum de Folin, 1868: ¡

Caecum clarkii Carpenter, 1858: ¡

Caecum trachea (Montagu, 1803): ¡

Familia HYDROBIIDAE

Ventrosia ventrosa (Montagu, 1803)

Familia IRAVADIIDAE

Ceratia proxima (Forbes y Hanley, 1850 ex Alder ms.): fango y detrito coralígeno, c-b

Hyala vitrea (Montagu, 1803): fango y detrito coralígeno, c-b

Familia TORNIDAE

Tornus subcarinatus (Montagu, 1803): i

Familia TRUNCATELLIDAE

Truncatella subcylindrica (Linnaeus, 1767): i

Familia APORRHAIDAE

Aporrhais pespelecani (Linnaeus, 1758): fango, c

Aporrhais serresianus (Michaud, 1828): fango, cb

Familia VANIKORIDAE

* Tolassia dagueneti (de Folin, 1873): b

Familia CALYPTRAEIDAE

Colyptraea chinensis (Linnaeus, 1758): en conchas muertas, cb

Crepidula fornicata (Linnaeus, 1758)

Crepidula unguiformis Lamarck, 1822: en conchas muertas, cb

Familia CAPULIDAE

Capulus ungaricus (Linnaeus, 1758): sobre conchas, c-b

Familia XENOPHORIDAE

Xenophora crispa (Koenig, 1825): fango y fondos detríticos, b

Familia VERMETIDAE

Vermetus triquetrus Bivona, 1832: rocas, ¡

Serpulorbis arenaria (Linnaeus, 1767): rocas, ¡

Familia CYPRAEIDAE

Erosaria spurca (Linnaeus, 1758)

Luria lurida (Linnaeus, 1758)

Zonaria pyrum (Gmelin, 1791): múer, c

Familia OVULIDAE

Aperiovula adriatica (G. B. Sowerby |, 1828): c

Neosimnia spelta (Linnaeus, 1758): sobre Eunicella, c

Pseudosimnia camea (Poiret, 1789): c

Familia LAMELLARIIDAE

Lamellaria latens (0. F. Múller, 1776): ¡

Familia TRIVIIDAE

Trivia arctica (Pulteney, 1789): ¡

Trivia monacha (da Costa, 1778): ¡

Trivia multilirata (G. B. Sowerby 11, 1870)

Erato voluta (Montagu, 1803)

Familia NATICIDAE

Naticarius cruentatus (Martyn, 1784): arena y fango, ic

Naticarius dillwyni (Payraudeau, 1826): arena, i

Naticarius punctatus (Chemnitz in Karsten, 1789)

Naticarivs vittatus (Gmelin, 1791)

Tectonatica filosa (Philippi, 1844): fango, c

Lunatia fusca (Blainville, 1825): fango, c-b

Lunatia guillemini (Poyraudeau, 1826): fango, ic

Lunatia macilenta (Philippi, 1844): arena y fango, ic

Lunatia nitida (Donovan, 1804): arena y fango, i-c

Payraudeautia intricata (Donovan, 1804): fango y múerl, i-c

(p), v, Ai

(p), v Ai

==" MES

Iberus, 15 (1), 1997

Familia TONNIDAE

Tonna galea (Linnaeus, 1758) +

Familia CASSIDAE

Galeodea echinophora (Linnaeus, 1758): fango, c HE 0V

Galeodea rugosa (Linnaeus, 1771): fango, cb ++ (p),v

Phalium granulatum (Born, 1778): fango, c H+ 0V

Phalium saburon (Bruguiére, 1792): fango, c ++ Y

Familia RANELLIDAE

Ranella olearia (Linnaeus, 1758) + pf

Cymatium corrugatum (Lamarck, 1816): fango, c HH V

Cymatium parthenopeum parthenopeum (von Salis, 1793): rocas, ¡ + Y

Cobestana cutacea cutacea (Linnaeus, 1767): fango y rocas, ic ++ v

Charonia lampas lampas (Linnaeus, 1758) +

Familia ATLANTIDAE

Atlanta peronii Lesueur, 1817: fango y detrito coralígeno, b ++ (p), Ai

Oxygyrus keraudrenii (Lesueur, 1817): b Hop

Familia TRIPHORIDAE

Marshallora adversa (Montagu, 1803): fondos detríticos, ¡ ++ vA

Monophorus erythrosomus (Bouchet y Guillemot, 1978): ¡ ++

Monophorus perversus (Linnaeus, 1758): ¡ ++

* Obesula marinostri Bouchet, 1985: b + op

Similiphora similior (Bouchet y Guillemot, 1978): i +

Metaxia metaxae (delle Chiaje, 1828): ¡-b ++ (p), (f)

Familia CERITHIOPSIDAE

* Cerithiopsis diadema Monterosato, 1874 ex Watson ms.: b Fig. 22 + Pp

Cerithiopsis jeffreysi Watson, 1885: b Fig.23 ++ (p), (1)

Cerithiopsis minima (Brusina, 1865): i Fig. 24 +++

Cerithiopsis nana Jeffreys, 1867: ib Figs. 25,29 ++ (p), (1

Cerithiopsis scalaris (Monterosato, 1877): b Fig. 26 +++ (p), (f)

* Cerithiopsis tiara Monterosato, 1874 ex Watson ms.: b Fig. 27 + (p), (1)

Cerithiopsis tubercularis (Montagu, 1803): +b Figs. 28,30 +++ (p), (f)

Familia ACLIDIDAE

*Adlis attenuans Jeffreys, 1883: fango y detrito coralígeno, b ++ (p), v Ai

Aclis gulsonae (W. Clark, 1850): fango y detrito coralígeno, b + pvA

Cima minima (Jeffreys, 1858): b ++ (p)

* Cioniscus gracilis Monterosato, 1874, ex Jeffreys ms.: b ++ (p)

Graphis albida (Kanmacher, 1798): ib Figs. 32, 33,34 ++ (p)

Familia EPITONIDAE

* Epitonium aculeatum (Allan, 1818): fango y detrito coralígeno, b ++ (p),v Ai

Epitonium olgerianum (Weinkauff, 1866): b ++ (p)

*Epitonium celesti (Aradas, 1854): b + p

Epitonium clathratulum (Kanmacher, 1798): b Fig.39 ++ (p)

Epitonium commune (Lamarck, 1822): rocas y fango, i-c +++ v A

*Epitonium dendrophylliae Bouchet y Warén, 1986: b 509

* Epitonium hispidulum (Monterosato, 1874): b ++ (p)

* Epitonium linctum (de Boury y Monterosato, 1890): b Fig. 40 + p

Epitonium pulchellum (Bivona, 1832): ¡ ++

Epitonium turtonis (Turton, 1819): fango, c +++ vw A

Cirsotrema cochlea (6. B. Sowerby Il, 1844): ¡ +

Gyroscala lamellosa (Lamarck, 1822): rocas y arena, i st

*Opalia abboti Clench y Turner, 1952: b 5

Opalia crenata (Linnaeus, 1758): i ++

Opalia hellenica (Forbes, 1844): b 0d)

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Familia EULIMIDAE

Eulima bilineata Alder, 1848: fango y detrito coralígeno, b

Eulima glabra (da Costa, 1778): ¡

*Crinophtheiros sp.: fango, b

* Entoconcha mirabilis Múller, 1852: b

Melanella alba (da Costa, 1778): i

Melanella boscii (Payraudeau, 1827): ¡

Melanella praecurta (Pallary, 1904): fango, b

Parvioris ibizenca (Nordsieck, 1968): ¡

Sticteulima jeffreysiana (Brusina, 1869): detrito coralígeno, b

Vitreolina perminima (Jeffreys, 1883): detrito coralígeno, b

Vitreolina sp.: detrito coralígeno, b

Familia MURICIDAE

Bolinus brandaris (Linnaeus, 1758): fango, arena y piedras, ¡

Hadriania craticuloides (Vokes, 1964): fango y detrito coralígeno, c-b

Hexaplex trunculus (Linnaeus, 1758): fango, rocas y espigones, ¡

Murexul aradasii (Poirier, 1883 ex Monterosato ms.)

Muricopsis cristatus (Brocchi, 1814): rocas y detritos, ic

Ocenebra erinaceus (Linnaeus, 1758): rocas y detritos, ix

Ocinebrina aciculata (Lamarck, 1822): fango y piedras, ix

Ocinebrina edwardsi (Payraudeau, 1826): rocas y piedras, i

*Trophon barvicensis (Johnston, 1825)

Trophon echinatus (Kiener, 1840): b

Trophon sp.: b

Irophon muricatus (Montagu, 1803): fango y detrito coralígeno, b

Familia BUCCINIDAE

Buccinum humphreysianum Bennet, 1824: b

Buccinum undatum Linnaeus, 1758: fango, c

Buccinulum cormeum (Linnaeus, 1758): fango y rocas, ic

Chauvetia brunnea (Donovan, 1804): ¡

Chauvetia turritellata (Deshayes, 1835): i

Colus jeffreysianus (Fischer, 1868): i

Neptunea contraria (Linnaeus, 1771)

Pisania striata (Gmelin, 1791)

Contharus dorbignyi (Payraudeau, 1826): i

Familia CORALLIOPHILIDAE

Coralliophila meyendorffi (Calcara, 1845): piedras y fango, i

Coralliophila panormitana (Monterosato, 1869): b

Coralliophila squamosa (Bivona, 1831): fango, cb

Familia FASCIOLARIIDAE

Fusinus pulchellus (Philippi, 1844): fango, ¡

Fusinus rostratus (Olivi, 1792): fango, c

Fusinus rudis (Linnaeus, 1758): fango, i

Familia NASSARIIDAE

Nassarius corniculus (Olivi, 1792): piedras, ¡

Nassarius cuvierii (Payraudeau, 1826): arena, ¡

Nassarius incrassatus (Stróm, 1768): piedras, ¡

Nassarius mutabilis (Linnaeus, 1758): arena y fango, i

Nassarius nitidus (Jeffreys, 1867): i

Nassarius pygmaeus (Lamarck, 1822): fango y piedras, ix

Nassarius reticulatus (Linnaeus, 1758): arena, i

Nassarius unifasciatus (Kiener, 1835)

Naytiopsis granum (Lamarck, 1822): arena, ¡

Cyclope neritea (Linnaeus, 1758)

Figs. 37, 38 ++

Figs. 35,36 +++

Fig. 45 +

Figs. 41,42. ++

Figs. 43,44 ++

Figs. 46,47 +++

(p), y Ai

Iberus, 15 (1), 1997

Familia THAIDIDAE

Orania fusulus (Brocchi, 1814)

Stramonita haemastoma (Linnaeus, 1766): rocas, ¡

Familia COLUMBELLIDAE

Columbella rustica (Linnaeus, 1758): rocos y algas, i

Mitrella minor (Scacchi, 1836)

Mitrella scripta (Linnaeus, 1758)

Familia COSTELLARIIDAE

Vexillum ebenus (Lomarck, 1811): ¡

Vexillum tricolor (Gmelin, 1790)

Familia MARGINELLIDAE

Gibberula caelata (Monterosato, 1877)

bibberula miliaria (Linnaeus, 1758): arena, ¡

Gibberula philippii (Monterosato, 1877): ¡

Gibberulo turgidula (Locard y Caziot, 1900): fango, e

Volvarina mitrella (Risso, 1826)

Granulina clandestina (Brocchi, 1814): b

Familia MITRIDAE

Mitra zonata Marryot, 1818: fango y múerl, ic

Familia CANCELLARIIDAE

Concellaria cancellata (Linnaeus, 1767): arena y fango, ic

Cancellaria similis Sowerby, 1833: fango, cb

Familia CONIDAE

Conus ventricosus (Gmelin, 1791): i

Familia TURRIDAE

Bela brachistoma (Philippi, 1844): fango y detrito coralígeno, c-b

Belo laevigata (Philippi, 1836): arena, i

Bela menkhorsti van Aartsen, 1988: b

Bela nebula (Montagu, 1803): arena fangosa, i

Bela ornata (Locard, 1897): fango, ic

Bela zonata (Locard, 1892): fango, c

Mongelia attenuata (Montagu, 1803): fango y detrito coralígeno, b

Mangelia costata (Donovan, 1804): fango y detrito coralígeno, cb

Mangelia cfr. goodalii Reeve, 1846: i

Mangelia nuperrima (Tiberi, 1855): fango y detrito coralígeno, cb

Mangelia paciniana (Calcara, 1839): arena, ¡

Mangelia serga (Dall, 1881): fango y detrito coralígeno, c-b

Mangelia smithi (Forbes, 1844): arena, fango y detrito coralígeno, cb

Mangelia stossiciana (Brusina, 1869): ¡

Mangelia unifasciata Deshayes, 1835: arena y fango, ¡

Mangelia vauquelini (Payraudeau, 1826): orena, ¡

Mangiliella bertrandii (Poyraudeau, 1826): ¡

Mangiliella taeniata (Deshayes, 1835): ¡

Taranis moerchi (Malm, 1861): detrito coralígeno, b

Taranis sp.: detrito coralígeno, b

Microdrilia loprestiana (Calcara, 1841): fongo y detrito coralígeno, cb

Haedropleura septangularis (Montagu, 1803): en Posidonia, i

**Pleurotomella coeloraphe (Dautzenberg y Fischer, 1896): b

*Pleurotomella demosia (Duutzenberg y Fischer, 1896): b

Crassopleura maravignae Bivona, 1838: ic

Mitrolumna olivoidea (Cantraine, 1835): ¡

Raphitoma aequalis Jeffreys, 1867: fango, c-

Raphitoma bicolor (Risso, 1826): arena, ¡

Raphitoma concinna (Scacchi, 1836): arena, ¡

Figs. 50, 51

Figs. 52, 53

Figs. 54, 55

Figs. 57, 58, 59

Figs. 60, 61, 62

Fig. 56

Figs. 63, 64, 65

Figs. 66, 67, 68

(p), Ai

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

*Raphitoma cordieri (Payraudeau, 1826): ¡

Raphitoma echinata (Brocchi, 1814): i

Raphitoma horrida Monterosato, 1844: en Posidonia, ¡

Raphitoma leufroyi (Michaud, 1828): arena, ¡

Raphitoma linearis (Montagu, 1803): arena, ¡

Raphitoma cfr. nivea (Marshall in Sykes, 1906): i

*Raphitoma pupoides (Monterosato, 1884): i

Comarmondia gracilis (Montagu, 1803): fango y arena, i-c

Teretia teres (Forbes, 1844): fango y detrito coralígeno, b

Familia TJAERNOEIDAE

Tjaernoeia exquisita (Jeffreys, 1883): b

Familia ARCHITECTONICIDAE

Basisulcata lepida (Bayer, 1942): mier, c

Heliacus alleryi (Seguenza, 1876): b

Heliacus architae (0. 6. Costa, 1867): b

Familia MATHILDIDAE

Mathilda cochlaeformis Brugnone, 1873: b

Familia OMALOGYRIDAE

Omalogyra atomus (Philippi, 1841): ¡

Ammonicera fischeriana (Monterosato, 1869): i

Familia PYRAMIDELLIDAE

Tiberia minuscula (Monterosato, 1880): fango y detrito coralígeno, c-

Chrysallida brattstroemi Warén, 1991: b

Chrysallida brusinaí (Cossmann, 1921): i

Chrysallida dollfusí (Kobelt, 1903): b

Chrysallida emaciata (Brusina, 1866): ¡

Chrysallida excavata (Philippi, 1836): ¡

Chrysallida fenestrata (Jrffreys, 1848): c

Chrysallida flexuosa (Monterosato, 1874 ex Jeffreys): fango y detrito coralígeno, cb

Chrysallida ghisottii van Aartsen, 1984: i

Chrysallida indistincta (Montagu, 1808): ¡

Chrysallida intermixta (Monterosato, 1884)

Chrysallida interstincta (J. Adams, 1797):

Chrysallida juliae (de Folin, 1872): fango, c

Chrysallida palazziiMicali, 1984: fango y detrito coralígeno, cb

Chrysallida pellucida (Dillwyn, 1817)

Chrysallida suturalis (Philippi, 1844): fango y detrito coralígeno, c-b

Odostomella doliolum (Philippi, 1844): ib

Euparthenia bulinea (Lowe, 1841): arena, ¡

Euparthenia humboldti (Risso, 1826)

Eulimella acicula (Philippi, 1836)

Eulimella ataktos Warén, 1991: fango y detrito coralígeno, cb

Eulimella bogii van Aartsen, 1994: b

Eulimella scillae (Scacchi, 1835): fango y detrito coralígeno, cb

Eulimella unifasciata (Forbes, 1844): fango y detrito coralígeno,

Eulimella ventricosa (Forbes, 1844): fango y detrito coralígeno, c-b

Puposyrnola minuta (H. Adams, 1869): fango y detrito coralígeno, b

Odostomia acuta Jeffreys, 1848: fango y arena, ic

Odostomia afzelii (Warén, 1991): fango y detrito coralígeno, c-b

Odostomia carrozzai van Aartsen, 1987: ¡

Odostomia clavulus (Lovén, 1846): fango y detrito coralígeno, cb

Odostomia conoidea (Brocchi, 1814): fango y detrito coralígeno, ib

Odostomia erjaveciana Brusina, 1869: arena, ¡

Odostomia eulimoides Hanley, 1844: ¡

Fig. 69 +++

p, Y, Ai

(p), v Ai

Iberus, 15 (1), 1997

Odostomia hansgei (Warén, 1991): fango y detrito coralígeno, b

Odostomia kromi van Aartsen, Menkhorst y Gittenberger, 1984: ¡

Odostomia lukisii Jeffreys, 1859: i

Odostomia megerlei (Locard, 1886)

Odostomia plicata (Montagu, 1803): ¡

Odostomia scalaris MacGillivray, 1843: ¡

Odostomia striolata Forbes y Hanley, 1850: ib

Odostomia suboblonga Jeffreys, 1884: b

Odostomia turriculata Monterosato, 1869: ¡

Odostomia turrita Honley, 1844: ib

Odostomia umbilicaris Malm, 1863: fango y detrito coralígeno, c-b

Odostomia unidentata (Montagu, 1803): fango y detrito coralígeno, ib

Odostomia verduini van Aartsen, 1987: ¡

Noemiamea dolioliformis (Jeffreys, 1848): ¡

Ondina dilucida (Monterosato, 1844): fango, c

Ondina obliqua (Alder, 1844): ¡

Turbonilla acuta (Donovan, 1804): ¡

Turbonilla acutissima Monterosato, 1884: b

Turbonilla jeffreysii (Jeffreys, 1848): i

Turbonilla pusilla (Philippi, 1844)

Turbonilla rufa (Philippi, 1836): ¡

Turbonilla sinvosa (Jeffreys, 1884): ¡

Turbonilla striatula (Linnaeus, 1758)

Ebala nitidissima (Montagu, 1803)

Ebala pointeli (de Folin, 1868): i

Ebala sp.: b

Familia ACTEONIDAE

Acteon tornatilis (Linnaeus, 1758): arena y fango, ic

Crenilabrum exilis (Forbes in Jeffreys, 1870): b

Familia DIAPHANIDAE

Diaphana minuta Brown, 1827: b

Familia RETUSIDAE

Retusa semisulcata (Philippi, 1836): ¡

Retusa truncatula (Bruguiére, 1792): i

Cylichnina umbilicata (Montagu, 1803): fango, ic

Familia RINGICULIDAE

Ringicula auriculata (Ménard de la Groye, 1811): fango, c

*Ringicula cfr. leptocheila Brugnone, 1873: b

Familia BULLIDAE

Bulla striata Bruguiére, 1792: arena, ¡

Familia HAMINAEIDAE

Haminaea hydatis (Linnaeus, 1758): i

Haminaea orbignyana (Férussac, 1822): ¡

Weinkauffia turgidula (Forbes, 1844): b

Familia PHILINIDAE

Philine aperta (Linnaeus, 1767): ¡

Philine catena (Montagu, 1803): i

Philine scabra (0. F. Miller, 1776): ib

*Lgona pruinosa (Clark, 1827): fango y detrito coralígeno, cb

Familia SCAPHANDRIDAE

Cylichna cylindracea (Pennant, 1777): fango y detrito coralígeno, cb

Roxania utriculus (Brocchi, 1814): fango, c

Scaphander lignarivs (Linnaeus, 1758): fango, c

Scaphander punctostriatus (Mighels y Adams, 1841): b

Figs. 80,81 ++

Figs. 78, 79 +

Fig. 82. +++

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Familia CAVOLINIIDAE

Cavolinia inflexa (Lesueur, 1813)

Clio cuspidata (Bosc, 1802)

Clio pyramidata Linnaeus, 1767

Creseis acicula Rang, 1828

Styliola subula (Quoy y Gaimard, 1827)

Familia LIMACINIDAE

* Limacina bulimoides (d'Orbigny, 1836): b

Limacina inflata (d'Orbigny, 1836): b

*Limacina retroversa (Fleming, 1822): b

Familia PERACLIDAE

Peracle reticulata (d'Orbigny, 1836): b

Familia ELYSIIDAE

Elysia viridis (Montagu, 1810)

Familia HERMAEIDAE

Stiliger sp.

Familia UMBRACULIDAE

Umbraculum mediterraneum (Lamarck, 1819): fango, c

Familia APLYSIIDAE

Aplysia depilans Gmelin, 1791

Apiysia fasciata Poiret, 1789: arena, i

Aplysia punctata Cuvier, 1803: arena, ¡

Familia TRITONIIDAE

Tritonia hombergi Cuvier, 1803

Familia DOTIDAE

Doto koenneckeri Lemche, 1976

Familia TRIOPHIDAE

Kaloplocamus ramosus (Cantraine, 1835)

Familia POLYCERIDAE

Polycera quadrilineata (0. E. Miller, 1776): piedras, i

Familia DORIDIDAE

Doris verrucosa Linnaeus, 1758: detritos y piedras, ¡

Familia ARCHIDORIDIDAE

Archidoris tuberculata (Cuvier, 1804)

Familia DISCODORIDIDAE

Taringa faba (Ballesteros, Llera y Ortea, 1984): bajo piedras, zona detrítica, ¡

Familia CENTRODORIDIDAE

Jorunna tomentosa (Cuvier, 1804)

Familia DENDRODORIDIDAE

Dendrodoris grandiflora (Rapp, 1827): bajo piedras, zona detrítica, ¡

Doriopsilla areolata Bergh, 1880

Familia ARMINIDAE

Armina maculata Rafinesque, 1814

Familia FLABELLINIDAE

Colmella cavolini (Verany, 1846)

Coryphella pedata (Montagu, 1822)

Familia TERGIPEDIDAE

lergipes tergipes (Forskal, 1775)

Familia EUBRANCHIDAE

Eubranchus exiguus (Alder y Hancock, 1848)

Eubranchus farrani (Alder y Hancock, 1844)

Familia FACELINIDAE

Cratena peregrina Gmelin, 1791

Facelina coronata (Forbes y Goodsir, 1839)

Facelina drummondi (Thompson, 1844)

Facelina sp.

DUDO OO

++ |

+ op

Fig. 83 ++++ p

Figs. 84, 85 + p

Ballesteros (1984)

+ Y

Ros (1975)

++ V

Ros (1975)

Ballesteros (1984)

Ros (1975)

H ov

Ros (1975)

++ V

Ballesteros (1984)

++ V

Asensi (1984)

Ballesteros (1981)

Ballesteros (1978)

Ballesteros (1984)

Asensi (1984)

Ballesteros (1984)

Ballesteros (1978)

Ballesteros (1984)

DS)

Iberus, 15 (1), 1997

Familia FAVORINIDAE

Favorinus branchialis (Rathke, 1806): bajo piedras, zona detrítica, ¡

Favorinus vitreus (Ortea, 1982): bajo piedras, zona detrítica, ¡ + ov

Familia AEOLIDIDAE

Aeolidiella alderi (Cocks, 1852) Ros (1975)

Berghia verrucicornis (0. G. Costa, 1864): bajo piedras, zona detrítica, ¡ H+ oV

Spurilla neapolitana (delle Chiaje, 1841): bajo piedras, zona detrítica, ¡ HH oV

Familia SIPHONARIIDAE

Williamia gussonii (O. 6. Costa, 1829): zonas detríticas, ¡ ++

Familia TRIMUSCULIDAE

Irimusculus mammilaris (Linnaeus, 1758): i ++

Familia ELLOBIIDAE

Auriculinella erosa (Jeffreys, 1829) +

Ovatella firminii (Payraudeau, 1826) ++

Ovatella myosotis (Draparnaud, 1801) ++

Close BIVALVIA

Familia NUCULIDAE

Nucula hanleyi Winckworth, 1930: fango, c +++ v A

Nucula nitidosa Winckworth, 1930: fango y detrito coralígeno, c-b +++ (p), v, Ao

Nucula cfr. nucleus (Linnaeus, 1758): fango, c +++ v A

Nucula sulcata Bronn, 183: fango y detrito coralígeno, cb +++ (p), y Ao

* Ennucula aegeensis (Forbes, 1844): fango y detrito coralígeno, c-b ++ (p),v Ai

Familia NUCULANIDAE

Nuculana commutata (Philippi, 1844): fango y detrito coralígeno, cb + lp), Ai

Nuculana pella (Linnaeus, 1767): fango, c ++ Ao

Familia YOLDIIDAE

* Yoldiella lucida (Lovén, 1846): b Fig.86 ++ pA

*Yoldiella nana (M. Sars, 1865): fango y detrito coralígeno, b Fig.87. ++ (p), Ai

*Yoldiella philippiana (Nyst, 1845): fango y detrito coralígeno, b Figs. 8893 — +++ (p), v Ai

Familia ARCIDAE

Arca noae Linnaeus, 1758: rocas, ¡ H ov

Arca tetragona Poli, 1795 +

Barbatia barbata (Linnaeus, 1758): rocas, i + oV

Barbatia clathrata (Defrance, 1816): b Hop

Anadara diluvii (Lamarck, 1819): fango, c H+ oV

Bathyarca pectunculoides (Scacchi, 1834): b Fig. 94 +++ (p),v

Bathyarca philippiana (Nyst, 1848): b Fig. 95 +++ (p)

Familia NOETIIDAE

Striarca lactea (Linnaeus, 1758): piedras, i-c Sa 0

Familia GLYCYMERIDAE

Glycymeris glycymeris (Linnaeus, 1758): fango, c: Hov

Glycymeris insubrica (Brocchi, 1814): arena, i H+ V

Familia MYTILIDAE

Myrilus galloprovincialis Lamarck, 1819: rocas y espigones, i HH V

Mytilaster minimus (Poli, 1795): rocas, m HH V

*Crenella pellucida (Jeffreys, 1850): detrito coralígeno, b Fig.96 ++ p,v

Gregariella subclavata (Libassi, 1859): rocas, ¡ TV

Gregariella petagnae (Scacchi, 1832): rocas, i HE v

Musculus costulatus (Risso, 1826) +.

Musculus subpictus (Cantraine, 1835): fango, c +H ov

Lithophaga lifhophaga (Linnaeus, 1758): en piedras, ic H ov

Modiolus adriaticus Lamarck, 1819 +

Modiolus barbatus (Linnaeus, 1758): rocas, i +++

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

*Idas cfr. ghisottii Warén y Carrozza, 1990: madera

*Idas simpsoni (Marshall, 1900): esqueletos de peces y cetáceos, ix

Modiolula phaseolina (Philippi, 1844): detrito coralígeno, b

Familia PINNIDAE

Atrina pectinata (Linnaeus, 1758): fango, c

Pinna nobilis Linnaeus, 1758: praderas de Posidonia, ¡

Familia PTERIIDAE

Pteria hirundo (Linnaeus, 1758): gorgonias y restos de redes, c

Familia PECTINIDAE

Palliolum incomparabile (Risso, 1826)

Delectopecten vitreus (Gmelin, 1791): b

Pseudamussium septemradiatum (0. F. Múller, 1776): fango, c

Peplum clavatum (Poli, 1795): fango, c

Karnekampia bruei (Payraudeau, 1826): b

Manupecten pesfelis (Linnaeus, 1758): cb

Chlamys islandica (0. E. Múller, 1776)

Chlamys multistriata (Poli, 1795): fango, ic

Chlamys varia (Linnaeus, 1758): roca y fango, i

Lissopecten hyalinum (Poli, 1795)

Flexopecten flexuosus (Poli, 1795): arena y fango, ic

Flexopecten glaber (Linnaeus, 1758)

Aequipecten opercularis (Linnaeus, 1758): fango, c

Perapecten commutatus (Monterosato, 1875): fango, c

Pecten jacobaeus (Linnaeus, 1758): arena y fango, i-c

Similipecten similis (Loskey, 1811): fango y detrito coralígeno, c-b

*Propeamussium lucidum (Jeffreys in Thompson, 1873): detrito coralígeno, b

Propeamussium fenestratum (Forbes, 1844): detrito coralígeno, b

*(yclopecten hoskynsi (Forbes, 1844): detrito coralígeno, b

Familia SPONDYLIDAE

Spondylus gaederopus Linnaeus, 1758

Familia ANOMIIDAE

Anomia ephippium Linnaeus, 1758: conchas de moluscos

*Heteranomia squamula (Linnaeus, 1758): b

Monia patelliformis (Linnaeus, 1761)

Familia LIMIDAE

Limaria hians (Gmelin, 1791): rocas y espigones, ¡

Limaria inflata Link, 1807: ¡

Notolimea crassa (Forbes, 1844): detrito coralígeno, b

Limatula subauriculata (Montagu, 1808): detrito coralígeno, b

*Limatula cfr. gwyni (Sykes, 1903): detrito coralígeno, b

Familia OSTREIDAE

Ostrea edulis Linnaeus, 1758: rocas, ¡

Fomilia GRYPHAEIDAE

Neopycnodonte cochlear (Poli, 1795): cb

Familia LUCINIDAE

Ctena decussata (0. 6. Costa, 1829): arena, i

Loripes lacteus (Linnaeus, 1758): arena, ¡

Lucinella divaricata (Linnaeus, 1758): arena, ¡

Lucinoma borealis (Linnaeus, 1767): fango y detrito coralígeno. cb

Myrtea spinifera (Montagu, 1803): ¡

Familia THYASIRIDAE

Thyasira (Thyasira) biplicata (Philippi, 1836): b

*Thyasira (Thyasira) obsoleta (Verrill y Bush, 1898): b

*Thyasira (Parathyasira) granulosa (Mont., 1874 ex Jeffreys): b

Fig. 97

Fig. 98

Fig. 99

Figs. 101, 102

p, (1), v

Iberus, 15 (1), 1997

*Thyasira (Parathyasira) subovata (Jeffreys, 1881): detrito coralígeno, b

*Thyasira (Leptaxinus) incrassata (Jeffreys, 1876): b

*Thyasira (Axinulus) croulinensis (Jeffreys, 1847): b

*Thyasira (Axinulus) eumyaria (M. Sars, 1870): b

Thyasira (Mendicula) ferruginea (Locard, 1886): fango y detrito coralígeno, b

Familia CHAMIDAE

Chama gryphoides Linnaeus, 1758: en piedras o Microccosmus, ¡

Pseudochama gryphina (Lamarck, 1819): rocas, i

Familia ERYCINIDAE

Scacchia ovata Philippi, 1844: ¡

Familia KELLIIDAE

Bornia sebetia (0. 6. Costa, 1829): ¡

Kellia suborbicularis (Montagu, 1803): b

Familia LASAEIDAE

Hemilepton nitidum (Turton, 1822): b

Familia MONTACUTIDAE

* Mancikellia pumila (Sowerby, 1846): b

* Montacuta phascolionis Dautzenberg y Fischer, 1925: b

Montacuta substriata (Montagu, 1808): b

Mysella bidentata (Montagu, 1803): i

Mysella obliguata (Chaster, 1897)

Tellimya ferruginosa (Montagu, 1808): b

Epilepton clarkiae (Clark, 1852): b

Epilepton sp.: detrito coralígeno, b

Familia NEOLEPTONIDAE

*Arculus sp.: detrito coralígeno, b

Familia CARDITIDAE

Venericardia antiquata (Linnaeus, 1758): fango, c

Glans aculeata (Poli, 1795): fango y detrito coralígeno, cb

Glans trapezia (Linnaeus, 1758): fango y piedras, c

Familia ASTARTIDAE

Astarte fusca (Poli, 1795): fango, c

Astarte sulcata (da Costa, 1778): fango, c

Goodallia triamgularis (Montagu, 1803): i. b

Goodallia sp.: detrito coralígeno, b

Familia CARDIIDAE

Acanthocardia aculeata (Linnaeus, 1758): arena,

Acanthocardia echinata (Linnaeus, 1758): fango, c

Acanthocardia paucicostata (Sowerby, 1834): fango, ic

Acanthocardia tuberculata (Linnaeus, 1758): arena, ¡

Parvicardium exiguum (Gmelin, 1791): ¡

Parvicardium minimum (Philippi, 1836). fango y detrito coralígeno, cb

Parvicardium ovale (6. B. Sowerby, 1840): i

Parvicardium roseum (Lamarck, 1819): fango, c

Plagiocardium papillosum (Poli, 1795): arena, i

Laevicardium crassum (Gmelin, 1791): fango, c

Loevicardium oblongum (Chemnitz, 1782): fango, cb

Cerastoderma glaucum (Poiret, 1789): arena fangosa, ¡

Familia MACTRIDAE

Mactra glauca (Bor, 1778): arena, ¡

Mactra stultorum (Linnaeus, 1758): arena, ¡

Spisula subtruncata (da Costa, 1778): arena, ¡

Lutraria angustior Philippi, 1844

Lutraria lutraria (Linnaeus, 1758): ic

Lutraria magna (da Costa, 1778): Posidonia y fango, ix

Fig. 100

Fig. 103

Fig. 104

Fig. 105

Fig. 108

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Familia MESODESMATIDAE

Donacilla corea (Poli, 1795)

Ervilia castanea (Montagu, 1803): fango, c

Familia SOLENIDAE

Solen marginatus Pulteney, 1799: fango, i

Familia PHARIDAE

Ensis ensis (Linnaeus, 1758): arena, ¡

Ensis minor (Chenu, 1843): arena, ¡

Pharus legumen (Linnaeus, 1758): arena, ¡

Familia TELLINIDAE

Arcopagia balaustina (Linnaeus, 1758): fango, cb

Arcopagia crassa (Pennant, 1777): fango y rocas, ic

Gastrana fragilis (Linnaeus, 1758)

Macoma cumana (0. 6. Costa, 1829): arena, ¡

Tellina donacina Linnaeus, 1758: arena y fango, ic

Tellina incarnata Linnaeus, 1758: areno, ¡

Tellina nitida Poli, 1791: arena, ¡

Tellina planata Linnaeus, 1758: arena, i

Tellina pulchella Lamarck, 1818: arena y fango, ic

Tellina serrata Brocchi, 1814: fango, c-b

Tellina tenvis da Costa, 1778: arena, ¡

Familia DONACIDAE

Donax semistriatus Poli, 1795: arena, i

Donax trunculus Linnaeus, 1758: arena, ¡

Familia PSAMMOBIIDAE

Gari fervensis (Gmelin, 1791)

Familia SCROBICULARIIDAE

Serobicularia coftardi (Payraudeau, 1826)

Familia SEMELIDAE

Abra longicallus (Scacchi, 1834): detrito coralígeno, b

Familia SOLECURTIDAE

Solecurtus scopula (Turton, 1822): fango, c

Solecurtus strigilatus (Linnaeus, 1758): fango, c

Azorinus chamasolen (da Costa, 1778): fango, c

Familia ARCTICIDAE

Arctica islandica (Linnaeus, 1767)

Familia KELLIELLIDAE

Kelliella abyssicola (Forbes, 1844): fango, cb

Familia TRAPEZIIDAE

Coralliophaga lithophagella (Lamarck, 1819): rocas, c

Familia GLOSSIDAE

Glossus humanus (Linnaeus, 1758): fango, c

Familia VENERIDAE

Callista chione (Linnaeus, 1758): arena, i

Chamelea gallina (Linnaeus, 1758): arena, ¡

Clausinella fasciata (da Costa, 1778): fango,

Dosinia exoleta (Linnaeus, 1758): fango, c

Dosinia lupinus (Linnaeus, 1758): arena y fango, ic

Globivenus effosa (Bivona, 1836)

Gouldia minima (Montagu, 1803): b

lrus irus (Linnaeus, 1758): interior de piedras calcáreas, mi

Paphia aurea (Gmelin, 1791): i

Paphia rhomboides (Pennant, 1777): fango, ic

Pitar mediterranea Tiberi, 1855: b

v Al

Iberus, 15 (1), 1997

Pitar rudis (Poli, 1795): fango, cb

Tapes decussatus (Linnaeus, 1758): arena fangosa, ¡

Timoclea ovata (Pennant, 1777): fango, cb

Venerupis corrugata (Gmelin, 1791): arena, i

Venus casina Linnaeus, 1758: fango, c

Venus nux Gmelin, 1791

Venus verrucosa Linnaeus, 1758: fango, c

Familia PETRICOLIDAE

Petricola lifhophaga (Retzivs, 1786): interior de piedras calcáreas, mii

Mysia undata (Pennant, 1777): fango, ic

Familia CORBULIDAE

Corbula gibba (Olivi, 1792): fango y detrito coralígeno, c-b

Lentidium mediterraneum (0. 6. Costa, 1829): arena, i

Familia GASTROCHAENIDAE

Gastrochaena dubia (Pennant, 1777): interior de piedras calcáreas, ¡

Familia HIATELLIDAE

Hiatella arctica (Linnaeus, 1767): restos de conchas y piedras, ic

Hiatella rugosa (Linnaeus, 1767): restos de conchas, piedras y detrito coralígeno, ib

Panopea norvegica (Spengler, 1793)

Familia PHOLADIDAE

Barnea candida (Linnaeus, 1758): arena con bloques de fango, i

Pholas dactylus Linnaeus, 1758: arena y fango, ¡

Familia TEREDINIDAE

*Bankia carinata (Gray, 1827): madera

*lyrodus pedicellatus (Quatrefages, 1849): madera

Nototeredo norvegica (Spengler, 1792): madera

Familia XYLOPHAGIDAE

Xylophaga dorsalis (Turton, 1819): madera y detrito coralígeno

Familia THRACIDAE

Thracia convexa (Wood, 1815): fango, c

Thracia corbuloides Deshayes, 1830: fango, c

Thracia papyracea (Poli, 1795): arena, i

Thracia pubescens (Pulteney, 1799): fango, c

Familia PANDORIDAE

Pandora inaequivalvis (Linnaeus, 1758): arena, ¡

Pandora pinna (Montagu, 1803): fango y detrito coralígeno, b

Familia POROMYIDAE

Poromya granulata (Nyst y Westendorp, 1839): fango y detrito coralígeno, b

Familia CUSPIDARIIDAE

* Cardiomya striolata (Locard, 1898): fango y detrito coralígeno, b

* Cuspidaria abbreviata (Forbes, 1843): b

Cuspidaria cuspidata (Olivi, 1792): b

Cuspidaria rostrata (Spengler, 1793): b

Clase SCAPHOPODA

Familia DENTALIIDAE

Dentalium agile Sars, 1872

Dentalium inaequicostatum Dautzenberg, 1891: fango y detrito coralígeno, cb

Dentalium panormum Chenu, 1842: b

Dentalium vulgare da Costa, 1778: arena y detrito rocoso, ¡

Fustiaria rubescens (Deshayes, 1825): ¡

Familia SIPHONODENTALIDAE

Pulsellum lofotense (Sars, 1865): fango y detrito coralígeno, b

Codulus jeffreysi (Monterosato, 1875): b

Entalina teftragona (Brocchi, 1814): b

Fig. 109 ++

Fig. 110 +++

Fig. 111 +++

Fig. 108 +++

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figura 2. Concha de Rissoa gemmula (Sitges, 1,7 mm).

Figure 2. Shell of Rissoa gemmula (Sitges, 1.7 mm).

DISCUSIÓN

COMENTARIOS SOBRE ALGUNOS TÁ-

XONES: De la mayoría de especies citadas

para el Garraf existen fotografías y des-

cripciones actualizadas en la literatura,

aunque algunas un poco dispersas. En

este apartado nos hemos limitado a

comentar algunos de los taxones que nos

han parecido de mayor interés, ya sea por

su rareza, por su importancia pesquera-

comercial en el Garraf, por la escasa

documentación bibliográfica existente, o

bien por su importancia biológica en la

zona. De muchas de estas especies repor-

tamos fotografías, la mayoría al M.E.B.,

de la concha o de la protoconcha, según

convenga para su identificación.

Clase GASTROPODA

Familia LEPETIDAE

lothia fulva (O. E. Muller, 1776)

Esta especie atlántica ha sido citada

para el Mediterráneo por TAvIANI (1974),

más concretamente para el Adriático, en

fondos de fango entre 180 y 320 m de

profundidad, aunque como comenta el

propio autor, seguramente se trataba de

una concha semifósil del Wuúrmiense.

CECALUPO Y GIUustTI (1986) citan otro

ejemplar en buenas condiciones de la

Isla de Capraia, entre 400 y 440 m de

profundidad. Nuestro único ejemplar

también parece ser un fósil Wúrmiense,

puesto que se trata de una concha mal

conservada, procedente del detrito de

“El Parrusset” entre 250 y 350 m de pro-

fundidad.

Uber

Familia LEPETELLIDAE

Lepetella cfr. espinosae Dantart y Luque, 1994

No se han encontrado ejemplares

vivos de esta especie, y la diagnosis

sólo es posible estudiando la morfolo-

gía del animal. De todas formas las

otras dos especies con las que podría

confundirse, Lepetella sierrai Dantart y

Luque, 1994 y L. barrajoni Dantart y

Luque, 1994, no se han encontrado en el

Mediterráneo (ver DANTART Y LUQUE,

1994).

Familia ADDISONIDAE

Addisoniía excentrica Tiberi, 1857 (Fig. 3)

MCLEAN (1985) señala que la princi-

pal diferencia entre A. paradoxa Dall,

1882 del Atlántico occidental, y A. excen-

trica (Tiberi, 1857) es el tamaño del

adulto, dando 20, 3 mm de talla máxima

para la primera y 10, 5 mm para la segu-

nada. DANTART Y LUQUE (1994), tras una

detallada discusión, consideran a A.

paradoxa sinónimo posterior de A. excen-

trica, y reportan ejemplares de esta

última de hasta 12 mm. Nosotros hemos

encontrado un ejemplar de A. excentrica

de 20 mm, por lo que ratificamos esta

sinonimia.

Familia FISSURELLIDAE

Emarginula pustula Thiele in Kuester, 1913 (Fig. 4)

Esta especie ha sido considerada

por PIANI (1984) como un “endemismo

del archipiélago Toscano y de la costa

Sarda oriental”, pero el hallazgo de un

ejemplar en el detrito de “El Parrusset”,

amplía su distribución al Mediterráneo

occidental, hecho éste que era de espe-

rar.

Familia SCISSURELLIDAE

Anatoma aspera (Philippi, 1844)

Esta especie ha sido considerada si-

nónina de A. crispata Fleming, 1828, o

como una subespecie de ésta (SCHIRO,

1986), pero presenta una espira más alta,

y parece ser que A. crispata no vive al

sur de Escocia (Gofas, com. pers.), y ade-

más ambas especies presentan diferen-

cias en la rádula (DANTART, com. pers.).

Familia RISSOIDAE

Alvania cimicoides (Forbes, 1844) y Alvania testae (Aradas y Maggiore, 1843)

(ies 790)

Los miles de ejemplares de A. testae

hallados en los contenidos estomacales de

Astropecten irregularis, contrastan con los

tan sólo cuatro de A. címicoides (hallados

todos ellos en la misma estrella), a pesar

de que en sedimentos como el de “El Pa-

rrusset” A. cimicoides es aproximadamente

tres veces más frecuente que A. testae.

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figura 3. Addisonia excentrica (El Parrusset), 20 mm. Figura 4. Emarginula pustula (El Parrusset),

1,22 mm. Figura 5. Protoconcha de Danilia otaviana (El Parrusset). Figura 6. Obtusella macilenta

(Vilanova). Figura 7. Alvania testae (Vilanova), 2,16 mm. Figura 8. Alvania subsoluta (isla de

Capraia, Italia), 2,1 mm.

Figure 3. Addisonia excentrica (El Parrusset), 20 mm. Figure 4. Emarginula pustula (El Parrusset), 1.22

mm. Figure 5. Protoconch ofDanilia otaviana (El Parrusset). Figure 6. Obtusella macilenta. (Vilanova). Fi-

gure 7. Alvania testae (Vilanova), 2.16 mm. Figura 8. Alvania subsoluta (Capraia Island, Italy), 2.1 mm.

Iberus, 15 (1), 1997

Alvania zylensis Gotas y Warén, 1982 (Figs. 13, 14)

Esta especie fue descrita por GOFAS Y

WARÉN (1982) para las costas atlánticas

de Marruecos. AARTSEN, MENKHORST Y

GITTENBERGER (1984) la citan en la Bahía

de Algeciras, y posteriormente, BOGI,

COPPINI Y MARGELLI (1989) la mencionan

por primera vez para el Mediterráneo,

en el Tirreno. En el detrito de “El Parrus-

set” hemos encontrado algunas conchas

que asociamos a esta especie, concreta-

mente a la forma de profundidad des-

crita por BOGI ET AL. (1989), que presenta

una teleoconcha con una escultura débil.

El diámetro máximo de la protoconcha

es de 530 mm. Aportamos, además, la fo-

tografía de un ejemplar de la Isla de Al-

borán (Fig. 14), que aunque presenta una

teleoconcha idéntica a la de los ejempla-

res de Vallcarca, tiene una protoconcha

más pequeña, de 450 mm de diámetro.

Obtusella macilenta (Monterosato, 1880) (Fig. 6)

Es una especie abundante en todos los

fondos fangosos del Garraf, mientras que

sólo hemos hallado unos pocos ejempla-

res de O. intersecta (Wood, 1857). Sinembar-

go, lo normal en fondos similares de otras

regiones es que la proporción sea inversa.

Rissoa gemmula Fischer in de Folin, 1871 (Figs. 2, 20, 21)

Se han encontrado dos conchas en el

litoral de Sitges, más una en Es Caló (For-

mentera, Islas Baleares). Aunque no se ha

estudiado el material tipo, los tres ejem-

plares se corresponden con la descripción

aparecida en FOLIN (1871), que reprodu-

cimos a continuación, y con la figura

representada en NORDSIECK (1972):

“... Long 1*/3 millim. Coquille conique-

allongée, blanche, subdiaphane, ornée de có-

tes longitudinales obsoletes, a peine indi-

quées, et de stries spirales, visibles a la partie

inférieure des tours. Sept tours de spire ven-

trus: les trois premiers translucides, brillants,

globuleux, papilliformes; le quatrieme dilaté,

proportionnellement trés large; les dernieurs

peu dilatés; suture bordée, ornée en dessous

d'une petite zone transverse, brune, inte-

rrompue de blanc; dernier tour orné, a sa

partie moyenne, d'une zonule de meme co-

loration; overture petite, ovale; periostoma

simple. Observation. - On ne pourrait rap-

procher ce Rissoa que du R. dolium (Nyst),

(Nassa Philippi); mais notre espece est plus

élancée, plus petite, á cótes obsoletes, et sa

coloration est spéciale, comme la présence de

la zone suturale et de la zone médiane du

dernier tour”.

En cuanto a la protoconcha, es lisa

de 2!/4 vueltas de espira, con un diáme-

tro máximo de 390 mm. Hemos incluído

en el trabajo un dibujo detallado de la

concha (Fig. 2), aparte de las fotografías

realizadas al M.E.B. (Figs. 20, 21).

Además, hemos fotografiado las espe-

cies de Rissoidae a nuestro juicio más cer-

canas, Pusillina inconspicua (Alder, 1844)

(Figs. 17-19) y P. philippi (Aradas y Mag-

giore, 1844) (Figs. 15, 16), de las que se

diferencia por la coloración y forma.

Familia CALYPTRAEIDAE

Crepidula fornicata (Linnaeus, 1758)

Aunque se ha hallado una única

concha, su presencia en la zona puede

explicarse por la introducción artificial

adherida, al casco de un barco. Ade-

más, en el puerto de Barcelona (a tan

sólo 45 km de distancia) han aparecido

numerosos ejemplares vivos de esta

especie.

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figuras 9-14. Género Alvania, protoconchas. 9, 10: A. testae (Vilanova); 11, 12: A. subsoluta (isla de

Capraia, Italia); 13: A. zylensis (El Parrusset); 14: A. zylensis (isla de Alborán). Escalas, 9, 11-14: 200

um; 10: 100 ym.

Figures 9-14. Genus Alvania, protoconchs. 9, 10. A. testae (Vilanova); 11, 12: A. subsoluta (Capraia

Island, Italy); 13: A. zylensis (El Parrusset); 14: A. zylensis (Alborán Island). Scale bars, 9, 11-14: 200

ym; 10: 100 um.

Iberus, 15 (1), 1997

Figuras 15, 16. Pusillina philippi (cala Montjoy, Roses, Girona). 15: ejemplar de 2,1 mm; 16: pro-

toconcha. Figuras 17-19. Pusillina inconspicua (lossa de Mar, Girona). 17: ejemplar de 1,7 mm; 18,

19: protoconcha. Figuras 20, 21. Ríssoa gemmula (Sitges). 20: ejemplar de 1,7 mm; 21: protocon-

cha. Escalas, 16, 19, 21: 100 um; 18: 200 pm.

Figures 15, 16. Pusillina philippi (cala Montjoy, Roses, Girona). 15: shell of 2.1 mm; 16: protoconch.

Figures 17-19. Pusillina inconspicua (Tossa de Mar, Girona). 17: shell of 1.7 mm; 18, 19: protoconch.

Figures 20, 21. Rissoa gemmula (Sitges). 20: shell of 1.7 mm; 21: protoconch. Scale bars: 16, 19, 21:

100 ym, 18: 200 um.

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figuras 22-28. Protoconchas de Cerithiopsis. 22: C. diadema (Isla de Alborán); 23: C. jeffreysi

(bahía de Almería); 24: C. minima (Sitges); 25: C. nana (Sitges); 26: C. scalaris (bahía de Almería);

27: C. tiara (isla de Alborán); 28: C. tubercularis (La Herradura, Granada). Figura 29: C. nana

(Sitges), concha de 1,8 mm. Figura 30: C. tubercularis (La Herradura, Granada), concha de 2,4

mm. Escalas, 22-28: 200 um; 29, 30: 1 mm.

Figures 22-28. Cerithiopsis protoconchs. 22: C. diadema (Alborán Island); 23: C. jeftreysi (Almería

bay); 24: C. minima (Sitges); 25: C. nana (Sitges); 26: C. scalaris (Almería bay); 27: C. tiara

(Alborán Island); 28: C. tubercularis (La Herradura, Granada). Figure 29: C. nana (Sitges), shell of

1.8 mm. Figure 30: C. tubercularis (La Herradura, Granada), shell of 2.4 mm. Scale bars, 22-28:

200 um; 29, 30: 1 mm.

Iberus, 15 (1), 1997

Familia EULIMIDAE

La familia Eulimidae es una de las

más ricas en aguas profundas, quizás

más que la familia Turridae (BOUCHET Y

WARÉN, 1986). A diferencia de otras

regiones, la zona de estudio es pobre en

especies del género Vitreolina, que prin-

cipalmente viven en aguas infralitorales.

Los pocos ejemplares de este género

encontrados proceden de aguas profun-

das.

Crinophthetros sp. (Figs. 37, 38)

Se han encontrado tres ejemplares

frescos del género Crinophtheiros en con-

tenidos estomacales de Astropecten irre-

gularis, a profundidades superiores a

los 200 m. La especie C. comatulicola

(Graf, 1875) es frecuente en fondos

infralitorales, siempre asociada a Ante-

don mediterranea (Lamarck) (TEMPLADO,

com. pers.), pero a profundidades supe-

riores a 200 m, el crinoideo presente es

Leptometra phalangium (Muller), por lo

que podría tratarse de otra especie dife-

rente perteneciente al género Crinoph-

theiros.

Parvioris ibizenca (Nordsieck, 1968)

La especie Parvioris ibizenca ha sido

referida normalmente en la bibliografía

como P. microstoma (Brusina, 1864), pero

según Gofas (com. pers.) el nombre

correcto sería el primero de éstos,

porque P. microstoma ya está preocu-

pado, con lo que se considera sinoni-

mía.

Vitreolina sp.

La especie tipo del género Vitreolina

es Eulima incurva Bucquoy, Dautzen-

berg y Dollfus, 1883, especie poco clara

(BOUCHET Y WARÉN, 1986). Además,

hay una gran confusión con las especies

adscritas al género Vitreolina (eulímidos

de pequeño tamaño,con forma curva-

da), por lo que hemos preferido men-

cionar estos ejemplares como Vitreolina

sp.

Familia MURICIDAE

Trophon echinatus (Kiener, 1840) y Trophon sp. (Figs. 41-44)

T. echinatus presenta una considerable

variación de formas de la teleoconcha, espe-

cialmente con relación a la profundidad.

Sin embargo, no se ha descrito variabili-

dad en la protoconcha. En el detrito de “El

Parrusset” hemos encontrado dos tipos de

protoconcha que presentan tamaños muy

diferentes; una con un diámetro máximo

de unos 670 mm, que asignamos a T. echi-

natus, y otra con un diámetro máximo regis-

trado entre 770 y 830 mm, que denomi-

namos provisionalmente Trophon sp.

Trophon barvicensis (Johnston, 1825) (Fig. 45)

Según BOUCHET Y WARÉN (1985), no

han visto ningún especimen mediterrá-

neo que pueda asignarse inequívoca-

mente a este taxón. También comentan

que algunas citas de Trophonopsis richardi

(Dautzenberg y Fischer, 1896) para el

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Eigura 31. Talassia daguenetí (El Parrusset), concha juvenil de 0,88 mm. Figura 32. Graphis albida

(El Parrusset), 1,83 mm. Figuras 33, 34. Graphis albida (La Herradura, Granada). 33: concha de 2,4

mm, 34: protoconcha. Figuras 35, 36. Vitreolina perminima (El Parrusset). 35: concha de 2,2 mm;

36: protoconcha. Figuras 37, 38. Crinophtheiros sp. (Vilanova). 37: concha de 4,6 mm; 38: proto-

concha. Figuras 39, 40. Protoconchas de Epitonium. 39: E. clathratulum (Mijas Costa, Málaga); 40:

E. linctum (El Parrusset). Escalas, 34: 100 um; 36: 300 um; 38: 300 um; 39, 40: 200 pm.

Figure 31. Talassia dagueneti (El Parrusset), juvenile shell 00.88 mm. Figure 32. Graphis albida (El

Parrusset), 1.83 mm. Figures 33, 34. Graphis albida (La Herradura, Granada). 33: shell 0f2.4 mm; 34:

protoconch. Figures 35, 36. Vitreolina perminima (El Parrusset). 35: shell of 2.2 mm; 36: protoconch.

Figures 37, 38. Crinophtheiros sp. (Vilanova). 37: shell of 4.6 mm; 38: protoconch. Figures 39, 40. Pro-

toconchs ofEpitonium. 39: E. clathratulum (Mijas Costa, Málaga); 40: E. linctum (El Parrusset). Scale

bars, 34: 100 um; 36: 300 ym; 38: 300 um; 39, 40: 200 um.

Iberus, 15 (1), 1997

Figuras 41, 42. Trophon echinatus (El Parrusset). 41: concha juvenil de 4,75 mm; 42: protoconcha.

Figuras 43, 44. Trophon sp. (El Parrusset). 43: concha juvenil de 2,7 mm; 44: protoconcha. Figura

45. Trophon barviciensis (El Parrusset), 19 mm. Figuras 46, 47. Trophon muricatus. 46: concha juve-

nil de 5,9 mm; 47: protoconcha. Escalas 500 um.

Figures 41, 42. Trophon echinatus (El Parrusset). 41: juvenile shell of 4.75 mm; 42: protoconch.

Figures 43, 44. Trophon sp. (El Parrusset). 43: juvenil shell of 2.7 mm; 44: protoconch. Figure 45.

Trophon barviciensis (El Parrusset), 19 mm. Figures 46, 47. Trophon muricatus. 46: juvenil shell of

5.9 mm; 47: protoconch. Scale bars 500 um.

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figuras 48, 49. Bela brachystoma (Vilanova). 48: ejemplar de 3,6 mm; 49: protoconcha. Figuras 50,

51. Mangelia attenuata (Vilanova). 50: ejemplar de 2,5 mm; 51: protoconcha. Figuras 52, 53.

Mangelia nuperrima (El Parrusset). 52: ejemplar de 7 mm; 53: protoconcha. Figuras 54, 55.

Mangelia serga (El Parrusset). 54: ejemplar de 2,75 mm; 55: protoconcha. Escalas 500 um.

Figures 48, 49. Bela brachystoma (Vilanova). 48: shell of 3.6 mm; 49: protoconch. Figures 50, 51.

Mangelia attenuata (Vilanova). 50: shell 0f2.5 mm; 51: protoconch. Figures 52, 53. Mangelia nupe-

rrima (El Parrusset). 52: shell of 7 mm; 53: protoconch. Figures 54, 55. Mangelia serga (El Parrusset).

54: shell 0f 2.75 mm; 55: protoconch. Scale bars 500 yum.

Iberus, 15 (1), 1997

Mediterráneo (ver DI GERONIMO Y PaA-

NETTA, 1973 y FRANCHINI Y FRILLI, 1970),

podrían corresponder a T. barvicensis.

Nuestro único ejemplar proveniente de

“El Parrusset” parece un ejemplar nor-

mal de T. barvicensis (Warén, com. pers.).

Familia TURRIDAE

Mangelia costata (Donovan, 1804)

AARTSEN ET AL. (1984) diferencian M.

coarctata (Forbes, 1840) de M. costata,

siendo la primera más grande y alar-

gada, con una o dos costillas más y pre-

sentando una coloración uniforme. De

todas maneras, dichos autores no des-

cartan la posibilidad de que M. costata

sea la forma litoral y M. coarctata la de

aguas profundas de una misma especie.

El análisis de más de 300 ejemplares del

Mediterráneo español de la colección de

uno de los autores, desde aguas someras

hasta profundidades de 350 m, y cu-

briendo una área geográfica desde el

Mar de Alborán hasta Cataluña, nos

sugiere que, efectivamente, ambas son

formas de una misma especie, con una

ligera tendencia a aumentar el tamaño y

a atenuar la coloración a medida que

aumenta la profundidad. Además, los

escasos ejemplares de coloración uni-

forme encontrados en aguas profundas

son ejemplares subfósiles. Por lo tanto, y

atendiendo sólo a las características de

la concha, consideramos a M. coarctata

como sinónimo de M. costata.

Mangelia attenuata (Montagu, 1803) (Figs. 50, 51)

Del mismo modo que en el caso

anterior, creemos que M. tenuicostata

(Brugnone, 1868) es sinónimo de M. atte-

nuata. Esta especie, que se encuentra

desde el litoral hasta los 250 m, dismi-

nuye de tamaño al aumentar la profun-

didad, y atenúa la coloración. Además,

las vueltas de espira se van haciendo

más escalonadas y las costillas más mar-

cadas, pero la protoconcha no sufre

variación alguna. Las Figuras 50 y 51

ilustran la forma de aguas profundas.

Mangelia nuperrima (Tiberi, 1855) (Figs. 52, 53)

Esta especie se diferencia de Man-

gelia serga (Dall, 1881) por poseer

vueltas de espira más redondeadas y

una boca más ancha. Se ha encontrado

un ejemplar fresco en el contenido es-

tomacal de Astropecten irregularis, y

cuatro conchas en el detrito de “El Pa-

rrusset”.

Mangelia serga (Dall, 1881) (Figs. 54, 55)

Esta especie se ha citado pocas veces

en el Mediterráneo; una vez para Cerdeña

(CECALUPO, 1984) y otra vez para el Tirreno

Central (SMRIGLIO, MARIOTTINI Y GRAVINA,

1987b). Se ha encontrado un ejemplar con

restos de partes blandas en contenidos

estomacales de Astropecten irregularis y

tres conchas en el detrito de “El Parrusset”.

Taranis moerchi (Malm, 1861) (Figs. 57, 58, 59)

Esta especie, de concha extremada-

mente variable, no es rara en el Medite-

rráneo. Las medidas de la protoconcha

ilustrada son las siguientes: 500 mm de

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figura 56. Microdrillia loprestiana (EL Parrusset), 3,1 mm. Figuras 57-59 Taranis moerchi (El

Parrusset). 57: concha de 4,4 mm, 58, 59: protoconcha. Figuras 60-62. Taranis sp. (El Parrusset).

60: concha de 2,8 mm; 61, 62: protoconcha. Escalas, 58, 61, 62: 500 um; 59: 200 um.

Figure 56. Microdrillia loprestiana (EL Parrusset), 3.1 mm. Figures 57-59. “Taranis moerchi (El

Parrusset). 57: shell of 4.4 mm; 58, 59: protoconch. Figures 60-62. Taranis sp. (El Parrusset). 60: shell

of 2.8 mm; 61, 62: protoconch. Scale bars, 58, 61, 62: 500 ym; 59: 200 ym.

Iberus, 15 (1), 1997

altura y 550 mm de diámetro máximo.

Se han encontrado 18 ejemplares (inclu-

yendo juveniles) en el detrito de “El

Parrusset”.

Taranis sp. (Figs. 60, 61, 62)

Junto a los ejemplares de T. moerchi,

hemos encontrado una concha de

aspecto similar a ésta, pero que difiere

en la protoconcha, que es mucho más

grande (650 mm de altura y 730 mm de

diámetro máximo), y además presenta

una escultura puntiforme alineada en

cordones desde el mismo ápice de la

protoconcha, mientras que en T. moerchi

los cordones de puntos se desordenan

en el ápice. Este ejemplar mide 2, 8 mm

de longitud.

Microdrillia loprestiana (Calcara, 1841) (Fig. 56)

Esta especie es común en contenidos

estomacales de estrellas a partir de los

60-80 metros, y en el detrito de “El

Parrusset”.

Pleurotomella demosia (Dautzenber y Fischer, 1896) (Figs. 66, 67, 68)

Nuestros ejemplares se ajustan a la

descripción y figuras aportadas por BOu-

CHET Y WARÉN (1980), aunque el diá-

metro de la protoconcha es menor. Las

medidas que presenta la protoconcha fo-

tografiada (Figs. 67, 68) son las siguien-

tes: 325 mm de diámetro de la P1 y 580

mm de diámetro de la P2.

Esta especie fue citada por primera

vez para el Mediterráneo por BoGI (1985),

y posteriormente por CECALUPO (1988)

para Cerdeña y por BOGI ET AL. (1989)

para el Tirreno. Hemos encontrado dos

ejemplares frescos en contenidos estoma-

cales de Astropecten irregularis y dos

conchas en el detrito de “El Parrusset”.

Pleurotomella coeloraphe (Dautzenberg y Fischer, 1896) (Figs. 63, 64, 65)

De esta especie, de aspecto más glo-

boso que P. eurybrocha y que P. demosia,

sólamente se conoce el material de la

zona batial de Azores, recolectado en va-

rias estaciones de la expedición MO-

NACO y en una estación de la expedi-

ción PORCUPINE (BOUCHET Y WARÉN,

1980). Presenta una protoconcha similar

a la de P. eurybrocha en cuanto a forma,

pero se diferencia en que la protoconcha

embrionaria es reticulada (como en P. de-

mosia) y no granulada como en P. eury-

brocha. Las medidas que presenta la pro-

toconcha fotografiada son las siguientes:

210 mm de diámetro de la P1 y 460 mm

de diámetro de la P2. El diámetro de la

protoconcha a 100 mm del ápice es de

200 mn, al igual que en P. eurybrocha.

Hemos encontrado 4 ejemplares

juveniles en el detrito de “El Parrusset”.

Se trata por tanto de la primera cita de

esta especie para el Mediterráneo.

Familia TJAERNOEIDAE

Familia monogenérica de Hetero-

branchia descrita por WARÉN (1991) en

base a la anatomía externa del animal.

Previamente, el género Tjaernoeia, des-

crito por WARÉN Y BOUCHET (1988), ha-

bía sido situado provisionalmente en la

familia Pyramidellidae por los mismos

autores.

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figuras 63-65. Pleurotomella coeloraphe (El Parrusset). 63: juvenil de 1,6 mm; 64, 65: protoconcha.

Figuras 66-68. Pleurotomella demosía (El Parrusset). 66: juvenil de 2,1 mm; 67, 68: protoconcha.

Figura 69. Teretía teres (El Parrusset), 4,4 mm. Escalas, 64, 67: 500 pm; 65, 68: 200 ym.

Figures 63-65. Pleurotomella coeloraphe (El Parrusset). 63: juvenile shell of 1.6 mm; 64, 65: proto-

conch. Figures 66-68. Pleurotomella demosia (El Parrusset). 66: juvenile shell of 2.1 mm; 67, 68: pro-

toconch. Figure 69. Teretia teres (El Parrusset), 4.4 mm. Scale bars, 64, 67: 500 pm; 65, 68: 200 ym.

Iberus, 15 (1), 1997

Figuras 70-77. Familia Pyramidellidae. 70: Chrysallida brattstroemi (El Parrusset), 1,1 mm. 71: Chry-

sallida dollfusi (LEscala, Girona), 2,8 mm. 72: Eulimella ataktos (Vilanova), 2,7 mm. 73: Eulimella

ventricosa (Isla de Alborán), 2,36 mm. 74: Enlimella unifasciata (Blanes, Girona), 5,5 mm. 75: Odos-

tomia afzelii (Vilanova), 1,5 mm. 76: Odostomia hansgei (Vilanova), 1,6 mm. 77: Turbonilla acutis-

sima (Mijas, Málaga), 4,4 mm. Escalas, 70: 200 pm; 71, 72, 73, 75, 76: 500 um; 74, 77: 1 mm.

Figures 70-77. Family Pyramidellidae. 70: Chrysallida brattstroemi (El Parrusset), 1.1 mm. 71: Chry-

sallida dollfusi (L'Escala, Girona), 2.8 mm. 72: Eulimella ataktos (Vilanova), 2.7 mm. 73: Eulimella

ventricosa (Alborán Island), 2.36 mm. 74: Eulimella unifasciata (Blanes, Girona), 5.5 mm. 75: Odos-

tomia afzelii (Vilanova), 1.5 mm. 76: Odostomia hansgei (Vilanova), 1.6 mm. 77: Turbonilla acutis-

sima (Mijas, Málaga), 4.4 mm. Scale bars, 70: 200 um; 71, 72, 73, 75, 76: 500 um; 74, 77: 1 mm.

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figuras 78, 79. Ringicula cfr. leptocheila (El Parrusset). 78: concha de 2,37 mm; 79: protoconcha.

Figuras 80, 81. Ringicula auriculata (Vilanova). 80: concha de 3,75 mm; 81: protoconcha. Figura

82. Cylichnina umbilicata (Vilanova), 1,3 mm. Figura 83: Limacina retroversa (El Parrusset), 2,7

mm. Figuras 84, 85. Peracle reticulata (El Parrusset). 84: concha de 2 mm; 85: protoconcha. Escalas,

79, 85: 200 um; 81: 500 um.

Figures 78, 79. Ringicula cfr. leptocheila (El Parrusset). 78: shell 02.37 mm; 79: protoconch. Figures 80,

81. Ringicula auriculata (Vilanova). 80: shell of 3.75 mm, 81: protoconch. Figure 82. Cylichnina umbili-

cata (Vilanova), 1.3 mm. Figure 83: Limacina retroversa (El Parrusset), 2.7 mm. Figures 84, 85. Peracle

reticulata (El Parrusset). 84: shell of 2 mm; 85: protoconch. Scale bars, 79, 85: 200 ym; 81: 500 ym.

Iberus, 15 (1), 1997

Tjaernoeia exquisita (Jeffreys, 1883)

Especie poco frecuente en el Medite-

rráneo, en donde WARÉN (1991) la

señala entre 25 y 200 m. Nosotros

hemos encontrado ejemplares a más

profundidad en el detrito de “El Parrus-

set”.

Familia ARCHITECTONICIDAE

Basisulcata lepida (Bayer, 1942)

Sólo se han encontrado dos ejempla-

res vivos en “La Mar de Nit”, a unos 40 m

de profundidad, en una zona detrítica con

grandes colonias de Bolinus brandaris (L.,

1758), cuyas conchas estaban recubiertas

por la anémona Calliactis parasitica (Couch).

Familia PYRAMIDELLIDAE (Figs. 70-77)

En el trabajo sobre Pyramidellidae

del Mediterráneo español de PEÑAS,

TEMPLADO Y MARTÍNEZ (1996) se aporta

una gran cantidad de datos sobre la pre-

sencia de esta familia en fondos del

Garraf. Asimismo, se citaba por primera

vez para el Mediterráneo las siguientes

especies: Chrysallida brattstroemi Warén,

1991, Eulimella ataktos Warén, 1991,

Odostomia afzelii (Warén, 1991), y Odosto-

mia hansgei (Warén, 1991), las cuatro

comunes en esta zona, y ninguna

hallada hasta ahora en ninguna otra

localidad del Mediterráneo español.

Además, en el mismo trabajo, numero-

sas especies son citadas por primera vez

para el Mediterráneo español, también

procedentes del Garraf.

Las especies que no habían sido cita-

das para el Garraf en el trabajo anterior de

PEÑAS ET AL. (1996), y que hemos hallado

posteriormente, son: Chrysallida dollfusi

(Kobelt, 1903), Eulimella unifasciata (Forbes,

1844), Eulimella ventricosa (Forbes, 1844),

y Turbonilla acutissima Monterosato, 1884.

El hallazgo en “El Parrusset” de una

concha subfósil de Chrysallida pellucida

(Dillwyn, 1817), no contradice la hipóte-

sis de PEÑAS ET AL., (1996), que comen-

tan que, en la actualidad, esta especie

atlántica no penetra en el Mediterráneo

más allá del mar de Alborán.

Familia SCAPHANDRIDAE

Scaphander punctostriatus (Mighels y Adams, 1841)

Esta especie ha sido citada raras

veces en el Mediterráneo (MONTERO-

SATO, 1880; Ros, 1975; BOUCHET Y

TAvIANI, 1989), y ha sido tradicional-

mente considerada una especie de aguas

profundas del Atlántico. Se han encon-

trado 6 conchas en el detrito de “El

Parrusset”.

Familia DISCODORIDIDAE

Taringa faba (Ballesteros, Llera y Ortea, 1984)

Esta especie se encuentra en la

playa del búnker de Cubelles, bajo

piedras a menos de 50 centímetros de

profundidad. Ésta es la localidad tipo

de la especie, descrita por BALLESTE-

ROS, LLERA Y ORTEA (1984), donde se

puede encontrar con relativa frecuen-

cia.

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figura 86. Yoldiella lucida, 4,25 mm. Figura 87. Yoldiella nana, 2,9 mm. Figuras 88-93. Yoldiella

philippiana. 88, 89: juveniles de 2,9 y 2,3 mm; 90: adulto de 4,3 mm; 91: juvenil, detalle de los

dientes de la charnela; 92: charnela; 93: protoconcha. Todos las especies de El Parrusset, contenido

estomacal de Astropeceten, 250-350 m. Escalas; 92: 200 pm; 93: 100 pm.

Figure 86. Yoldiella lucida, 4.25 mm. Figure 87. Yoldiella nana, 2.9 mm. Figures 88-93. Yoldiella phi-

lippiana. 88, 89: juvenile shells of 2.9 and 2.3 mm; 90: adult shell of 4. 3 mm; 91: juvenil shell, hinge

lateral view; 92: hinge; 93: protoconch. All the especies from El Parrusset, gut content of Astropecten,

250-350 m. Scale bars, 92: 200 ym; 93: 100 ym.

Iberus, 15 (1), 1997

Familia FAVORINIDAE

Favorinus vitreus (Ortea, 1982)

Esta especie fue descrita por ORTEA

(1982) en Tenerife (Islas Canarias), a par-

tir de dos ejemplares, y únicamente se

ha citado 1 ejemplar para el Mediterrá-

neo, en al Cabo de Palos (Murcia) en ri-

zomas de Posidonia a 5 m de profundi-

dad (TEMPLADO, 1982). Se ha recolec-

tado un ejemplar de esta especie en Ju-

nio de 1993 en Cubelles, debajo de una

piedra a unos 20-30 cm de profundidad.

Clase BIVALVIA

Familia NUCULIDAE

Nucula cfr. nucleus (Linnaeus, 1758)

Gofas (com. pers.) opina que su pre-

sencia en el Mediterráneo es dudosa,

siendo N. hanleyi Winckworth, 1930, la

especie más normal, que presenta un

periostraco más brillante, con estrías

radiales oscuras, y más alargada ante-

riormente. SALAS (1996) ilustra un ejem-

plar de N. nucleus del Mediterráneo.

Hemos encontrado ejemplares juveniles

con una protoconcha que coincide con la

descripción de GOFAS Y SALAS (1996), y

ejemplares adultos que, aunque han

perdido parte del periostraco, asigna-

mos a N. nucleus.

Familia YOLDIIDAE

La familia Yoldiidae es la principal

familia de Nuculanoidea en el Medite-

rráneo, y en el Garraf sólo hemos ha-

llado el género Yoldiella. Las otras fami-

lias de Nuculanoidea no han sido reco-

lectadas en este estudio por presentar

una distribución batimétrica de mayor

profundidad. Aunque tenemos constan-

cia de la existencia de especies de otras

familias en zonas más profundas frente

a la costa del Garraf (Dantart, com. pers.).

Hemos tratado de recopilar aquí las

citas modernas de las especies de la sub-

familia Yoldiellinae en el Mediterráneo

debido a que la informacion sobre el

género es muy dispersa. Sin embargo,

no se han tenido en cuenta descripcio-

nes originales, citas antiguas ni sinoni-

mias, que se pueden encontrar en los

artículos mencionados en esta sección.

De todas las citas recopiladas para el

Mediterráneo, reconocemos sólo 5 espe-

cies. Se han dado entre corchetes los

nombres utilizados incorrectamente, o

los que han sido considerados como

sinonimias.

Yoldiella lucida (Lovén, 1846) (Fig. 86)

Di Geronimo y Panetta (1973): Golfo de Taranto, 940-1000 m

Warén (1989): distribucion Mediterránea entre 100 y 1000 m de profundidad

Es la especie de Yoldiella menos fre-

cuente de las halladas en Garraf, habién-

dose encontrado sólo tres conchas, dos

en contenidos estomacales de estrellas y

una en el sedimento, todas del detrito

de “El Parrusset”, entre 250 y 350 m de

profundidad, por lo que se deduce que

vive en la zona.

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figura 94. Bathyarca pectunculoides (El Parrusset), 1,3 mm. Figura 95. Bathyarca philippiana (El

Parrusset), 1,3 y 1,6 mm. Figura 96. Crenella pellucida (El Parrusset), 710 um. Figura 97. Modiolula

phaseolina (El Parrusset), 1,96 mm.

Figure 94. Bathyarca pectunculoides (El Parrusset), 1.3 mm. Figure 95. Bathyarca philippiana (El

Parrusset), 1.3 and 1.6 mm. Figure 96. Crenella pellucida (El Parrusset), 710 pm. Figure 97.

Modiolula phaseolina (El Parrusset), 1.96 mm.

Iberus, 15 (1), 1997

Yoldiella nana (M. Sars, 1865) (Fig. 87)

Cecalupo y Giusti (1986): Isla de Capraia, 400-440 m [Portlandia frigida (Torell, 1859)]

Bogi et al. (1989): Capo Corso, 150 m [Portlandia frigida (Torell, 1859)]

Warén (1989): Banyuls, 650-770 m

Se han encontrado varias conchas en

contenidos estomacales de estrellas

entre 100 y 350 m de profundidad, y

otras en el detrito de “El Parrusset”, por

lo que se deduce que también vive en la

zona.

Yoldiella philippiana (Nyst, 1845) (Figs. 88-93)

Di Geronimo y Panetta (1973): Golfo de Taranto, 350-1000 m [Yoldiella tenuis (Philippi)]

Di Geronimo (1974): Jónico [Yoldiella tenuis (Philippi)]

Cecalupo y Giusti (1986): varios ejemplares vivos, Isla de Capraia, 400-440 m

Warén (1989): Mediterráneo, 100-300 m

Bonfitto y Sabelli (1995): Cerdeña, 245-1707 m

Es la especie más abundante, apare-

ciendo tanto en sedimentos como en

contenidos intestinales de Astropecten,

desde los 60 hasta los 350 m de profun-

didad. Se han encontrado numerosos

juveniles vivos.

Las otras dos especies mediterráneas

de este género, Yoldiella messanensis

(Seguenza, MS, Jeffreys, 1870) y Yoldiella

seguenzae Bonfitto y Sabelli, 1995, no se

han encontrado en la zona de estudio.

La primera ha sido citada en el Medite-

rráneo en general (150-3000 m) y en el

mar de Alborán (60-1235 m) por

TERRENI (1980) CECALUPO Y GIUST1

(1986), WARÉN (1978, 1989), ALLEN Y

HANNAH (1989) y SALas (1996). La se-

gunda sólo se conoce de Cerdeña -(245-

1707 m) y mar de Alborán, entre 480 y

1005 m (BONFITTO Y SABELLL 1995;

SALAS 1996).

Familia MYTILIDAE

El género Idas en el Mediterráneo

está representado por tres especies (Wa-

RÉN, 1991), asociadas siempre a restos

de esqueletos de cetáceos. Además de

este hábitat, 1. argenteus Jeffreys, 1876 e 1.

ghisotti Warén y Carozza, 1990 también

pueden encontrarse en restos de troncos

(WARÉN, 1991, 1993). La utilización del

género Idas Jeffreys, 1878, en vez de Ida-

sola Iredale, 1915, contrariamente a lo

propuesto por DELL (1987), se basa en lo

expuesto por WARÉN (1991).

Idas cfr ghisottii Warén y Carrozza, 1990

En un tronco de madera arrojado a

la playa tras una tormenta, se ha

encontrado un ejemplar que asignamos

a esta especie, junto con varios ejem-

plares de Teredinidae. Esta especie sólo

se ha encontrado asociada a restos de

madera en el Mediterráneo por

CARROZZA (1984), que la identificó

erróneamente como Myrina modiolaefor-

mis Sturany, 1896. Fue corregido poste-

riormente por WARÉN Y CARROZZA

(1990), que la describieron como nueva

especie. De todas formas, y debido a la

pérdida del único ejemplar hallado, la

asignación de este especimen podría

ser errónea, aunque estamos seguros

de que no se trata de /. simpsoni (Mars-

hall, 1900).

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figura 98. Thyasira (1) biplicata (El Parrusset), 4,9 mm. Figura 99. Thyasira (1) obsoleta (El Parrusset),

1,46 y 1,87 mm. Figura 100. Thyasira (P) subovata (El Parrusset), 1 y 1,16 mm. Figuras 101, 102.

Thyasira (Parathyasira) granulosa. 101: ejemplar de 5.4 mm. 102: detalle de la granulación de la concha.

Escala 400 um.

Figure 98. Thyasira (T.) biplicata (El Parrusset), 4.9 mm. Figure 99. Thyasira (T.) obsoleta (El Parrusset),

1.46 and 1.87 mm. Figure 100. Thyasira (P.) subovata (El Parrusset), 1 and 1.16 mm. Figures 101, 102.

Thyasira (Parathyasira) granulosa. 101: shell of5.4 mm. 102: shell granulation detail. Scale bar 400 ym.

Iberus, 15 (1), 1997

Idas simpsoni (Marshall, 1900)

Se trata de otro Mytilidae interesante,

que vive en restos orgánicos de esquele-

tos de cetáceos o peces, aunque mucho

más común que el anterior y de distribu-

ción más amplia. Se han encontrado va-

rios ejemplares asociados a esqueletos.

Modiolula phaseolina (Philippi, 1844) (Fig. 97)

Se han encontrado unos pocos ejem-

plares vivos en el detrito de “El Parrus-

set”, aunque esta especie destaca por la

gran cantidad de conchas Wúrmienses

de gran tamano (más de 20 mm) encon-

tradas en esta localidad.

Familia PECTINIDAE

Hemos incluido en esta familia las

especies clasificadas por algunos auto-

res como Propeamussiidae (o Amussii-

dae). De todas formas, esta clasificación

es provisional hasta que se realice un es-

tudio filogenético que resuelva la cues-

tión. Hemos seguido la sistemática utili-

zada por WAGNER (1991), excepto en la

separación de Pectinidae y Amussiidae.

Para algunas especies de “Amussiidae”,

hemos seguido a SMRIGLIO Y MARIOT-

TINI (1990).

Pseudamussium septemradiatum (O. E. Muller, 1776)

Se han encontrado ejemplares vivos

de esta especie en fondos de fango entre

150 y 200 m de profundidad, mientras

que VinYas (1981) la consideraba como

una especie de aguas frías, que supues-

tamente se había extinguido del Medite-

rráneo, aunque abundante en los sedi-

mentos Wúrmienses (MArs, 1958; MAR-

TINELL Y JULIA-BRUGUES, 1973; VINYAS,

1981; DOMENECH Y MARTINELL, 1982).

SALAS (1996) la cita viva para el mar de

Alborán.

Flexopecten glaber (Linnaeus, 1758)

Gofas (com. pers.) ha comentado que se

trata de una especie más bien lagunar,

rara en el Mediterráneo occidental, y que

algunos juveniles grandes de E. flexuosus,

que no presentan aún la flexuosidad típica

de la especie, pueden ser confundidos con

F. glaber. Hemos encontrado una única

valva de esta especie, de 33 mm de altura

con 10 costillas principales, claramente

diferente de FE. flexuosus, cuyos mayores

ejemplares encontrados en la zona miden

28 mm y presenta 5 costillas principales.

Familia LIMIDAE

Limatula cfr. gwyni (Sykes, 1903)

Se ha encontrado una única valva

fresca procedente del detrito de “El

Parrusset”. El ejemplar se destruyó

durante el proceso de montaje para el

M.E.B. El color era blanco y presentaba

una forma y tamaño similar a L.

subovata, aunque totalmente lisa y sin

restos de costillas radiales.

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figura 103. Thyasira (Leptaxinus) incrassata (El Parrusset), 1,66 mm. Figura 104. Thyasira

(Axinulus) croulinensis (El Parrusset), 1,26 mm. Figura 105. Thyasira (A.) eumyaria (El Parrusset),

2,9 mm. Figura 106. 7hyasira (Mendicula) ferruginea (El Parrusset), 1,46 mm. Figura 107. Arculus

sp. (El Parrusset), 1,02 mm y 1,16 mm.

Figure 103. Thyasira (Leptaxinus) incrassata (El Parrusset) 1.66 mm. Figure 104. Thyasira

(Axinulus) croulinensis (El Parrusset), 1.26 mm. Figure 105. Thyasira (A.) eumyaria (El Parrusset),

2.9 mm. Figure 106. Thyasira (Mendicula) ferruginea (El Parrusset), 1.46 mm. Figure 107. Arculus

sp. (El Parrusset), 1.02 mm and 1.16 mm.

Iberus, 15 (1), 1997

Familia LUCINIDAE

Lucinoma borealis (Linnaeus, 1767)

Se han encontrado ejemplares vivos de

gran tamaño (hasta 47,9 mm, un ejemplar

de la coleccion de J. L. Ferrer), en fondos

de fango entre 60 y 80 m de profundidad.

Familia THYASIRIDAE (Figs. 98-106)

Los tiasíridos son la principal familia

de bivalvos que habita en el talud.

Muchas especies viven desde el circalito-

ral hasta profundidades abisales. Las

especies atlánticas han sido reciente-

mente revisadas a nivel morfológico por

PAYNE Y ALLEN (1991), pero son pocos los

trabajos sobre este grupo en el Mediterrá-

neo. La descripción de una nueva especie

mediterránea por CARROZZA (1981), así

como las redescripciones de algunas

especies de Monterosato publicadas por

GAGLINI (1991) son las obras modernas

más importantes para este mar. Además,

existen algunas citas dispersas (p. e. DI

GERONIMO Y PANETTA, 1973; TERRENI,

1980; CARROZZA, 1984; CIANFANELLI Y

TALENTI, 1987; CECALUPO Y GIusTI, 1989),

pero no hay ningún trabajo taxonómico

de revisión importante. Tampoco preten-

demos aquí realizar una revisión taxonó-

mica del grupo, pero hemos querido foto-

grafiar las 8 especies aparecidas en el

detrito de “El Parrusset” (Figs. 98-106),

constituyendo 6 de ellas la primera cita

para el Mediterráneo español.

Thyasira biplicata (Philippi, 1836) (Fig. 98)

Esta especie, fue descrita por PHI-

LIPPI (1836) para zonas abisales del Mar

Mediterráneo, pero ha sido referida en

la literatura como T. flexuosa (Montagu,

1803), que es una especie litoral atlántica

con la ondulación posterior más suave

(Gofas, com. pers.), de presencia dudosa

en el Mediterráneo.

Familia MONTACUTIDAE

Epilepton clarkiae (Clark, 1852)

Todos los ejemplares se han encon-

trado en una comunidad dominada por

Turritella communis Risso, 1826, a unos

60 m de profundidad.

Epilepton sp.

Este pequeño Epilepton, aparecido en

el detrito coralígeno de “El Parrusset”

está en este momento en proceso de des-

cripción.

Familia NEOLEPTONIDAE

Arculus sp. (Fig. 107)

Hemos identificado provisionalmen-

te estos ejemplares, hallados también en

“El Parrusset”, dentro del género Arcu-

lus.

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Figura 108. Xylophaga dorsalis (Vilanova): exterior valva derecha, 3,4 mm; interior valva izquierda,

1,6 mm. Figura 109. paleta de Bankia carinata (Vilanova), 4,7 mm. Figura 110. Lyrodus pedicellatus

(Vilanova): exterior valva izquierda, 1,9 mm; interior valva derecha, 2,16 mm; paleta de 2,5 mm. Fi-

gura 111. Nototeredo norvegica (Vilanova): exterior valva izquierda, 1, 38 mm; interior valva derecha,

1,5 mm; paleta de 1,7 mm.

Figures 108. Xylophaga dorsalis (Vilanova): outside of right valve, 3.4 mm; inside of lefi valve, 1.6 mm.

Figure 109. pallet ofBankia carinata (Vilanova), 4.7 mm. Figures 110. Lyrodus pedicellatus (Vilanova):

outside of lefi valve, 1.9 mm; inside of right valve, 2.16 mm; pallet of 2.5 mm. Figure 111. Nototeredo

norvegica (Vilanova): outside of lefi valve, 1.38 mm; inside of right valve, 1.5 mm), pallet of 1.7 mm.

Mars SANS

Familia ASTARTIDAE

Goodallia sp.

En el detrito de “El Parrusset” han

aparecido ejemplares pertenecientes

al género Goodallia, similares al taxon

Los MOLUSCOS DE “EL PARRUSSET”:

Como se comentaba anteriormente, “El

Parrusset” es una biocenosis de coral

blanco (PÉRES Y PICARD, 1964) con fan-

go, cuyas especies predominantes son

los madreporarios Dendrophyllia corni-

gera (Lamarck) y Caryophyllia sp., aso-

ciada a una tanatocenosis del Wur-

miense, con la presencia de algunas es-

pecies subfósiles que actualmente están

extintas en el Mediterráneo. En esta lo-

calidad se han obtenido numerosas es-

pecies de moluscos de profundidad,

siendo Trophon barvicensis y Pleurotomella

coeloraphe la primera vez que se citan

para el Mediterráneo, y otras la primera

vez que se citan para el litoral Medite-

rráneo español.

Una comunidad muy similar a la de

“El Parrusset” ha sido descrita por SMRI-

GLIO, MARIOTTINI Y GRAVINA (1987a;

1987b) para el mar Tirreno Central,

donde estudiaron una biocenosis de co-

ral blanco con detrito fangoso situada

entre 400 y 600 m de profundidad, en la

que la especie de madreporario predo-

minante es Dendrophyllia cornigera (La-

marck). Aunque se trate de dos comuni-

dades muy parecidas desde un punto de

vista biológico y paleontológico, las es-

pecies de moluscos que se encuentran

en la localidad del Tirreno Central (Sm-

RIGLIO ET AL., 1987a; 1987b, 1988a; 1988b;

1989; 1993) son características de aguas

más profundas. Sin embargo, algunas

de las especies son coincidentes en am-

bas biocenosis, como por ejemplo Micro-

drillia lopestriana (Calcara, 1841), Mange-

lia serga (Dall, 1881) y Teretia teres (For-

bes, 1844), entre los túrridos.

En cuanto a los bivalvos actuales en-

contrados en esta biocenosis, los grupos

mayoritarios son Nuculidae, Yoldielli-

dae, Bathyarca, Astarte, Veneridae, Thya-

siridae, Pectinidae y Xylophaga, siendo

88.

G. macandrewi Smith, 1881, que posi-

blemente se trate de una nueva espe-

cie.

Kelliella abyssicola (Forbes, 1844) la espe-

cie viva más abundante, tanto en detri-

tos como en contenidos estomacales de

estrellas. ALLEN (1979) en un trabajo so-

bre bivalvos abisales Atlánticos, co-

menta que el 95% de especies de bival-

vos de sustratos blandos son Protobran-

chia, Septibranchia y Thyasiridae,

mientras que para sustratos duros, pre-

dominan las especies con biso, como

Bathyarca, Limopsis y Dacrydium. Ade-

más, encuentra especies pequeñas, ge-

neralmente menores de 5 mm. En este

trabajo se han encontrado prácticamente

los mismos grupos, aunque representa-

dos por especies o géneros de menor

profundidad y por norma general, de ta-

llas mayores.

El primer yacimiento Wúrmiense

conocido para el litoral catalán fue el

Cap de Creus (Girona), descrito por

PRUVOT Y ROBERT (1897), posteriormente

estudiado por Mars (1958). Otro yaci-

miento situado en el Cap de Begur

(Girona) ha sido descrito por MARTINELL

Y JULIA-BRUGUES (1973). Trabajos poste-

riores sobre estos yacimientos Wur-

mienses de la costa de Girona son, entre

otros, los de VINYAS (1981) y DOMENECH

Y MARTINELL (1982). En cuanto a los

fondos blandos del litoral catalán, la

primera fauna malacológica Wúrmiense

fue descrita por MARTINELL, DOMENECH

Y DE VILLALTA (1986) para el delta del

Ebro (Tarragona).

La tanatocenosis Wurmiense que

aquí se describe es el origen de algunas

de las especies subfósiles que se encuen-

tran en el área. Éste es el caso de los Gas-

terópodos: lothia fulva (O. E. Múller, 1776),

Calliostoma zizyphinum (Linnaeus, 1758),

Danilia otaviana (Cantraine, 1835), Capulus

ungaricus (Linnaeus, 1758), Trivia multili-

rata (Sowerby, 1870), Erato voluta (Mon-

tagu, 1803), Ranella olearia (Linnaeus,

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

1758), Buccinum undatum Linnaeus, 1758,

Neptunea contraria (Linnaeus, 1771), y

Murexul aradasi (Poirier, 1883 ex Montero-

sato ms.), que se encuentran sólo como

especies subfósiles en el área de estudio.

Entre los Bivalvos, destacan los grandes

ejemplares de Modiolula phaseolina (Phi-

lippi, 1844), Chlamys islandica (O. F. Mú-

ller, 1776), Pseudamussium septemradiatum

(O. F. Múller, 1776), Arctica islandica (Lin-

naeus, 1767), Glossus humanus (Linnaeus,

1758), Globivenus effosa (Bivona, 1836),

Gouldia minima (Montagu, 1803), Pitar me-

diterranea Tiberi, 1855, Venus casina Linna-

eus, 1758 y Panopea norvegica (Spengler,

1793). Al contrario que los Gasterópodos,

muchas de estas especies también se en-

cuentran vivas a menos profundidad, o

incluso en el mismo sedimento, como es

el caso de M. phaseolina. Las especies de

Bivalvos que se encuentran en forma

subfósil exclusivamente son C. islandica,

A. islandica, y P. norvegica, especies de

aguas frías que se han extinguido del Me-

diterráneo (VINYAS, 1981), y G. effosa, que

vive en zonas profundas del Mediterrá-

neo (Mar de Alborán).

La malacofauna del sedimento Wir-

miense de fondos blandos que aquí se-

ñalamos, tiene una composición similar

a la de los yacimientos de la costa de Gi-

rona descritos por los autores menciona-

dos más arriba. De todas formas, la ma-

yoría de los trabajos anteriores están

únicamente basados en especies grandes

debido a la metodología de muestreo

empleada (Martinell, com. pers.). En los

moluscos de “El Parrusset” destaca la

ausencia de Buccinum humphreysianum

Bennet, 1824 (aunque se han encontrado

conchas en otras zonas del Garraf), y de

Modiolus modiolus (Linnaeus, 1758), pre-

sentes en casi todos los otros yacimien-

tos de la costa catalana, así como la au-

sencia de Colus islandicus (Gmelin, 1791),

abundante en los yacimientos de la

costa de Girona.

CONTENIDOS ESTOMACALES DE AS-

TROPECTEN: Los contenidos estomacales

de Equinodermos y otros depredadores

han sido investigados por diversos mala-

cólogos, como método sencillo para reco-

lectar numerosas especies, principalmente

micromoluscos de profundidad. Trabajos

científicos de este tipo han sido realizados

en el Mediterráneo español, concretamente

en las islas Baleares (GASULL Y CUERDA,

1974) y en la bahía de Almería (SIERRA,

GARCÍA Y LLORIS, 1978), así como en la

costa atlántica española, en la Ría de Ares

(Galicia) (CRISTOBO-RODRÍGUEZ, TRON-

COSO, URGORRI-CARRASCO Y RíOs-LÓPEZ,

1988), entre otros.

En este trabajo, el número de espe-

cies de Moluscos encontradas en los

contenidos estomacales de Astropecten

spp. es de 118: 1 Poliplacóforo, 90 Gaste-

rópodos, 25 Bivalvos y 2 Escafópodos).

Si bien no se ha recopilado información

cuantitativa en los muestreos, sí se tie-

nen datos cualitativos, de los que se ha

podido extraer la siguiente información:

- En el Garraf A. aranciacus es más

común entre los 40 y los 80 m de pro-

fundidad en fondos fangosos, mientras

que A. irregularis es abundante en un

rango batimétrico mucho más amplio y

en todo tipo de fondos. Casi todas las

especies halladas en A. aranciacus se

encuentran en A. irregularis, aunque en

esta última se trata de individuos más

pequeños o juveniles.

- El Gasterópodo predominante en

todas las profundidades estudiadas fue

Alvania testae (Aradas y Maggiore, 1843),

representando aproximadamente el 50%

de individuos encontrados.

- El Bivalvo predominante hasta los

80 m fue Timoclea ovata (Pennant, 1777),

y a partir de esta profundidad predo-

minó Kelliella abyssicola (Forbes, 1844).

- En “El Parrusset”, existe una rela-

ción directa entre el número de ejempla-

res vivos de Nuculidae, Nuculanidae y

Yoldiidae encontrados en los contenidos

estomacales y los hallados en el sedi-

mento, pero ésto no ocurre en el resto de

grupos estudiados.

- En general, se observó una reduc-

ción del tamaño de muchas especies (so-

bre todo apreciable en grupos abundan-

tes, como Pyramidellidae) a medida que

aumentó la profundidad. Este hecho ha

sido comentado por ALLEN (1979), que

encuentra una disminución de las tasas

de crecimiento relacionada con el incre-

mento de la profundidad.

Iberus, 15 (1), 1997

NUEVAS CITAS PARA EL MEDITERRÁ-

NEO ESPAÑOL: Como se señaló anterior-

mente, en la lista de especies se identifi-

can con un asterisco aquellas que, de

acuerdo con la bibliografía, se citan por

primera vez en el litoral Mediterráneo

español, y con dos asteriscos las que se

citan por primera vez para el Meditérra-

neo en general. En este apartado no se

han tenido en cuenta las citas del estre-

cho de Gibraltar como pertenecientes al

Mediterráneo.

La publicación del catálogo provisio-

nal no crítico de los Bivalvos del Medi-

terráneo español de BONNIN Y RODRÍ-

GUEZ-BABÍO (1990), basado en la biblio-

grafía, nos ha facilitado el trabajo de

recopilación de citas. En cuanto a los

Gasterópodos, no existe ningún catá-

logo actualizado, exceptuando algunas

monografías de varios grupos, como

Cocculiniformia (DANTART Y LUQUE,

1994); Pyramidellidae (PEÑAS ET AL,,

1996); Opistobranchia (CERVERA, TEM-

PLADO, GARCÍA-GÓMEZ, BALLESTEROS,

ORTEA, GARCÍA, ROs, Y LUQUE, 1988); o

el género Mitrella (LUQUE, 1986). Por

ello, la constatación de nuevas citas se

hace dificultosa y requiere la consulta

de numerosos trabajos muy dispersos.

En total se reportan 53 nuevas citas

para el Mediterráneo español, siendo la

mención de Trophon barvicensis y Pleuro-

tomella coeloraphe las primeras citas para

el Mediterráneo en general.

CONCLUSIONES

Este estudio confirma la presencia en

el Garraf de gran parte de la malaco-

fauna típica del Mediterráneo español

(exceptuando la zona de Alborán). Sin

embargo, se ha constatado la ausencia

de algunas especies consideradas comu-

nes en áreas cercanas como Scissurella

costata d'Orbigny, 1824, Sinezona cingu-

lata (O. G. Costa, 1861), Gibbula ardens

(von Salis, 1793), G. adansonil (Payrau-

deau, 1826), G. rarilineata (Michaud,

1829), G. umbilicaris (Linnaeus, 1758), Ju-

jubinus gravinae (Dautzenberg, 1881),

Rissoa variabilis (von Muhlfeldt, 1824),

Alvania mamillata Risso, 1826, o Mitrella

gervillei (Payraudeau, 1826), o por ejem-

plo, la presencia de sólo 2 skenéidos

pertenecientes a dos especies diferentes,

y que no se haya encontrado ningún

ejemplar de esta familia en fondos de

maérl, en los que son comunes en otras

localidades.

Finalmente, podemos concluir que la

comarca del Garraf es una zona rica en

moluscos marinos debido a las particu-

laridades de sus fondos. Prueba de ello

son las 53 nuevas citas para el Medite-

rráneo español. Muchas de estas espe-

cies viven probablemente en otras zonas

de nuestras costas, pero, debido a su

pequeño tamaño, podrían haber pasado

desapercibidas. Los autores consideran

que la prospección con medios adecua-

dos de las zonas más profundas de esta

comarca podría contribuir a ampliar el

conocimiento de la malacofauna del

litoral español, y que sería interesante

recolectar vivas algunas de las especies

que aquí se han encontrado, de las que

no se conoce el animal.

AGRADECIMIENTOS

Los autores quieren mostrar su agra-

decimiento a los hermanos J. y F. Ayza,

pescadores ya jubilados, que nos han

proporcionado numerosas muestras, así

como comentarios sobre los tipos de

fondos, etc., y a M. Roca y a sus hijos Je-

sús y Pavel que nos facilitaron desinte-

resadamente numerosos ejemplares de

asteroideos y los sedimentos de “El Pa-

rrusset”. Sin ellos, este trabajo no hu-

biera podido llevarse a cabo. También

queremos agradecer a A. Tubau el ha-

bernos cedido información de su colec-

ción. Especialmente agradecemos a C.

Palacín y M. Ballesteros (Dpt. de Biolo-

gia Animal, Universitat de Barcelona) y

a J. Templado (Museo Nacional de Cien-

cias Naturales, Madrid) sus comentarios

sobre el manuscrito original. Las correc-

ciones y comentarios constructivos de S.

Gofas (Muséum National d'Histoire Na-

turelle, Paris) y de otro revisor anónimo

han contribuido notablemente en la me-

jora de este trabajo. También agradecer a

L. Dantart (Dpto. de Biologia Animal,

GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf

Universitat de Barcelona) y C. Salas

(Dpto. de Biología Animal, Universidad

de Málaga) por la determinación de al-

gunos ejemplares. A. Warén (Swedish

Museum of Natural History, Stockholm)

y S. Gofas realizaron comentarios sobre

algunas especies, y Y. -R. Kim (Ameri-

can Museum of Natural History, New

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(1-4): 1-18.

SMRIGLIO, C., MARIOTTINL, P. Y GRAVINA, F.,

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ritrovamento di Typhlomangelia nivalis (Lovén,

1846). Contributo 1. Bollettino Malacologico,

23 (1-4): 47-52.

SMRIGLIO, C., MARIOTTINL, P. Y GRAVINA, F.,

1987b. Molluschi del Mar Tirreno Centrale:

segnalazione di alcuni Turridi provenienti da

una biocenosi a coralli bianchi. Contributo II.

Bollettino Malacologico, 23 (11-12): 381-390.

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1988a. Molluschi del Mar Tirreno Centrale:

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G. Seguenza, 1876, Fissurisepta granulosa Jef-

freys, 1883 e Propilidium ancyloide (Forbes,

1840). Contributo III. Bollettino Malacologico,

24 (1-4): 1-6.

SMRIGLIO, C., MARIOTTINI, P. Y GRAVINA, F.,

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segnalazione di Pleurotomella packardi Verrill,

1872. Contributo V. Bollettino Malacologico,

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SMRIGLIO, C., MARIOTTINI, P. Y GRAVINA, F.,

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opilina zografi (Dautzenberg éz Fischer, 1896).

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SMRIGLIO, C., MARIOTTINI, P. Y GRAVINA, F.,

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WARÉN, A. Y CARROZZA, F., 1990. Idas ghisottii

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Malacologico, 26 (1-4): 19-24.

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O Sociedad Española de Malacología Iberus, 15 (1): 95-111, 1997

Scaphopoda from the Spanish coasts

Escafópodos de las costas españolas

Gerhard STEINER*

Recibido el 20-IX-1996. Aceptado el 18-X1-1996

ABSTRACT

The scaphopod molluscs collected at 29 stations of the Fauna Iberica projects | - 1Il along

the entire Spanish mainland coast and the Balearic Islands belong to 12 species from 7

genera and 5 families. These species are listed together with their synonyms, original des-

criptions, and geographic and bathymetric ranges in this material and from the literature.

None of the species are found to extend either range in these samples. On the other hand,

some otherwise common scaphopods are not represented. Some of the species have

extensive fossil records, and in Entalina tetragona and Antalis inaequicostata, problems of

synonymy and historical biogeography are discussed.

RESUMEN

En el presente trabajo se relacionan aquellas especies de escafópodos encontradas en 29

estaciones de las costas españolas de la península Ibérica y de las islas Baleares durante

las campañas FAUNA |-1II. En total se han hallado 12 especies pertenecientes a 7 géneros

y 5 familias. Para cada especie se proporciona una lista de sinónimos, la descripción ori-

ginal y los rangos de distribución geográfica y batimétrica, obtenidos a partir de este

material y de la bibliografía. Los datos que proporcionan estas muestras no permiten

extender rangos de distribución de ninguna de las especies. Por otra parte, algunas de

las especies comúnes de escafópodos en el área no están aquí representados. Algunas de

las especies tienen registros fósiles extensos, y en Entalina tetragona y Antalis inaequi-

costata se discuten problemas relacionados con su sinonimia y biogeografía histórica.

KEY WORDS: Scaphopoda, systematic biogeography, Iberian Peninsula and Balearic Islands

PALABRAS CLAVE: Escafópodos, sistemática biogeografía, península Ibérica e islas Baleares.

INTRODUCTION

The main purpose of this paper is

the systematic treatment of Scaphopoda

collected by recent sampling cruises of

the Fauna Ibérica programme along the

Spanish coasts. The cruises Fauna I

(July, 1989), Fauna II (June - July, 1991)

and Fauna lll (June - July, 1994) covered

the Sea of Alborán, the Gulf of Cádiz,

the Bay of Biscay, the Atlantic coast off

South Galicia, and the waters around

the Balearic Islands. Scaphopod speci-

mens are represented in 29 stations of

these cruises. A list of stations of the

“Fauna I” cruise is already published

(TEMPLADO, GUERRA, BEDOYA, MORENO,

REMÓN, MALDONADO AND RAMOS,

* Institute of Zoology, University of Vienna. Althanstr. 14, A-1090 Vienna, Austria.

Iberus, 15 (1), 1997

1993); publications with the stations of

the remaining cruises are in preparation

(Iemplado, pers. comm.). Sources of

primary faunistic information on Scap-

hopoda are reports of the marine expe-

ditions “Travailleur” and “Talisman”

(LocarD, 1898), “Porcupine”, “Valo-

rous” and two smaller cruises (JEFFREYS

1870, 1877, 1882), and ALZURIA (1986,

1987). Faunistic and systematic accounts

of the Eastern Pyrenean Seas were made

by BucQuoY, DAUTZENBERG AND

DOLLFUS (1882) and Mars (1965). The

only synopsis of Iberian and Balearic

SYSTEMATIC ACCOUNT

The samples revealed 12 species from

7 genera and 5 families. Both orders of

Scaphopoda, Dentaliida and Gadilida are

represented, with 8 and 4 species respec-

tively. Each species is listed with sy-

nonyms, original description, type locality,

molluscs to date (HIDALGO, 1917) also

includes Scaphopoda. The fundamental

monography of PILSBRY AND SHARP

(1897), the comprehensive studies of

CAPROTTI (1965, 1968, 1979), and the

classification of higher taxa Of STEINER

(1992, 1996) and SCARABINO (1995) form

the basis of the systematic treatment.

Complementary information on syno-

nyms is drawn from MONTEROSATO

(1875) and STORK (1934). The material

studied in this paper is deposited in the

Museo Nacional de Ciencias Naturales,

Madrid.

distribution in the present material (with

distribution maps) and as reported in the

literature, and with the earliest fossil oc-

currence. The numbers of empty shells (e)

and of shells with soft body (b) are given

for each station.

Order DENTALIDA Da Costa, 1776

Family DENTALIDAE Gray, 1847

Genus Antalis H. and A. Adams, 1854

Antalis agilis (M. Sars, 1872) (Figs. 1, 4A)

Synonyms

Dentalium incertum Philippi 1844, Enum. Moll. Sicil. 1: 207, non Deshayes 1825.

Dentalium abyssorum var. agilis Jeffreys 1870, Ann. Mag. N. Hist. Ser. 4 VI: 74.

Dentalium agile M. Sars 1872, Some remarkable forms etc., Christiania 1872: 34.

Dentalium fusticulus Brugnone 1876, Misc. Malac. 11: 21.

Dentalium vagina Jeffreys 1877, Ann. Mag. N. Hist. Ser. 4, XIX: 155.

Antalis agilis (M. Sars): G. O. Sars 1878, Moll. Reg. Arct. Norv., Christiania 1878: 102.

Dentalium (Antalis) calabrum Crema 1910, p. 68.

Original description: Shell slender,

very narrow, slightly curved, almost

straight, gradually attenuated towards the

apex. White, faint luster, posterior part fre-

quently darker. Apex very narrow, obli-

quely truncated, with a tolerably deep inci-

sion and a short, hardly protruding sup-

plementary tube. Shell surface with cir-

cular growth lines, rarely longitudinally

striated in the posterior part, the striae

being little distinct and never prominent

ribs. [... ] Largest shell 58 mm long and 4

mm in diameter, 1 mm at the apex.

Type locality: Lofoten Islands,

North Atlantic, at 360-540m.

Present material: 5 stations; Gulf of

Cádiz, 500-546 m (76A: 1b, 20e; 77A: 3b);

Cape Finisterre, 129-133 m (91A: 1b);

Biscay, 540-1025 m (124A: 2b, 9e; 1594:

2b, 14e).

Reported distribution: North Atlan-

tic: Portugal to Lofoten, Halifax to Cuba,

Gulf of Mexico, Azores; Mediterranean;

?Red Sea; 60-5000 m.

STEINER: Scaphopoda from the Spanish coasts

Earliest fossil: Pliocene.

Remarks: The largest specimen from

station 159A is 65 mm long, which exce-

eds indications of SARS (1872), PILSBRY

AND SHARP (1897) and CAPROTTI (1965),

but matches those of LOCARD (1898).

There are no shells with apical slits in the

present material, but most of the speci-

mens are empty shells and fragments.

The shell surface in the apical region

may be eroded by boring organisms

even in live animals. The shell then has a

chalky and often crackled aspect, as was

also remarked by Sars (1872). Young in-

dividuals are often more or less dis-

tinctly ribbed near the apex. The ribs

intercalate to about 20 in number and

then gradually become obsolete. The

younger parts of the shell (towards the

anterior opening) with an intact surface

are glossy and bear closely spaced

growth lines. LOCARD (1898) distinguis-

hed a number of varieties according to

size, curvature and sculpture. Antalis pa-

norma (Chenu, 1842-47), similar in size

and shape to A. agilis and reported from

the Mediterranean and the Bay of Biscay,

differs in being more curved and having

a more solid shell with 12 narrow but

pronounced primary ribs.

Some of the shells from station 76A

have bore holes of naticid gastropods

being, apart from different fishes,

important predators of scaphopods in

the Mediterranean.

Antalis entalis (Linné, 1758) (Figs. 1, 4B)

Synonyms

Dentalium entalis Linné 1758, Syst. Nat. (10): 785.

Dentalium entalum L.: Blainville 1819, Dict. Sc. Nat. XUl: 70.

Dentalium labiatum Brown 1827, 111. Conch. Gr. Brit. and Irel.: pl. 1, fig. 4.

Dentalium striolatum Stimpson 1851, Proc. Bost. Soc. Nat. Hist. IV: 114. non Jeffreys, Watson, Sars,

Risso.

Entalis striolata (Stimpson 1851): Gould-Binney 1870, Invert. of Mass.: 266.

Original description: Shell smooth,

moderately curved, continuous, not

fractured.

Type locality: Atlantic Ocean.

Present material: 2 stations; La

Coruña, 151-152 m (101A: le); Biscay,

119-122 m (112DH: 2e).

Reported distribution: North Atlan-

tic from Spain north to Spitzbergen and

Maine, Massachusetts to Bay of Fundy;

6-3500 m.

Earliest fossil: Pliocene.

Remarks: The shell is up to 42 mm

long, solid, white, sometimes glossy, mode-

rately curved but in the apical region, the

wider, anterior part of the shell being only

slightly curved. The shell surface is

smooth, very fine longitudinal striae may

be present in the apical region only.

Towards the anterior opening growth lines

become more distinct. There may be a

shallow apical notch on the convex side.

This species is infrequently cited

from the Mediterranean as well, due to

confusion with the apically striated An-

talis vulgaris. However, HIDALGO (1917)

retains Gibraltar and Mataró / Catalunya

as localities for A. entalis.

Antalis dentalis (Linné 1766) (Figs. 1, 4C)

Synonyms

Dentalium dentalis Linné 1766, Syst. Nat. XII: 1263.

Dentalium dentale L.: Locard 1886, Ann. Soc. Agricult., Lyon Ser. 5, IX: 145.

Dentalium linnaeum Locard 1886, Ann, Soc. Agricult., Lyon Ser. 5, IX: 145.

Dentalium mutabile Dóderlin in Hórnes 1856, Abhandl. K. -K. Geol. Reichsanst. Il: 654.

Iberus, 15 (1), 1997

Original description: Shell striated,

moderately curved, fractured.

Type locality: Mediterranean Sea.

Present material: 1 station; Gulf of

Cádiz, 13-15 m (714: 1e).

Reported distribution: Mediterra-

nean Sea, East Atlantic from Galicia to

Cape of Good Hope (?), Azores, Canary

Islands; 0-300 m.

Earliest fossil: Miocene.

Remarks: This rather small species

is up to 24 mm long (13 mm from

station 71A), mostly white, only the

apex sometimes with a rose tinge. There

are about 10 sharp and narrow primary

ribs, becoming doubled by intercalation

towards the anterior opening. The

secondary ribs are of about the same

height as the primary ribs. The intercos-

tal spaces are much wider than the ribs,

and smooth except for widely spaced

growth lines. Antalis dentalis is often

confused with A. inaequicostata (see

below).

Antalis inaequicostata (Dautzenberg, 1891) (Figs. 1, 4D)

Synonyms

Dentalium dentalis Lamarck 1818, Anim. sans vert. V: 344; Deshayes 1825, Anat. et Monogr. du genre

Dentale: 33; Risso 1826, Hist. Nat. Europ. Merid. IV: 398; Philippi 1836, Enum. Moll. Sicil. 1: 243;

Jeffreys 1870, Ann. Mag. Nat. Hist. VI: 10; Monterosato 1872, Not. Int. alle conch. Medit.: 28; non

Linné 1766.

Dentalium fasciatum Lamarck 1818, Anim. sans vert. V: 343; non Gmelin 1790

Dentalium pseudo-antalis Scacchi 1836, Catal. Conch. Regni Neap.: 17; non Lamarck 1818

Dentalium novem-costatum Réquien 1848, Coq. de Corse: 90; non Lamarck 1818.

Dentalium novemcostatum var. tenuis Monterosato 1878, Enum. et Sinon.: 16; non Lamarck 1818.

Dentalium novemcostatum Réquien: Monterosato 1884, Nom. Gen. e Spec.: 31; non Lamarck 1818.

Dentalium alternans Bucquoy, Dautzenberg and Dollfus 1891, Moll. Mar. Roussillon 1: 561; non

Chenu 1842.

Dentalium inaequicostatum Dautzenberg 1891, Mem. Zool. Soc. France 1891: 53.

Antale novemcostatum (Réquien): Sacco 1897, Moll. terr. terz. Piemonte e delle Liguria XXI: 104.

Dentalium (Antalis) inaequicostatum B. D. D.: Pilsbry and Sharp 1897-98, Man. Conch. XVII: 52.

Caprotti 1965, Atti. Soc. Ital. Sci. Nat. Milano 105: 343.

Dentalium novemcostatum var. inaequicostata Fantinet 1959, Serv. Carte Geol. Algérie 1: 46.

Dentalium (Antalis) novemcostatum Réquien: Caprotti 1961, Atti. Soc. Ital. Sci. Nat. Milano 100: 353.

Dentalium (Antalis) mutabile inaequicostatum Dautzenberg: Caprotti 1979, Boll. Malacol. Milano 15: 233.

Original description: Shell solid,

opaque, slightly to moderately curved,

straightening towards the anterior aper-

ture. Sculpture of 9 or 10 primary ribs

alternating with same number of wider

and less protruding secondary ribs. All

ribs become obsolete towards the ante-

rior. Numerous transverse growth lines,

sometimes with irregular fractures or

interruptions. Anterior shell aperture

slightly polygonal. Posterior aperture

truncated, polygonal, with an oval,

short central pipe. No slit or notch.

Colour light rose, more intense at the

posterior end, transversal bands of

lighter and darker colour. Shell 35 mm

long, 5 mm at anterior aperture.

Type locality: Mediterranean Sea.

Present material: 4 stations; Gulf of

Cádiz, 13-28 m (44A: 2e; 66A: 9b, 3e;

69A: le; 71A: about 10b, many empty

shells).

Reported distribution: Mediterra-

nean from Greece to Algeria; 5-120m.

Earliest fossil: Miocene (BUCQUOY ET

AL., 1886) or Pliocene (CAPROTTI, 1979).

Remarks: This species is extremely

variable in its longitudinal sculpture

causing considerable confusion in

studies of both fossil and recent scapho-

STEINER: Scaphopoda from the Spanish coasts

MW Antalis agilis

O Antalis dentalis

Ba Antalis novemcostata

A Antalis entalis

O Antalis inaequicostata

e Antalis vulgaris

Figure 1. Localities where Antalis agilis, A. entalis, A. dentalis, A. inaequicostata, A. novemcostata, and

A. vulgaris were collected.

Figura 1. Localidades donde se encontraron las especies Antalis agilis, A. entalis, A. dentalis, A. inae-

quicostata, Á. novemcostata y A. vulgaris.

pods. Small individuals of this species

closely resemble Antalis dentalis. The

latter differs, however, in having conspi-

cuous transversal striae between the

ribs, the ribs themselves are less acute,

and the intercostal space is not as wide.

If PILSBRY AND SHARP (1897-98, p. 52)

describe “... 9-12 strong primary ribs

towards the apex, narrower than their

intervals... “ for A. inaequicostata, they

obviously did not consider direct com-

parison with A. dentalis. The atlantic A.

novemcostata is stouter and has more

conspicuous transverse striae.

Many of the empty shells shows cha-

racteristic signs of sipunculid occupa-

tion. Phascolion strombus (Montagu,

1804) is known to close the anterior ope-

nings of scaphopod and other mollusc

shells by agglutinations of sediment,

leaving open only a small tube for their

introvert (TÉTRY, 1959).

Iberus, 15 (1), 1997

Antalis novemcostata (Lamarck, 1818) (Figs. 1, 4E)

Synonyms

Dentalium novemcostatum Lamarck 1818, Anim. sans vert., V: 344.

Dentalium dentalis Risso 1826, Hist. Nat. Europ. Mérid., IV: 398.

Dentalium dentale Risso: Weinkauff 1862 (partim), J. Conch., X: 364.

Antale novemcostatum (Lamarck): Sacco 1896, Boll. Mus. Zool. Anat. Comp. Univ. Torino, XI: 97.

Dentalium (Antalis) novemcostatum (Lamarck): Pilsbry and Sharp 1897-98, Man. Conch., 17: 51

Original description: Shell small,

greenish-white, with nine ribs, subde-

cussate transverse striae.

Type locality: Atlantic coast near La

Rochelle.

Present material: 1 station, Gulf of

Cádiz, 110-112 m (694: 1e).

Reported distribution: East Atlantic

from La Rochelle to South Spain; 20-300 m.

Earliest fossil: Pliocene.

Remarks: The rather stout shell is up

to 32 mm long and has 8 to 10 rounded

ribs decreasing in heigth towards the ante-

rior opening. The intercostal spaces are

more concave than in the other Antalis

species of the region and show faint lon-

gitudinal striae. Transverse striae are not

always developed, although BUCQUOY ET

AL. (1886) and CAPROTTI (1965) take the

strong transverse sculpture as an impor-

tant character to distinguish A. novem-

costata from the mediterranean A. inae-

quicostata. The apex often has a plug with

a central pipe in larger specimens.

Antalis novemcostata seems to be

living on the european Atlantic coast

only, although HIDALGO (1917) lists

several mediterranean locations for this

species. This seems to be due to misiden-

tifications of A. inaequicostata (CAPROTTI,

1961, 1965; Mars, 1965). According to

CAPROTTI (1965), the extremely rare A.

novem-costatum from the Italian Pliocene

could be intermediate between A. inae-

quicostata and A. novemcostata.

Antalis vulgaris (Da Costa, 1778) (Figs. 1, 4F)

Synonyms

Dentale vulgare Da Costa 1778, Brit. Conch.: 24.

Dentalium fasciatum Gmelin 1791, Syst. Nat., 13: 3737.

Dentalium striatum Montagu 1803, Test. Brit., II: 492, non Born 1780, Test. Mus. Caes. Vindob.: 431.

Dentalium tarentinum Lamarck 1818, Anim. sans vert., V: 345. Forbes and Henley 1853, Hist. Brit.

Moll., II: 451. Sowerby 1860, Thes. Conch., 11: 100. Jeffreys 1882, Brit. Conch., II: 195. Clessin 1896,

Conchyl. Cab.: 3.

Dentalium politum Blainville 1819, Dict. Sci. Nat., XI: 70. Turton 1819 (partim), Conch. Dict. Brit.

Sh.: 38.

Dentalium labiatum Turton 1819 (partim), Conch. Dict. Brit. Sh.: 38. Brown 1827, Illustr. Conch. Gr.

Brit.: 117.

Dentalium striolatum Risso 1826, Hist. Nat. Europ. Mérid., IV: 398.

Dentalium multistriatum Risso 1826, Hist. Nat. Europ. Mérid., IV: 398, non Deshayes 1825, Anat. et

Monogr. du genre Dentale.

Dentalium affine Biondi 1859, Atti Accad. Gioenia Sci. Nat. (2), XIV: 120.

Original description: Dentalium

with a slender, smooth, glossy, subar-

cuated shell, tapering to a small point,

pervious: sometimes marked with a few

circular wrinkles or annulations: colour

white or yellowish. Length an inch and

100

a half [38 mm]; diameter at the larger

end two-tenths of an inch [5 mm]; and

one fourth as much [1.25 mm] at the

smaller end. [... ] A variety is marked

with dusky bands; and sometimes a

little striated towards the point.

STEINER: Scaphopoda from the Spanish coasts

E Fissidentalium capillosum

O Entalina tetragona

Xx Dischides politus

A Episiphon filum

O Pulsellum lofotense

e Cadulus jeffreysi

Figure 2. Localities where Fissidentalium capillosum, Episiphon filum, Entalina tetragona, Pulsellum

lofotense, Dischides politus, and Cadulus jeffreysi were collected.

Figura 2. Localidades donde se encontraron las especies Fissidentalium capillosum, Episiphon filum,

Entalina tetragona, Pulsellum lofotense, Dischides politus y Cadulus jeffreysi.

Type locality: British shores, espe-

cially Scilly Islands, Cornwall, Devons-

hire, Hampshire.

Present material: 5 stations; Biscay,

119-122 m (112DH: 1e); Balearic Islands,

5-59 m (190B: 2e; 192A: 1b; 203B: 1e;

258B: 3b).

Reported distribution: Mediterranean

Sea and East Atlantic Ocean; 5-1100 m.

Earliest fossil: Miocene.

Remarks: The shell is up to 60 mm

long, white with a rose-coloured apex,

rather broad, and moderately curved in

the posterior half. There are about 30

longitudinal striae near the apex, oblite-

rating gradually towards the anterior

opening. The apical opening is entire

and may have a plug with a short central

pipe.

101

Iberus, 15 (1), 1997

Genus Fissidentalium Fischer, 1885

Fissidentalium capillosum (Jeffreys, 1876) (Figs. 2, 4G)

Synonyms

Dentalium capillosum Jeffreys 1876, Proc. Roy. Soc., 25: 185. (nomen nudum)

Dentalium capillosum Jeffreys: Jeffreys 1877, Ann. Mag. Nat. Hist. Ser. 4, 19: 153.

Dentalium (Fissidentalium) capillosum Jeffreys: Pilsbry and Sharp 1897-98, Man. Conch., 17: 77.

Original description: Shell tapering

to a fine point, slightly curved, rather solid,

opaque, and mostly lusterless; sculpture:

numerous and sharp (not rounded) lon-

gitudinal striae, some of which are inter-

mediate and smaller than the rest; they

disappear towards the posterior or narrow

end, which is quite smooth and glossy for

a quarter of an inch [6.4 mm]; colour

whitish; margin at the posterior end having

a short and narrow notch. L [length]: 1.4

[35.6 mm]. B [maximum diameter]: 0.15

[3.8 mm]. (...) This appears to attain a size

considerably exceeding that given in the

above description, as fragments measure

nearly 0. 4 inch [10 mm] in breadth.

Type locality: North Atlantic, Valo-

rous st. 12, 13, 16; 1242-3213 m.

Present material: 1 station; Biscay,

925-1025 m (159A: 5e).

Reported distribution: North Atlan-

tic, Caribbean Sea to Portugal, Azores to

Hebrides; 400-3500 m.

Earliest fossil: No fossil record.

Remarks: This species can be 81 mm

long. The shell is white or grey and may

be somewhat eroded. There are about 65

fine ribs throughout most of the length.

Pilsbry and SHARP (1897-98) supplement

JEFFREYS' (1877) description saying that

the ribs are sharply cut but rounded on

the top. This is particularly obvious in

the anterior part of the shell where the

ribs become wider.

Family GADILINIDAE Chistikov, 1975

Subfamiliy EPISIPHONINAE Chistikov, 1975

Genus Episiphon Pilsbry and Sharp, 1897-98

Episiphon filum (Sowerby, 1860) (Figs. 2, 4H)

Synonyms

Dentalium filum Sowerby 1860, Thes. Conch., III: 89.

Dentalium gracile Jeffreys 1870, Ann. Mag. Nat. Hist. (4), VI: 74. Fischer 1873, Journ. Conchyl.: 140.

Pseudantalis filum (Sowerby): Monterosato 1884, Nom. Gen. Spec. Conch. Medit.: 33.

Dentalium rufescens Weinkauff 1868 (partim), Conch. Mittelm., Il: 420.

Original description: Shell slender,

very narrow, thin, finely pointed, mantle

reddish brown, apex entire.

Type locality: Gibraltar.

Present material: 2 stations; Biscay,

104-132 m (152A: 2b, le; 153A: 2b).

Reported distribution: North Atlan-

tic from Florida to Cape Hatteras,

Algeria to Biscay; Mediterranean Sea

from Aegean to Gibraltar; 20-4784 m.

102

Earliest fossil: Miocene.

Remarks: This is a very characte-

ristic species, being very narrow,

hardly tapering and almost straight.

The length is up to 13 mm. The shell

is white and glossy, semitransparent

and extremely fragile. The sculpture

consists of growth lines only. Typical

for the genus is a long pipe at the

apex continuous with the shell. It

may be wanting because of its fragi-

lity.

STEINER: Scaphopoda from the Spanish coasts

Figure 3. Bathymetric ranges of species reported in literature (shaded) and in the present material

(black).

Figura 3. Rangos batimétricos de las especies: datos bibliográficos (gris), datos del presente material (negro).

Order GADILIDA Starobogatoy, 1982

Suborder ENTALIMORPHA Steiner, 1992

Family ENTALINIDAE Chistikov, 1979

Subfamily ENTALININAE Chistikov, 1979

Genus Entalina Monterosato, 1872

Entalina tetragona (Brocchi, 1814) (Figs. 2, 5A)

Synonyms

Dentalium tetragonum Brocchi 1814, Conch. foss. subapen., Milano: 627.

Dentalium quinquangulare Forbes 1844, Rep. Brit. Ass. Adv. Sci. for 1844: 188.

Siphonodentalium pentagonum M. Sars 1865, Forh. Videsk. Selsk. Christiania 1864: 307.

Dentalium quinquangulatum Reeve 1872, Conch. Icon.: pl. 5, fig 45.

Siphonodentalium quinquangulare (Forbes): Jeffreys 1867, Ann. Mag. Nat. Hist. Ser. 3, XX: 251.

Weinkauff 1868, Conch. Mittelm., 1: 421. Locard 1886, Ann, Soc. Agricult., Lyon Ser. 5, IX: 149.

Dautzenberg 1891, Mem. Soc. Zool. France, IV: 609. Friele and Grieg 1901, Norv. N. Atlant. Exp.

etc., Christiania 1901, Mollusca III: 50.

Siphonentalis tetragona (Brocchi): G. O. Sars 1878, Moll. Reg. Arct. Norv., Christiania 1878: 105.

103

Iberus, 15 (1), 1997

Siphodentalium tetragonum (Brocchi): Norman 1879, J. Conch., London, 2: 49.

Entalina tetragona (Brocchi): Monterosato 1880, Bull. Soc. Malac. Ital., VI: 64. Caprotti 1961, Atti

Soc. Ital. Sci. Nat. Mus. Civ. Stor. Nat. Milano, Vol. C, IV: 356.

Siphodentalium quinquangulare (Forbes): Jeffreys 1882, Proc. Zool. Soc. London, 1882: 662.

Siphonentalis quinquangularis (Forbes): Carus 1889, Prodr. faunae Medit., 11: 176.

Pulsellum quinquangulare (Forbes): Norman 1893, Ann. Mag. Nat. Hist. Ser. 6, XII: 344, 362.

Entalina quinguangularis (Forbes): Hidalgo 1917, Trab. Mus. Nac. Cienc. Nat. Ser Zool. 30: 306.

Chistikov and Sagaidachniy 1982, Zool. Zhurn., 60: 38.

Original description: Shell four-

angled, finely longitudinally striated,

sides weakly carinated.

Type locality: Pliocene of Italy

(BROCcHI, 1814), Aegean Sea (FORBES,

1844).

Present material: 6 stations; Medite-

rranean, 1001-1005 m (247A: le); Sea of

Alborán, 276-306 m (15A: 2e); Gulf of

Cádiz, 500-546 m (76A: 3e; 77A: 2e);

Galicia, 81-84 m (86DL: 1b), Biscay, 540-

543 m (124A: 5b, 15€e).

Reported distribution: Mediterra-

nean, East Atlantic from Biscay to Nort-

hern Norway; 10-2664 m.

Earliest fossil: Miocene.

Remarks: The shell is strongly

curved, at least in the apical half. It has

five primary ribs, four of them forming

almost right angles, the fifth rib on the

midline of the concave side forms an

obtuse angle. In specimens larger than

about 10 mm, 3 - 25 secondary ribs gra-

dually appear, the pentagonal form of the

cross-section smoothing out to become

subcircular. The anterior opening is

oblique, the apex simple and entire. The

shells may be up to 93 mm long but are

usually around 15 mm.

There is some confusion in the Euro-

pean species and the use of the names te-

tragona Brocchi, pentagona Sars and quin-

quangularis Forbes. BROCCHI (1814) des-

cribed tetragona as a Pliocene fossil from

Piemont and the Vienna Basin. FORBES

(1844) described the extant species from

the Aegean Sea as quinquangularis. MiI-

CHAEL SARS (1865) described pentagona

from the Norwegian coasts, without refe-

rring to either Brocchi or Forbes. His son,

G. O. SArs (1878) considers pentagona and

quinquangularis as janior synonyms of te-

tragona. MONTEROSATO (1880) comes to the

same conclusion claiming the fossil and

recent mediterranean species being iden-

tical. JEFFREYS (1870, 1882), however, is of

different opinion and gives priority to

quinquangularis for the recent Atlantic and

Mediterranean form. PILSBRY AND SHARP

(1897-98), LOCARD (1898) and FRIELE AND

GRIEG (1901) agree with this view.

CAPROTTI (1968), finally, comparing re-

cent specimens with the type material,

confirmes the synonymity of quinquangu-

laris and tetragona, stressing the priority

of Brocchi's name. Later, CHISTIKOV AND

SAGAIDACHNIY (1982), however, not only

separate tetragona and quinquangularis, but

also split the latter into the Mediterranean

quinguangularis and the Atlantic pentagona

on grounds of shell and radula characters.

However, they do not include fossil spe-

cimens in their study.

Suborder GADILIMORPHA Steiner, 1992

Family PULSELLIDAE Scarabino in Boss, 1982

Genus Pulsellum Stoliczka, 1868

Pulsellum lofotense (M. Sars, 1865) (Figs. 2, 5B)

Synonyms

Siphonodentalium lofotense M. Sars 1865, Forh. Videsk. Selsk. Christiania 1864: 297.

Siphonentalis lofotensis (M. Sars): G. O. Sars 1878, Moll. Reg. Arct. Norv., Christiania 1878: 104. Mon-

terosato 1884, Nom. Gen. Spec. Conch. Medit.: 33.

104

STEINER: Scaphopoda from the Spanish coasts

Figure 4. A: Antalis agilis, B: Antalis entalis, C: Antalis dentalis, D: Antalis inaequicostata; E: Antalis

novemcostata; E: Antalis vulgaris, G: Fissidentalium capillosum; H: Episiphon filum. Scale bars, A, G:

10 mm; B-F: 2 mm; H: 1 mm.

Figura 4. A: Antalis agilis; B: Antalis entalis; C: Antalis dentalis; D: Antalis inaequicostata; E: Antalis

novemcostata; F: Antalis vulgaris; G: Fissidentalium capillosum; A: Episiphon filum. Escalas, A, G:

10 mm, B-E: 2 mm; H: 1 mm.

Iberus, 15 (1), 1997

Siphodentalium lofotense M. Sars: Jeffreys 1882, Ann. Mag. Nat. Hist., Ser. 5, XI: 395.

Siphonodentalium (Pulsellum) lofotense M. Sars: Pilsbry and Sharp 1897-98, Man. Conch., 17: 138.

Siphonodentalium lofotensis M. Sars: Stork 1934, Thalassia, 1: 10.

Pulsellum lofotensis (M. Sars): Emerson 1962, Journ. Paleontol. 36: 475.

Original description: Shell smooth,

moderately curved, anteriorly wide and

tapering to the posterior, white, walls

transparent or semitransparent, thin,

shiny, very fine and dense obliquely

transverse growth lines well visible,

posterior shell margin entire [plain].

Length 5-6 mm, basal width 0.66 mm,

apical about 0.33 mm. [Description of

soft body omitted.]

Type locality: Lofoten Isl., Norway;

90-216 m.

Present material: 4 stations; Gulf of

Cádiz, 535-546 m (76A: 3e); Galicia, 80-

120 m (1684: 3e; 171A: 3e); Biscay, 129-

132 m (153A: 5b, about 20e).

Reported distribution: Mediterra-

nean Sea; North Atlantic from Spain to

Finmark, Ireland, New England; 26-

3500 m.

Earliest fossil: Pliocene.

Remarks: The small shell is rather

fragile in the anterior third and easily

breaks into cylindric fragments upon

handling. Breakage occurs along the

oblique growth lines. The anterior ope-

ning is circular as the apical opening but

slightly oblique. Most live animals have

perfectly transparent shells, empty

shells are opaque. The apical rim is al-

ways entire without notches or lobes,

although JEFFREYS (1882) mentions spe-

cimens with regularly jagged tips. This

may have been a siphonodentaliid spe-

cies, also because he did not see a bul-

bous larval shell, which is well develo-

ped in Pulsellum lofotense (Steiner, 1995).

Family GADILIDAE Stoliczka, 1868

Subfamily SIPHONODENTALIINAE Simroth, 1894

Genus Dischides Jeffreys, 1867

Dischides politus (Wood, 1842) (Figs. 2, 5C)

Synonyms

Ditrupa polita Wood 1842, Ann. Mag. Nat. Hist., 9: 459.

Dentalium coarctatum Philippi 1844, Enum. Moll. Sicil., UI: 208, non Lamarck 1818, Anim. sans Vert., 5: 346.

Dentalium laevigatum de Rayneval, Hecke and Ponzi 1854, Cat. Foss. Mont Mario, Versailles, non

Schlotheim 1830.

Dentalium bifissum Jeffreys 1867, Ann. Mag. Nat. Hist., Ser. 3, XX: 251. Weinkauff 1868, Conch.

Mittelm., 1: 421. Monterosato 1884, Nom. Gen. Spec. Conch. Medit.: 34.

Dischides olivi Jeffreys 1870, Ann. Mag. Nat. Hist., Ser. 4, VI: 73.

Dischides bifissus (Jeffreys): Jeffreys 1882, Proc. Zool. Soc. London, 1882: 663.

Cadulus politus (Wood): Pilsbry and Sharp 1897-98, Man. Conch., 17: 144. Stork 1934, Thalassia, 1: 9.

Original description: Shell slightly

arcuated, thin, smooth, subcylindrical;

anterior opening plain, posterior cleft,

bilateral, with unequal terminations.

The body of the [... ] shell is not inflated

or enlarged like that of Dentalium gadus,

but has the posterior opening laterally

cleft, somewhat resembling that of Den-

talium coarctatum Deshayes [... ] but the

dorsal part of the posterior end of this

106

fossil is produced beyond the edge

beneath and rounded, the ventral edge

is shorter and truncated, an enamel-like

polish covers the exterior, and was pro-

bably when inhabited subhyaline, but is

now opaque. Length half an inch [12.7

mm] nearly.

Type locality: Coralline

England (Pliocene).

Crag,

STEINER: Scaphopoda from the Spanish coasts

Figure 5. A: Entalina tetragona; B: Pulsellum lofotense, C: Dischides politus, D: Cadulus jeffreysi. Scale

bars 1 mm.

Figura 5. A: Entalina tetragona; B: Pulsellum lofotense; C: Dischides politus; D: Cadulus jeffreysi.

Escalas 1 mm.

Present material: 1 station; Gulf of

Cádiz, Trafalgar, 34 m (58A: 1e).

Reported distribution: Northeast

Atlantic from Marocco to Biscay, Medi-

terranean; 9-324 m.

Earliest fossil: Pliocene.

Remarks: The recent members of

this species attain maximum lengths of

7 mm. The shell is white, glossy, mode-

rately curved and only slightly tapering.

The sculpture consists of growth lines

only. The greatest diameter of the shell

lies just behind the anterior opening.

The posterior opening has two lateral

notches producing a dorsal and a

ventral lobe. Tubes of the serpulid poly-

chaete Ditrupa sp. must not be confused

with Dischides politus. The polychaete

tubes are often reddish brown in colour,

the outer layer has a semitransparent

aspect. They are sharply constricted at

the anterior opening and lack growth

lines.

Dischides has repeatedly been chan-

ging between subgenus and genus

status. It was recently confirmed in its

generic rank and transferred to the sub-

family Siphonodentaliinae by SCARA-

BINO (1995).

1107

Iberus, 15 (1), 1997

Subfamily GADILINAE Stoliczka, 1868

Genus Cadulus Philippi, 1844

Cadulus jeffreysi (Monterosato, 1875) (Figs. 2, 5D)

Synonyms

Helonyx jeffreysi Monterosato 1875

Cadulus jeffreysi (Monterosato): Jeffreys 1882, Verrill 1882, Pilsbry and Sharp 1897-98, Muus 1959

Cadulus propinquus Verrill 1885, non G. O. Sars

Cadulus subfusiformis Stork 1934, non M. Sars

Original description: Anterior aper-

ture obliquely truncated, base or [?and]

posterior aperture compressed, slightly

deformed at each side.

Type locality: Aegean Sea, 234-450 m.

Present material: 1 station; Gulf of

Cádiz, 535-546 m (76A: 18€).

Depth range: 535-546 m.

Reported distribution: Mediterra-

nean; North Atlantic from Canary Ís.,

Biscay to Ireland and Norway, West

Atlantic from Martha's Vineyard to Bar-

bados, South Atlantic at St. Helena; 90-

2200 m.

Earliest fossil: Pliocene

DISCUSSION

Of the 19 valid species listed for the

Iberian coasts by HIDALGO (1917), 12 are

represented in the present material. Ta-

king into account the recent finding of

the Eastern Mediterranean species Anta-

lis rossati near Barcelona (Alzuria, 1986)

and the deep-water Biscayan material of

the “Talisman”, “Travailleur” (LOCARD,

1898) and “Valorous” (JEFFREYS, 1877)

expeditions, the species list of Iberian

scaphopods grows to 31. Species treated

in these reports but not present in this

material are listed in Table I.

The present findings provide no bio-

geographic novelties. Species living in

both the Atlantic and Mediterranen Sea

are Antalis agilis, A. dentalis, A. vulgaris,

Episiphon filum, Entalina tetragona, Pulse-

llum lofotense, Dischides politus and Cadu-

108

Remarks: Shell small, smooth and

shiny, moderately curved, with a conspi-

cuous swelling just anterior to the middle;

ventral side regularly curved, dorsal side

with distinct convex area due to the swe-

lling; anterior shell aperture obliquely trun-

cated facing downwards, slightly laterally

compressed; posterior aperture without

lobes, dorsoventrally depressed.

This species may be confused with

Cadulus subfusiformis (M. Sars 1865). It

differs in being larger, conspicuously

swollen near the middle of the shell and

having a distinct convex area in the

dorsal line. The anterior aperture is

slightly laterally compressed, the poste-

rior aperture dorsoventrally depressed,

while C. subfusiformis has a faintly dorso-

ventrally depressed anterior end and a

round posterior one.

lus jeffreysi. The only typical Mediterra-

nean forms in the material is A. inaequi-

costata. This latter shallow-water form

extends, however, beyond the Strait of

Gibraltar into the Gulf of Cádiz. On the

other hand, A. entalis, A. novemcostata

and Fissidentalium capillosum are known

from Atlantic waters only.

Comparing the bathymetric data of

this material with reported ranges (Fig.

3), only Antalis agilis, A. inaequicostata

and Entalina tetragona cover the greater

part of their ranges. Most of the other

species are found a one or two stations

only, which may explain their relatively

restricted bathymetric occurrence.

CAPROTTI (1968) considers Cadulus oli-

vii, C. strangulatus and C. tumidosus doubt-

ful species or mere variations of C. ovulus.

STEINER: Scaphopoda from the Spanish coasts

Table I. Scaphopoda reported from the Iberian coasts but not in present material. Asterisks mark

doubtful species (see Discussion).

Tabla 1. Escafópodos citados en las costas ibéricas pero no hallados en el presente material. Los asteríscos

indican especies dudosas (ver Discusión)

Species Area

Order DENTALIIDA

Family DENTALIIDAE

Antalis panorma (Chenu, 1842-47)

Antalis rossati (Caprotti, 1966)

Fissidentalium candidum (Jeffreys, 1877) Biscay

FUSTIARIIDAE

Fustiaria rubescens (Deshayes, 1825) Mediterranean

1917, Alzuria, 1987

Order GADILIDA

Suborder ENTALIMORPHA

Family ENTALINIDAE

Subfamily BATHOXIPHINAE

Bathoxiphus ensiculus (Jeffreys, 1877)

Subfamily HETEROSCHISMOINAE

Heteroschismoides subterfissum

Suborder GADILIMORPHA

Family GADILIDAE

Subfamily SIPHONODENTALIINAE

Siphonodentalium lobatum (Sowerby, 1860) Atlantic, Portugal

Subfamily GADILINAE

Cadulus artatus Jeftreys, 1880

Cadulus subfusiformis (M. Sors, 1865)

Atlantic, Biscoy

Cadulus gracilis Jeffreys, 1877 Biscoy

Cadulus propinquus 6. O. Sars, 1878 Biscay

Cadulus cylindratus Jeffreys, 1877 Biscay

Cadulus monterosatoi Locard, 1897

Codulus gibbus Jeffreys, 1882

Cadulus ovulus (Philippi, 1844)

Cadulus amphorus Jeffreys, 1882

*Codulus olivii (Scacchi, 1835)

*Cadulus tumidosus Jeffreys, 1877

*Cadulus strangulatus Locard, 1897

Biscoy

Biscay

The occurrence of C. subfusiformis, a wide-

spread Atlantic species, in the Mediterra-

nean remains controversary. Although

MONTEROSATO (1880) identifies a varia-

tion (var. abyssicola) from Palermo, CA-

PROTTI (1968) does not list this species as

constant inhabitant of the Mediterranean.

Recently, GAGLINI (1985) reconfirmed C.

subfusiformis, and also MIFSUD's (1996) pho-

Mediterranean, Northeast Spain, Balearic ls.

Mediterranean, Northeast Spain

Atlantic, Portugal, Biscay

Atlantic, Portugal,

Atlantic, Biscay, Mediterranean

Atlantic, Galicia, Portugal

Biscoy, Mediterranean

-Aílantic, South Portugal

Atlantic, Biscay, Galicia, Southern Portugal

Biscay, Mediterranean off Morseilles

Reference

Locard, 1898; Hidalgo, 1917

Alzuria, 1986

Jeffreys, 1877

Locard, 1898; Hidalgo,

Jeffreys, 1877

Locard, 1898

Jeffreys, 1882

Locard, 1898

Locard, 1898; Monterosato, 1875

Locard, 1898

Locord, 1898

Locard, 1898

Jeffreys, 1882

Locord, 1898

Locard, 1898

tograph of C. jeffreysi from Malta looks

more like subfusiformis. However, until

further reports come in, it remains possi-

ble that in the Mediterranean C. subfusi-

formis occurs in episodic pseudopopula-

tions only (BOUCHET AND TAVIANI, 1992).

Long lists of synonyms as for some

Mediterranean scaphopod species may

have several reasons. One of them, espe-

109

Iberus, 15 (1), 1997

cially when generic designations vary, is

progress in supraspecific systematics.

Another reason may be closely related but

morphologically variable species and / or

similar but not identical fossil forms. The

species complex of Antalis inaequicostata,

novemcostata and the Miocene mutabile (Do-

derlein in HÓRNES, 1856) is a good exam-

ple for a combination of these causes. An-

talis mutabile and A. inaequicostata are very

similar and both highly variable in their

shell features. BUCQUOY ET AL. (1886) re-

cognize the close relationship of mutabile

and inaequicostata (=alternans). PILSBRY AND

SHARP (1897-98) go further and list muta-

bile as synonym of both dentalis (p. 53) and

novemcostata (p. 211). RUGGIERI (1948) sug-

gests mutabile being ancestral to novem-

costata. CAPROTTI (1979) takes the alterna-

tive view and presents the recent inaequi-

costata as subspecies of the fossil mutabile.

On the other hand, he assigns species sta-

tus to novemcostata, although considering

it the Atlantic descendant of mutabile. Fi-

nally, PavIA's (1991) questionable assign-

ment of mutabile to the genus Fissidenta-

lium leaves little doubt about its species

status. In this case, there are two argu-

ments for treating the presumed ancestor

and the descendants as separate species.

First, according to CAPROTTI (1979: 232),

there is a “typical mutabile” to be distin-

guished from inaequicostata, the latter being

extremely variable. Second, there is no fos-

sil record of either species from the Ta-

bianian (Lower Pliocene). Thus, there is a

gap between the latest mutabile fossils from

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from the Miocene on.

ACKNOWLEDGEMENTS

I am indepted to José Templado,

Museo Nacional de Ciencias Naturales,

for making the material available and for

the information on station data. I also

thank Luitfried Salvini-Plawen for his

comments on the manuscript. An anony-

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111

ad mA d A ds m NE: A” 7 rr EM

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O Sociedad Española de Malacología —_—_—_—_——— Iberus, 15 (1): 113-123, 1997

Análisis parasitológico de gasterópodos acuáticos del delta

del Llobregat (Barcelona). Estadios larvarios de trematodos

digénidos

Parasitological study on aquatic gastropods from the Llobregat delta

(Barcelona). Larval stages of digenetic trematodes

Mercedes VILLA, Isabel MONTOLIU, Mercedes GRACENEA y Olga GON-

ZÁLEZ-MORENO*

Recibido el 8-VIII-1996. Aceptado el 20-XT-1996

RESUMEN

Se ha estudiado, durante los años 1990 a 1995, el índice de parasitación por estadios

larvarios de trematodos digénidos de 6. 184 gasterópodos acuáticos recolectados en el

delta del Llobregat, pertenecientes a 4 especies, 2 prosobranquios (Hydrobiidae) y 2 pul-

monados (Ellobiidae, Physidae). El hidróbido Mercuria confusa [Frauenfeld, 1863) ha

resultado ser el único parasitado por digénidos, mostrando una prevalencia de infestación

del 3,22% (168 positivos sobre 5.219 analizados). Se han detectado hasta 5 especies

de trematodos en estadio larvario, las cuales siguen dos modalidades de ciclo biológico:

a) emisión de cercarias (xifidiocercarias Lecithodendriidae Odhner, 1910 y Microphalli-

dae Travassos, 1920, cercarias inermes Notocotylidae Lúhe, 1957; b) ausencia de emer-

gencia cercariana (Microphallidae y Heterophyidae -Leiper, 1909- Odhner, 1914). La

alta densidad poblacional de M. confusa y su susceptibilidad de ser parasitado por lar-

vas de diversas especies de digénidos revelan su importancia en el mantenimiento de los

ciclos biológicos de trematodos, ratificando el significativo papel que ejercen los proso-

branquios como hospedadores intermediarios específicos en ambientes palustres.

ABSTRACT

Aquatic gastropods from the Llobregat delta were studied in order to detect infection pre-

valence by larval digenetic trematodes in the period 1990-1995. Specimens analysed

(6.194) belonged to 4 species, 2 prosobranchs (Hydrobiidae) and 2 pulmonates (Ellobii-

dae, Physidae), the hydrobiid M. confusa (Frauenfeld, 1863) being the only infected. Lar-

vae of 5 trematode species were detected following two life cycle modalities: a) cercarial

emergence (xifidiocercariae Lecithodendriidae Odhner, 1910 and Microphallidae Travas-

sos, 1920, unarmed cercariae Notocotylidae Lúhe, 1957); b)without cercarial emergence

(Microphallidae and Heterophyidae -Leiper, 1909- Odhner, 1914). The high density of M.

confusa population and its susceptibility to infection by several digenetic species show it to

be a specific intermediate host in the life cycle of these parasites, thus confirming the signi-

ficant role of prosobranchs in the maintenance of trematode life cycles in deltaic zones.

PALABRAS CLAVE: delta del Llobregat, Prosobranchia, Hydrobiidae, M. confusa, estadios larvarios de digéni-

dos, Lecithodendriidae, Microphallidae, Notocotylidae, Heterophyidae.

KEY WORDS: Llobregat delta, Prosobranchia, Hydrobiidae, M. confusa, larval stages of Digenea,

Lecithodendriidae, Microphallidae, Notocotylidae, Heterophyidae.

* Laboratorio de Parasitología, Facultad de Farmacia, Universidad de Barcelona, Avda. Diagonal, s/n, 08028

Barcelona.

113

Iberus, 15 (1), 1997

INTRODUCCIÓN

Las marismas del delta del Llobregat

acogen una amplia variedad de gasteró-

podos acuáticos y terrestres, distribui-

dos en las lagunas, canales y terrenos

colindantes, especies que se extienden

por toda la planicie litoral en abundan-

tes poblaciones y cuya distribución y

frecuencia ha quedado reflejada en el

estudio de ALTIMIRA (1969), autor que

describe hasta 79 especies presentes en

diversos hábitats deltaicos, destacando

asimismo los tratados malacológicos

generales elaborados por Haas (1929),

VIDAL ABARCA Y SUÁREZ (1985) y BECH

(1990).

La importancia parasitológica de

este grupo zoológico de invertebrados

reside en el destacado papel que osten-

tan sus especies como primeros hospe-

dadores intermediarios específicos,

albergantes de esporocistos, redias y

cercarias, de prácticamente todas las

especies parásitas de trematodos digéni-

dos de ciclos biológicos conocidos;

pudiendo representar un doble papel en

los ciclos abreviados, al intervenir como

primeros y segundos hospedadores

intermediarios simultáneamente, alber-

gantes además de metacercarias.

El proceso natural de eutrofización

existente en este tipo de ambiente

lagunar, caracterizado por la presencia

constante de agua salobre, rica en

materia orgánica, y la descomposición

de la vegetación helofítica, constituye el

factor primordial para el asentamiento

de moluscos eurihalinos. Otros peque-

ños invertebrados que conviven estre-

chamente con los anteriores (crustáceos

e insectos) pueden actuar como segun-

dos hospedadores intermediarios de

estos digénidos, sirviendo, asimismo,

como fuente de alimentación de la fauna

vertebrada (aves, micromamíferos,

peces), hospedadores de las formas

adultas del parásito.

El conocimiento de la biología de los

digénidos en este tipo de ambiente

palustre requiere un seguimiento previo

de las especies hospedadoras parasita-

das, de los biotopos que frecuentan y de

su comportamiento. En este sentido,

114

cabe destacar los numerosos trabajos

faunístico-ecológicos realizados en el

delta del Ebro, basados en las especies

vermidianas y sus hospedadores micro-

mamíferos (FELIU, TORRES, GÁLLEGO,

GOSÁLBEZ Y VENTURA, 1985; GRACENEA,

FELIU, MONTOLIU, TORRES Y GÁLLEGO,

1987; FELIU, TORRES, GRACENEA Y MON-

TOLIU, 1990), trabajos que han servido

como punto de referencia para el

estudio de los ciclos biológicos, princi-

palmente acuáticos, de digénidos que

tienen lugar en dicho enclave (MONTO-

LIU, GRACENEA, VILLA Y GONZÁLEZ-

MORENO, 1991). De la misma forma, los

llevados a cabo en pequeños mamíferos

del delta del Llobregat, menos numero-

sos y caracterizados por recoger una

fauna parasitaria cualitativamente y

cuantitativamente más pobre (GRACE-

NEA Y MONTOLIU, 1992; GRACENEA,

MONTOLIU Y DEBLOCK, 1993), han sido

ampliados con el estudio parasitológico

de diversas especies de moluscos, hos-

pedadoras de larvas de digénidos tanto

de ciclo acuático (MONTOLIU, GRACENEA

Y DEBLOCK, 1992), como de ciclo terres-

tre (GONZÁLEZ-MORENO, GRACENEA,

MONTOLIU Y VILLA, 1994). Continuando

en esta línea de investigación sobre

ciclos biológicos de trematodos parási-

tos, este trabajo muestra la diversidad

existente en la helmintofauna larvaria

asociada a gasterópodos acuáticos adap-

tados a hábitats deltaicos.

MATERIAL Y MÉTODOS

Recolección de caracoles y manteni-

miento en el laboratorio: El enclave

prospectado, ya señalado anteriormente

por GONZÁLEZ-MORENO ET AL. (1994),

comprende los márgenes de la laguna

de La Ricarda (Biotopo A) y terrenos

colindantes del margen derecho parcial-

mente inundados por las aguas (Biotopo

B) (ver Figura 1). Ambos puntos de

muestreo constituyen el hábitat de asen-

tamiento de diversas comunidades de

moluscos, especies que son capaces de

soportar la intensa fluctuación estacio-

VILLA ET A4L.: Larvas de tremátodos digénidos en gasterópodos acuáticos del Llobregat

BARCELONA

b 1,5 km

RÍO LLOBREGAT

PRAT DEL

LLOBREGAT

AEROPUERTO

Figura 1. Situación geográfica de la zona prospectada en el delta del Llobregat. A: laguna de La

Ricarda. B: localización de los biotopos estudiados.

Figure 1. Geographic situation of the prospected areas in the Llobregat delta. A: La Ricarda lagoon. B:

localization of the study areas.

nal del enclave, con aporte constante de

agua dulce procedente de los canales y

del acuífero superficial e infiltración de

agua marina.

Biotopo A: la vegetación natural pre-

dominante en esta comunidad helofítica

son los cañizares que bordean el margen

de la laguna, con una especie predomi-

nante, el cañizo (Phragmites australis),

cuyos tallos parcialmente sumergidos

permiten el ascenso de los gasterópodos

hacia la superficie. Agua de salinidad

variable (3-8%0)y pH entre 7 y 8.

Biotopo B: constituido por zonas ane-

gadas, poco profundas, cercanas al

margen derecho de la laguna, alimenta-

das esencialmente por los canales de

desagúe y las lluvias; con predominio

de la vegetación helofítica asociada a

especies halófilas. Junto a las gramíneas

(Phragmites) se encuentra la espadaña

(Typha angustifolia, T. latifolia), el esparto

(Spartina juncea), así como una comuni-

dad bentónica (Enteromorpha, Ulva,

Chara); dado el menor aporte de agua

marina la salinidad es menor (2-5%o) y

pH entre 7 y 8.

Las especies de caracoles recolecta-

das en ambas zonas de muestreo han

sido seis: 2 prosobranquios, Mercuria

confusa (Frauenfeld, 1863) y Potamopyr-

gus jenkinsi (Smith, 1889) (Hydrobiidae)

y 2 pulmonados, Ovatella (Myosotella)

myosotis (Draparnaud, 1801) (Ellobiidae)

y Physa acuta (Draparnaud, 1805) (Physi-

dae). En la Tabla I queda reflejado el

número de caracoles de cada especie

estudiado, el biotopo de prospección y

la época de recolección.

El estudio ha abarcado un periodo

de cinco años (1990-95) con muestreos

en primavera y otoño. Para ello, fueron

utilizados tamices metálicos con los que

se procedía a barrer el fondo fangoso o

bien se practicaban pequeñas sacudidas

de las partes sumergidas de la vegeta-

ción acuática. Una vez en el laboratorio

los moluscos eran estabulados, reprodu-

ciendo las características específicas de

cada zona de captura: con una salinidad

115

Iberus, 15 (1), 1997

Tabla I. Especies de gasterópodos estudiadas: distribución del número de ejemplares según época de

recolección y biotopo prospectado (A, B).

Table I. Gastropod species analysed: specimen distribution in the prospected biotopes (A, B) and annual

variation.

1990 1991 1992

A A A

Prosobranchia

Mercuria confusa IIS O 27]

Potamopyrgus ¡jenkinsi 12 21 9

Pulmonata

Ovatella ([Myosotella) myosotis 25 6 17

Physa acuta 4 40

y pH adecuados, dieta constituida por

lechuga seca y alimento para peces.

Detección y aislamiento de formas

larvarias de digénidos: La detección de

una posible emisión de cercarias al

medio externo se efectuó disponiendo

individualmente los gasterópodos en

pocillos de placa de cultivo celular con-

teniendo agua del biotopo y observán-

dolos posteriormente bajo la lupa en

búsqueda de cercarias nadando libre-

mente. Para la detección de esporocis-

tos, redias y metacercarias, se procedió a

la disección de los caracoles y a la obser-

vación de todos sus órganos.

Técnicas microscópicas: Las larvas

fueron inicialmente estudiadas in vivo

con el microscopio óptico y con la ayuda

del colorante vital rojo neutro, proce-

diéndose posteriormente a la fijación

con el líquido de Bouin, tinción con

carmín alumínico y tras cuidadosa des-

hidratación se montaron con Bálsamo

del Canadá.

RESULTADOS

El estudio de 5.219 ejemplares de M.

confusa ha puesto de manifiesto la capa-

cidad de este prosobranquio, el único

gasterópodo que ha resultado estar

parasitado, para albergar diferentes

formas larvarias de digénidos, en total

cinco especies caracterizadas por su

116

1993: 1994 1995 Total

A B ASNO B

E O MO CA O NO

224 40 7

36 : : E Aa

121 OA 1 589

ciclo biológico acuático. Todas las infes-

taciones por digénidos han mostrado

invasión a nivel del complejo glándula

digestiva-gónada, como hábitat de elec-

ción para la evolución de las larvas.

Las especies de digénidos detecta-

das, pertenecientes a 4 familias, presen-

tan dos modalidades de ciclos biológi-

cos (Fig. 2). En la primera de ellas, es

característica la emisión de cercarias al

medio externo, bien sea en ciclos trihete-

roxenos (con tres hospedadores) o dihe-

teroxenos (con dos hospedadores). En la

segunda modalidad, ciclo de tipo abre-

viado, el gasterópodo se comporta como

primer y segundo hospedador interme-

diario simultáneamente, no habiendo

emisión de cercarias.

I. Emisión de cercarias

I. a. Ciclos triheteroxenos

Cercarias Lecithodendriidae. Lecit-

hodendriidae gen. sp.: Cercarias xifi-

diocercas (con estilete), distomas, pro-

vistas de una cola recta más estrecha

que el cuerpo (leptocercas) y virguladas

(cuerpo: 150 x 90 um). Originadas en

esporocistos sacciformes (100-350 x 50-

100 um) provistos de poro de salida

musculoso a través del cual las cercarias

emergen según un patrón de emisión

predominantemente nocturno.

La morfoanatomía de estas cercarias

se ajusta a la descrita para la familia

Lecithodendriidae (YAMAGUTI, 1975;

SCHELL, 1985), destacando su afinidad

con xifidiocercarias Lecithodendriidae

VILLA ET 4Lz.: Larvas de tremátodos digénidos en gasterópodos acuáticos del Llobregat

o 'O) enquistamiento O) anfibios,

Cercarias me a e reptiles o

EMISIÓN Lecithodendriidae : 22H. 1 quirópteros

insectos ( 19) H'D

DE CERCARIAS (H. D.)

E SLCALIAS enquistamiento O) insectivoros

M. feliui ——Y» en crustáceos, 5 (C. russula)

E) anfípodos e (H. D.)

isópodos (2? H. L)

CICLOS TRIHETEROXENOS

CICLOS DIHETEROXENOS

| — Catia enquistamiento O) Aves

a : ——PY» en plantas = aos

E Notocotylidae p y > acuáticas

conchas del caracol (H. D.)

iS enquistamiento 2) o

M. fusiformis bip> q = =D> acuáticas

: : en esporocistos HD

NO EMISIÓN (Microphallidae) (H. D.)

peces o

reptiles

(H. D.)

DE CERCARIAS cercarias $ enquistamiento

Heterophyidae en el caracol

Figura 2. Ciclos biológicos de los digénidos detectados en Mercuria confusa. (—): fases conocidas.

(=>): fases presumibles. 2” H. L.: segundo hospedador intermediario. H. D.: hospedador definitivo.

1: penetración activa. 2: penetración pasiva. 3: huevos en heces del H. D.

Figure 2. Life cycles of digeneans detected in Mercuria confusa. (—): known phases. (==): likely phases.

2'H. L.: second intermediate host. H. D.: definitive host. 1: active penetration. 2: pasive penetration. 3:

eggs in definitive host feces.

detectadas en otros biotopos deltaicos emergencia cercariana se produce predo-

geográficamente próximos, el delta del minantemente en horas crepusculares.

Ebro (MONTOLIU ET AL., 1991) y costa La realización experimental del ciclo

francesa mediterránea (DEBLOCK, 1980). biológico, descrito sucintamente por los

Las cercarias probablemente infestan autores de la especie, ha permitido su

larvas de insectos (segundos hospeda- determinación sistemática. Las cercarias

dores intermediarios, en los que se infestan activamente a crustáceos anfí-

forma la metacercaria), desarrollándose podos e isópodos (segundos intermedia-

el adulto probablemente en anfibios y rios), evolucionando a metacercarias

reptiles (LLUCH, ROCA Y NAVARRO, 1986) enquistadas, los cuales son depredados

o en quirópteros (ESTEBAN, OLTRA- por el hospedador definitivo, el insectí-

FERRERO, BOTELLA Y GRANEL, 1993) por voro Crocidura russula (Hermann, 1780)

depredación, aunque sin descartar su en el delta del Llobregat, en el que se

posible presencia en micromamíferos originan los adultos a nivel intestinal

(GRACENEA ET AL., 1987). (GRACENEA Y MONTOLIU, 1992; GRACE-

NEA ET AL., 1993).

Cercarias Microphallidae. Mari-

trema feliui Gracenea, Montoliu et I. b. Ciclos diheteroxenos

Deblock, 1993: Xifidiocercarias, monos- Cercarias Notocotylidae. Notocoty-

tomas, anentéricas y leptocercas (cuerpo: lidae gen. sp.: Cercarias de gran tamaño

120 x 68 um), de fórmula excretora (cuerpo: 600-800 ym x 370 um) y gran

2((2+2)+(2+2)= 16 solenocitos. Se origi- opacidad, inermes (ausencia de estilete),

nan en esporocistos de aspecto sacci- oftalmocercas (con manchas oculares),

forme e irregular, (166-430 x 140-180 monostomas y leptocercas. Se originan

um), provistos de poro de salida. La en redias de aspecto fusiforme, provis-

Iberus, 15 (1), 1997

tas de faringe subterminal y ciego intes-

tinal de gran volumen. Las cercarias

acaban de madurar fuera de las redias,

emergiendo al medio externo y enquis-

tándose rápidamente en la vegetación o

sobre la misma concha del caracol, evo-

lucionando a metacercarias (cuerpo: 736

x 138 um) confinadas en quistes hemies-

féricos (diámetro, 150-300 um).

La morfología de las cercarias y el

ciclo diheteroxeno con enquistamiento

en el medio acuático son propios de la

familia Notocotylidae (YAMAGUTI, 1975;

SCHELL, 1985). Los adultos se desarro-

llan presumiblemente en aves acuáticas

deltaicas (hospedadores definitivos) al

alimentarse éstos de plantas acuáticas o

de moluscos.

II. No emisión de cercarias (ciclos abre-

viados)

Cercarias Microphallidae. Microp-

hallus fusiformis Reimer, 1963: Cerca-

rias rudimentarias (blastocercarias) de

reducidas dimensiones (50-70 x 30-37

ym), inmóviles y constituidas por

células indiferenciadas, sin apéndice

caudal ni esbozos de otras estructuras.

Se forman en esporocistos blanquecinos,

transparentes (200-500 x 150 um) y sac-

ciformes, en los que evolucionan a

metacercarias enquistadas (diámetro,

80-110 x 59-64 um).

La metacercaria desenquistada se

caracteriza por su pequeño tamaño (140-

160 x 60 um) y cuerpo fusiforme muy

espinulado; de carácter progenético, con

testículos y glándulas vitelógenas fun-

cionales; ovario diestro y metratermo

confluyendo a nivel de la pared lateral

del atrio genital La morfología de dichas

metacercarias, muy similar a la del

adulto (MONTOLIU ET AL., 1992). El hos-

pedador definitivo en el delta lo consti-

tuyen probablemente aves anseriformes

(REIMER, 1963).

Cercarias Heterophyidae. Hete-

rophyidae gen. sp.: Cercarias monosto-

mas, leptocercas y oceladas, con órgano

de penetración protráctil (cuerpo: 100-

120 x 50-60 ym) y sistema excretor pro-

visto de glándula post-vesical. Evolucio-

nan a partir de redias de aspecto cilín-

drico (158-370 x 52-103 ym), provistas

de faringe musculosa y un ciego corto.

Las cercarias, tras emerger de la redia,

se enquistan dentro del mismo caracol.

Las metacercarias están provistas de

una doble corona de espinas rodeando a

la boca. Son distomas, con testículos

homolaterales, vesícula excretora de

gran tamaño, encontrándose confinadas

en quistes (80x70 um) de doble cubierta.

Las aves acuáticas, peces o anfibios

podrían actuar como hospedadores defi-

nitivos del digénido (YAMAGUTI, 1975).

En la Tabla II se encuentran recopila-

das las prevalencias de infestación en

Mercuria confusa para cada una de las

especies descritas anteriormente.

DISCUSIÓN

Los estudios sobre la helmintofauna

larvaria de digénidos de ciclo acuático

señalan a los moluscos prosobranquios

como los principales hospedadores

intermediarios específicos para las espe-

cies Digenea de ambientes palustres.

Los datos obtenidos en nuestro estudio

han mostrado que el hidróbido Mercuria

confusa interviene como hospedador

intermediario específico de cinco espe-

cies de digénidos deltaicos. Este proso-

branquio ha sido estudiado parasitoló-

(Página derecha). Figura 3. Estadios larvarios de digénidos detectados en Mercuria confusa. A: cer-

caria Lecithodendriidae (rojo neutro). B: blastocercaria y quistes metacercarianos intrasporocísticos

de M. fusiformis. C: metacercaria enquistada Heterophyidae. D: cercaria de M. feliui (rojo neutro).

Escalas 25 um.

(Right page). Figure 3. Digenean larval stages detected ¿in Mercuria confusa. 4: cercaria

Lecithodendriidae (neutral red). B: intrasporocystic blastocercaria and metacercarial cysts of M. fusifor-

mis. C: encysted metacercaria Heterophyidae. D: cercaria of M. feliui (neutral red). Scale bars 25 pm.

118

VILLA ET AL.: Larvas de tremátodos digénidos en gasterópodos acuáticos del Llobregat

119

Iberus, 15 (1), 1997

Tabla II. Parasitación de M. confusa por estadios larvarios de digénidos según épocas de recolección

y biotopo prospectado (A, B). N=n" de especímenes estudiados (por emisión y/o por disección).

%=prevalencia de parasitación.

Table II. Infection of M. confusa with larvae of digenetic trematodes, related to year of collection and

prospected biotopes (A, B). N=number of examined snails (emergence andlor dissection). %=prevalence

of infection.

1990 1991 1992

A A A

% % %

1993 1994 1995 Total

% % % % % %

Cercarias+Esporocistos (o redias) (emisores de cercarias)

N=1215 N=656 N=271 N=989 N=708 N=146 N=1164 N=70 N=5219

Lecithodendriidae gen sp. : 0,40 0,42 - 0,28 - 0,15

Maritrema felivi OASIS ERA SOTA ZONA: - ISAAC O

Notocotylidae gen. sp. - OOO O 0,10

Cercarias+Esporocistos (o redias) + metacercarias (ciclos abreviados)

N=492 N=482 N=33 N=475 N=210 N=32 N=150 N=0 N=1874

Microphallus fusiformis IESDAE2IO : a - : 2,51

Heterophyidae gen. sp. 325 0,85

gicamente con anterioridad en el delta

del Ebro por MONTOLIU ET AL. (1991),

detectándose formas larvarias pertene-

cientes a 7 especies de digénidos. La

ausencia de parasitación en el proso-

branquio Potamopyrgus jenkinsi, en el

que sólo han sido estudiadas hembras

partenogenéticas, podría deberse a un

fenómeno similar al observado por

LiveLY (1989) para otras especies de

Potamopyrgus Stimpson, 1865 en Nueva

Zelanda, en las que se demuestra la

correlación positiva entre las poblacio-

nes sexuadas y la parasitación por

microfálidos, no estando nunca parasi-

tadas las poblaciones que se reproducen

exclusivamente por partenogénesis. En

lo que respecta a las especies de pulmo-

nados estudiadas, si bien éstas no se

encuentraron parasitadas por digénidos,

sí que existen numerosas citas de infes-

taciones por otras especies de digénidos

en ejemplares dulceacuícolas europeos

(MOUAHID Y MONÉ, 1988, entre otras).

La prevalencia de parasitación por

larvas de digénidos mas alta detectada

en M. confusa ha correspondido a la

familia Microphallidae. Los digénidos

de biología conocida de esta familia

incluyen a diversos moluscos proso-

branquios como hospedadores interme-

120

diarios, tanto en las especies triheteroxe-

nas como las diheteroxenas, de ciclo

abreviado. En la primera modalidad,

para las especies de Maritrema Nicoll,

1907 y Microphallus Ward, 1901, son los

prosobranquios pertenecientes a Litto-

rina Ferrusac 1822 (BENJAMIN Y JAMES,

1987; IRwIN, MAGUIRE Y SAVILLE, 1990;

GALAKTIONOV Y BUSTNERS, 1995) y a

Hydrobia Hartmann, 1821 (GARKAVI,

1972; PREVOT Y BARTOLI, 1977; DEBLOCK,

1978; SAVILLE Y IRwIN, 1991) los más fre-

cuentemente citados, y más puntual-

mente los prosobranquios Bythinella

Moquin-Tandon, 1855 (JOURDANE, 1979),

Pseudamnicola Paulucci, 1878 (KULKINA Y

BELYAKOVA, 1983), Cerithium Bruguiere,

1789, Bittium Leach, 1847 (PREVOT,

BARTOLI Y DEBLOCK, 1976; BARTOLI Y

Prevor, 1978) y Cerithidea Swainson,

1840 (ABDUL-SALAM Y SREELATHA, 1991).

El género Microphallus es el que engloba

el mayor número de especies con ciclos

abreviados, interviniendo habitual-

mente especies de Hydrobia y Littorina y

puntualmente de Bittium, incluyéndose

únicamente a especies de Hydrobia para

el género Maritrema (DEBLOCK, 1977;

LAUCKNER, 1984).

En lo que se refiere a la familia Lecit-

hodendriidae, los prosobranquios vuel-

VILLA ET AL.: Larvas de tremátodos digénidos en gasterópodos acuáticos del Llobregat

ven a ser citados frecuentemente en los

trabajos sobre trematodofauna larvaria,

destacando los realizados en especies de

Bithynia Leach, 1818 (YAMAGUTL 1975) y

Amnicola Gould y Haldemanmn, 1841 (Ca-

BLE, 1985). Con respecto a los hospeda-

dores intermediarios de notocotílidos,

cabe señalar a los prosobranquios Hydro-

bia (STUNKARD, 1966; DEBLOCK, 1980), Po-

tamopyrgus (BisseT, 1977), Bithynia (Y A-

MAGUTI, 1975; VASILEV Y KANEvV, 1984),

Littorina (GRANOVICH, MIKHALOVA Y

SERGIEVSKIIL, 1987), así como a especies

de melaníidos (KHALIFA Y EL-NAFFAR,

1979). En los digénidos heterófidos no

son frecuentes los ciclos abreviados

como el que tiene lugar en el delta del

Llobregat, habiéndose citado a una sola

especie de digénido, Metagonimoides ore-

gonensis Price, 1931, la cual infesta a di-

versas especies de prosobranquios pleu-

rocéridos, únicos hospedadores interme-

diarios del ciclo (YAMAGUTI, 1975).

El análisis cuantitativo de la parasita-

ción por larvas de M. confusa muestra

diferencias en las prevalencias para cada

una de las especies de digénidos halla-

dos (Tabla II). Los índices totales más

elevados mostrados por los microfálidos

(Maritrema feliui - 1,76 %; Microphallus

fusiformis - 2,51 %) parecen ajustarse a los

detectados en hidróbidos que habitan

zonas geográficas litorales próximas al

delta del Llobregat. MONTOLIU ET AL.

(1991) muestran para M. confusa en el

delta del Ebro unos niveles de parasita-

ción para Maritrema sp. (1,36 %) muy

similares a las del presente trabajo. Asi-

“mismo, en el estudio realizado por

DEBLOCK (1978) se observan índices de

parasitación por microfálidos en especies

de Hydrobia que oscilan entre el 2,5 y

0,5% en el litoral atlántico y entre el 6 y

0,5% en el litoral mediterráneo. En otros

prosobranquios litorales también muy

estudiados del género Littorina, la preva-

lencia de estadios larvarios Microphalli-

dae es muy alta, del orden del 23,8% en

la costa del Mar Báltico (LAUCKNER,

1984) y de hasta el 40-50% en las costas

soviéticas (SERGIEVSKIL, 1985).

La baja prevalencia por lecitodéndri-

dos en el Llobregat no parece ajustarse a

las tasas de infestación observadas en

M. confusa del delta del Ebro, en el que

se alcanzan índices elevados (5,83%)

(datos no publicados). Los bajos índices

de infestación detectados en el resto de

digénidos sí parecen coincidir con los

obtenidos para este hidróbido en el

Ebro, detectándose el 0,27% para cerca-

rias Heterophyidae y el 0,20% para cer-

carias Notocotylidae, y con los de

DEBLOCK (1978) en las costas francesas

para especies de Hydrobia, 0,07-1,67%

para heterófidos y 0,20-0,43% para noto-

cotílidos.

En cuanto a la dinámica del parasi-

tismo, las fluctuaciones temporales

observadas en el hidróbido no parecen

guardar relación directa con su densi-

dad poblacional, relativamente cons-

tante en todas las prospecciones realiza-

das, o con el biotopo de prospección.

Ello podría estar relacionado con el

comportamiento de los hospedadores

definitivos como factor de máxima

influencia en dichas variaciones. Mari-

trema feliui es la única especie de digé-

nido que parece mantenerse relativa-

mente constante en el tiempo, hecho que

puede explicarse por la presencia

regular del insectívoro Crocidura russula

en el delta (GRACENEA Y MONTOLIU,

1992). En cuanto a Notocotylidae gen.

sp. y Microphallus fusiformis, potencial-

mente parásitos de aves deltaicas, su

aparición más esporádica en los caraco-

les estaría condicionada por el carácter

migratorio de sus hospedadores defini-

tivos.

AGRADECIMIENTOS

Los autores agradecen a D. Serge

Gofas, malacólogo del Laboratoire de

Biologie des Invertébrés Marins et Mala-

cologie du Muséum National d'Histoire

Naturelle de Paris, la determinación de

los moluscos prosobranquios. Asimismo,

agradecen a D. Manuel Bertrand Vergés,

Presidente de la sociedad Ebysa propie-

taria de la finca prospectada, su consen-

timiento para la recolección del material

malacológico. Este trabajo ha sido finan-

ciado por el Proyecto PB 92-0517 de la

DGICYT.

121

Iberus, 15 (1), 1997

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O Sociedad Española de Malacología ——_—_——— Iberus, 15 (1): 125-130, 1997

On a floating egg mass of the diamond shaped squid 7)»y-

sanoteuthis rhombus (Cephalopoda: Thysanoteuthidae) in

the western Mediterranean

Observaciones sobre una puesta pelágica del calamar losange 7»y-

sanoteuthis rhombus (Cephalopoda, Thysanoteuthidae) hallada en

el Mediterráneo occidental

Angel GUERRA and Francisco ROCHA*

Recibido el 14-X1-1996. Aceptado el 3-XII-1996

ABSTRACT

This is the second record of a floating egg mass of the diamond shaped squid Thysanoteu-

this rhombus in the Mediterranean (37* 11.85” N - 1? 31.15 E). The first one was obser-

ved at the Strait of Messina in 1929. The egg mass was a dense, resilient oblong cylinder

with rounded tips appoximately 100 cm in length and about 20 cm in diameter. From a

small sample, egg capsules and paralarvae (1.85+0.08 mm MIL) are described. Some

complementary characters about this species paralarvae, such as the “arm formulae, the

presence of an incipient swimming keel-like shaped membrane on some arms, and the

mantle chromatophore pattern should assist in their identification.

RESUMEN

En este trabajo se informa sobre el segundo hallazgo de una puesta pelágica del calamar

losange Thysanoteuthis rhombus en el Mar Mediterráneo (37? 11,85" N - 1? 31,15 E). La

primera se observó en el estrecho de Mesina en 1929. La masa de huevos consistía en un

cilindro oblongo con los bordes romos, denso y elástico, de unos 100 cm de longitud y

20 cm de diámetro. A partir de una muestra pequeña que se pudo obtener se describen

las cápsulas ovigeras y las paralarvas (1,85+0,08 mm ML). Se proporcionan algunos

caracteres complementarios útiles para la identificación de estas paralarvas: la formula

braquial, la presencia de una carena natatoria incipiente aquillada en algunos brazos y

el patrón de cromatóforos en el manto.

KEY WORDS: Cephalopoda, Thysanoteuthis rhombus, egg mass, paralarvae, Mediterranean Sea.

PALABRAS CLAVE: Cephalopoda, Thysanoteuthis rhombus, puesta, paralarvas, mar Mediterráneo.

INTRODUCTION

The diamond shaped squid Thysano- partially subtropical waters of the World

teuthis rhombus Troschel, 1857 is an epi- Ocean including the Mediterranean,

pelagic inhabitant of warm tropical and often occurring in pairs or small schools

* Instituto de Investigaciones Marinas, C/ Eduardo Cabello 6, 36208 Vigo, España.

Iberus, 15 (1), 1997

(NISHIMURA, 1966; CLARKE, 1966; NIG-

MATULLIN, ARKHIPKIN AND SABIROV,

1995). It is a large oegopsid squid which

reaches up to 85 cm in mantle length

(ML) and 24 kg in body weight (NIGMA-

TULLIN ET AL., 1995). T. rhombus is one of

the fastest-growing squid: in approxi-

mately one year, it reaches its maximum

ML (NIGMATULLIN ET AL., 1995). This

species has high potential fecundity (up

to 4.8 million oocytes), but a rather

small maximum volume of oviducts (up

to 140,000 eggs) and egg masses (35,000

to 75,000), which suggest that T. rhombus

is an intermittent spawner with multiple

filling and evacuation of oviducts (NIG-

MATULLIN ET AL., 1991; NIGMATULLIN ET

AL., 1995).

T. rhombus is one of the few oegopsid

cephalopods in which the spawn is

known. Until recently, a total of 21 egg

masses had been observed. All were

found drifting in the surface water layer

of the tropical Atlantic, northwest and

southeast Pacific and the Mediterranean

(review in SABIROV, ARKHIPKIN, TSYGAN-

KOV AND SHCHETINNIKOV, 1987). Egg

masses of this species are gelatinous, sau-

sage-shaped, 60-180 cm long by 10-30 cm

diameter; containing a double spirally

arranged row of eggs embedded in the

surface layer of the mass (MISAKI AND

OKUTANI, 1976; SUZUKI, MISAKI AND

OKUTANI, 1979). The egg mass was pho-

tographed in natural environment for the

first time by SUZUKI ET AL. (1979). The

first and unique reference to an egg mass

of T. rhombus in the Mediterranean (Strait

of Messina) was given by SANZO (1929).

Although the occurrence of this

species in the Mediterranean is rare

(MORALES, 1981; BIaGL, 1982; MANGOLD

AND BOLETZKY, 1988) it seems that its

presence is increasingly frequent as by-

catch in some pelagic fisheries, particu-

larly near the coast (EZZEDDINE-NAJAL,

1996). A pair of animals, male and

female, were observed by divers in a

submarine cave off the coast of Almeria

(Southeast Spain) relatively near the

place where the sample reported in this

paper was collected (GUERRA, 1992).

This paper deals with the second

record of the egg mass of T. rhombus in

126

the Mediterranean after 67 years. A des-

cription of the planktonic paralarvae is

given, emphasising several characteris-

tics which may be used to identify the

early stages of the species.

MATERIAL AND METHODS

The egg mass was discovered by the

vessel “Toftevaag” at 08.27 h on August

o MIS ARAS NS ME SS 18

the western Mediterranean (Fig. 1). The

reported egg mass was accompanied by .

other drifted pleuston, such as jelly-fish.

The whole mass was so loose that it

could not be taken out of the water. But

a sample of gelatinous material contai-

ning 2 eggs in early stage of develop-

ment, 1 embryo within its egg capsule, 2

paralarvae within the egg capsules, and

32 practically fully developed paralar-

vae outside the egg capsules were

caught. This sample was fixed in 5% for-

malin. The identification was made

based on the paralarvae which, alt-

hough still embedded in the external

gelatinous mass, were near hatching.

These paralarvae have similar characte-

ristics to those reported by STEPHEN

(1992). Egg diameter, dorsal mantle

length (ML) and total length (TL) of

each paralarvae were measured using a

dissecting microscope fitted with an

eye-piece graticule.

RESULTS

The egg mass was a dense, resilient,

oblong cylinder with rounded ends (Fig.

2A). The size of the whole mass was

about 100 cm in length and about 20 cm

in diameter. It was observed that the

purple egg capsules lay in two rows,

spirally arranged around the cylinder.

The diameter of the egg capsules with

the well developed embryo ranged from

2.8 to 3.0 mm. The average ML of para-

larvae was 1.85+0.08 mm (n= 30) and its

TL varied between 2.50 and 2.75 mm.

All paralarvae observed had the

head inside the mantle cavity, only

showing the arms and tentacles exter-

GUERRA AND ROCHA: Egg mass of Thysanoteuthis rhombus in Western Mediterranean

Atlantic

Ocean

oVigo

Mind Western

Mediterranean

Strait of _-

Gibraltar

Black Sea

| N Strait of

Messina

Mediterranean

Figure 1. Thysanoteuthis rhombus. Location of the two floating egg masses collected in the

Mediterranean Sea. A: This paper; B: SANZO (1929).

Figura 1. Thysanoteuthis rhombus. Localización geográfica de las dos puestas pelágicas encontradas en

el Mar Mediterráneo. A: Presente estudio; B: SANZO (1929).

nally (Fig. 2B). The mantle is oval, stout,

short and blunt posteriorly. The anterior

margin of the mantle curves inwards in

both dorsal and ventral sides, but is

more pronounced ventrally. This may be

a result of the preservation process

which could had produced the retrac-

tion of the head inside the mantle cavity.

The fins are subterminal, small and

rounded; the fin length 19.5% ML (Figs.

2C, D). The paralarvae have broadly

separated, slightly protruding eyes, and

funnel locking cartilage (Fig. 2E) with a

short, broad, transverse groove and a

long, relatively wide, longitudinal

groove (sideways Fshaped). The tenta-

cles are short (about 33% of the ML),

stouter and slightly longer than the

longest arm (III); the I and IV pairs of

arms are rudimentaries. Brachial formu-

lae IMI>II>I=IV. Both, tentacles and deve-

loped arms, with small suckers, pro-

bably arranged in two rows. On arms II

and III, an incipient swimming keel-like

shaped membrane was present. Trabe-

culate protective membrane was absent

in arms and tentacles.

The paralarvae show two types of

chromatophores: a) large and pale-ochre

chromatophores densely concentrated

on dorsal, lateral and ventral mantle

sides; and b) small, subtriangular dark-

red chromatophores arranged in a single

row around the anterior margin of the

mantle. There is a light area between

both types of chromatophores. Slight

chromatophores were observed on the

dorsal and ventral sides of the head, the

tentacles and the arms. The fins lacked

chromatophores.

DISCUSSION

The egg mass reported was captured

near the surface in a zone were the in-

flow of Atlantic water into the Medite-

rranean is high due to the proximity of

the Strait of Gibraltar. The water move-

ments through this bottleneck are gover-

ned by an inflow of surface water into

the Mediterranean, and a countercurrent

of lesser volume carrying water of hig-

her salinity into the Atlantic (MANGOLD

AND BOLETZKY, 1988). The egg mass co-

llected by SANZO (1929) was in the Strait

of Messina where there are strong cu-

rrents. Elsewhere egg masses of Thysa-

noteuthis rhombus occurred in regions

with strong warm currents such as Ku-

roshio, Perú countercurrent and the

Equatorial countercurrent (YAMAMOTO

127

Iberus, 15 (1), 1997

ARS

añ Pe

Figure 2. Thysanoteuthis rhombus. Floating egg mass and paralarvae. A: egg mass; B: Egg with non-

developed embryo and paralarvae within the egg capsule; C: Funnel locking-cartilage of a newly hatched,

1.85 mm ML; D: Dorsal view of a newly hatched, 1.85 mm ML; E: Ventral view of the same specimen. *

Figura 2. Yhysanoteuthis rhombus. Puesta pelágica y paralarva. A: Puesta pelágica; B: Huevo con embrión muy

poco desarrollado y paralarva dentro de la cápsula ovígera; C: Cartilago de cierre en el sifón de un recién nacido

de 1,85 mm ML; D: Visión dorsal de un recién nacido de 1,85 mm ML; E: Visión ventral del mismo ejemplar.

128

GUERRA AND ROCHA: Egg mass of Thysanoteuthis rhombus in Western Mediterranean

AND OKUTANI, 1975; NIGMATULLIN ET

AL., 1995). Therefore, as in the Atlantic

and the Pacific Oceans, in the Medite-

rranean the species seems to spawn in

waters with strong currents.

NIGMATULLIN ET AL. (1995) indicated

that T. rhombus spawns throughout the

year in tropical waters, but during the

warm season (summer and early autumn)

in peripheral regions such as in the Me-

diterranean, which agree with the date

when the egg mass reported was observed.

The egg mass in the report had a

shape and a size which coincide with

those given for the other egg masses

illustrated (SANZO, 1929; MISAKI AND

OKUTANI, 1976) and photographed

(SUZUKI ET AL., 1979).

Considering the dimension of this

egg mass, the diameter of the egg capsu-

les measured and calculating the surface

of the egg mass as a cylinder (6,280 cm?),

an estimation gives a figure of about

66,800 eggs. This amount coincides with

the total number of eggs in each egg

mass calculated by SABIROV ET AL. (1987)

which ranged from 32,000 to 76,000 eggs.

The embryo and paralarvae found

have sizes, shapes and characters which

BIBLIOOGRAPHY

BIAGI, V., 1982. Sul rinvenimento di un gio-

vane esemplare di Thysanoteuthis rhombus

Troschel (Cephalopoda - Teuthoidea) in

acque Elbane. Bollettino Malacologico, 18 (7-

8): 137-144.

CLARKE, M. R., 1966. A review of the systema-

tics and ecology of oceanic squids. Advances

in Marine Biology, 4: 91-300.

EZZEDDINE-NAJAL, S., 1996, On the presence of

a new species of cephalopod Thysanoteuthis

rhombus Troschel, 1857 on the north and

south coasts of Tunisia. IV International Sym-

posium on Cephalopods - Present and Past. Gra-

nada, Spain: 64.

GUERRA, A., 1992. Mollusca, Cephalopoda. In

Ramos, M. A,, et al. (Eds.), Fauna Ibérica,

Vol. 1. Museo Nacional de Ciencias Natura-

les, CSIC, Madrid, 327 pp.

IsseL, R., 1920. Primo contributo alla conos-

cenza dello sviluppo nei cefalopodi medite-

rranei (Ihysanoteuthis - Chiroteuthis - Gali-

teuthis). Memorie Reale Comitato Talassogra-

fico Italiano, 73: 1-19.

largely coincide with those reported by

IsseL (1920), SANZO (1929) YAMAMOTO AND

OKUTANI (1975) and MISAKI AND OKUTANI

(1976) and summarised by CLARKE (1966)

and STEPHEN (1992). Some complemen-

tary information about this species para-

larvae, such as the arm formulae, the pre-

sence of an incipient swimming keel-like

membrane on some arms, and the mantle

chromatophore pattern should assist in

their identification.

ACKNOWLEDGEMENTS

We are particularly grateful to Ricardo

Sagarminaga and Ana Cañadas, owners

of the Vessel “Toftevaag” and promoters of

the “Alnitak” project for the specimen

collection. We thank Diego Moreno of the

University of Almería for valuable preli-

minary comments on the sample and

Alfredo López “Tokio” for drawing the

egg mass and paralarvae figures. We are

grately indebted to Prof. T. Okutani, Dr.

P. G. Rodhouse and Dr. A. F. González for

valuable suggestions on the manuscripts.

Our thanks also to lan Emmett for revie-

wing the English text.

MANGOLD, K. AND BOLETZKY, S. v., 1988. Me-

diterranean cephalopod fauna. In Clarke,

M. R. and Trueman, E. R. (Eds.), Vol. 12:

The Mollusca, Paleontology and neontology of

cephalopods. Academic Press, San Diego:

s19390)

MisaKI, H. AND OKUTANI, T., 1976. Studies on

early life history of decapodan Mollusca -

VI. An evidence of spawning of an oceanic

squid, Thysanoteuthis rhombus Troschel, in

the japanese waters. Venus, The Japanese Jour-

nal of Malacology, 35: 211-213.

MORALES, E., 1981. Presencia de Thysanoteu-

this rhombus Troschel, en el puerto de Ma-

hon (Menorca). Investigación Pesquera, 45:

17-20.

NIGMATULLIN, CH. M., ARKHIPKIN, A. l. ANDSA-

BIROV, R. M,, 1991. Structure of the repro-

ductive system of the squid Thysanoteuthis

rhombus (Cephalopoda: Oegopsida). Journal

of Zoology, London, 224: 271-283.

129

Iberus, 15 (1), 1997

NIGMATULLIN, CH. M., ARKHIPKIN, A. Í. ANDSA-

BIROV, R. M., 1995. Age, growth and repro-

ductive biology of diamond-shaped squid

Thysanoteuthis rhombus (Oegopsida: Thysa-

noteuthidae). Marine Ecolology Progress Se-

ries, 124: 73-87.

NISHIMURA, S., 1966. Notes on the occurence and

biology of the oceanic squid, Thysanoteuthis

rhombus Troschel, in Japan. Publications of the

Seto Marine Biological Laboratory, 13 (4): 327-

349.

SABIROV, R. M., ARKHIPKIN, A. I., TSYGANKOV,

YU AND SHCHETINNIKOV, A. S., 1987. Egg la-

ying and embrional development of dia-

mond-shaped squid Thysanoteuthis rhombus

(Oegopsida, Thysanoteuthidae). Zoological

Journal USSR, 66 (8): 1155-1163.

SANZO, L., 1929. Nidamento pelagico, uova elarve

di Thysanoteuthis rhombus Troschel. Memorie

Reale Comitato Talassografico Italiano, 161: 1-10.

130

STEPHEN, S. J., 1992. Family Thysanoteuthidae

Keferstein, 1866. In Sweeney, M. J. et al. (Eds.),

“Larval” and juvenile cephalopods: a ma-

nual for their identification. Smithsonian Con-

tributions to Zoology, 513: 121-123.

SUZUKI, S., MISAKI, H. AND OKUTANI, T., 1979.

Studies on early life history of decapodan

Mollusca - VILA supplementary note on flo-

ating egg mass of Thysanoteuthis rhombus

Troschel in Japan - The first underwater pho-

tography. Venus, The Japanese Journal of Ma-

lacology, 38 (2): 153-155.

YAMAMOTO, K. AND OKUTANI, T., 1975. Studies

on early life history of decapodan Mollusca

- V. Systematics and distribution of epipela-

gic larvae of decapod cephalopods in the

south-western waters of Japan during the

summer in 1970. Bulletin Tokai Regional Fis-

hery Research Laboratory, 83: 45-96.

O Sociedad Española de Malacología ——_—_—_—_——— Iberus, 15 (1): 131-138, 1997

Ontogenetic variation of statolith shape in the short-finned

squid lex coindetíz (Mollusca, Cephalopoda)

Variacion ontogénica del estatolito de la pota /llex coindetii

(Mollusca, Cephalopoda)

Ángel E GONZÁLEZ and Ángel GUERRA*

Recibido el 27-X1-1996. Aceptado el 10-11-1997

ABSTRACT

Changes in statolith morphology of Illex coindetii are described from specimens ranging

from 42 to 379 mm mantle length obtained during trawling activities in the North-eastern

Atlantic. The growth of the statolith was differentiated in five developmental stages. lt has

been observed that the wing of the statolith has two different growth patterns, from the

ventral zone to the dorsal one and viceversa. The rostral angle of the statolith varied

during its ontogenetic growth from an obtuse angle to a 90? angle. The dorsal zone of the

statolith is here named as dorsal dome, and the lateral and ventral zone the lateral dome.

RESUMEN

En este trabajo se describen los cambios en la morfología del estatolito de /llex coindeti.

El estudio se llevo a cabo mediante el análisis de 341 ejemplares con longitudes del

manto comprendidas entre 42 y 379 mm. Estos animales se capturaron en la pesquería

de arrastre desarrollada en el Atlántico noreste. El crecimiento del estatolito fue diferen-

ciado en cinco estadíos de desarrollo. Se observaron dos patrones diferentes de creci-

miento del ala del estatolito, desde la parte ventral a la dorsal y viceversa. El angulo ros-

tral varía en el crecimiento ontogénico del animal, desde un ángulo marcadamente obtuso

a un ángulo recto. El crecimiento en longitud máxima y en anchura del estatolito se enlen-

tece al llegar a la maduración de los animales. Se propone una variación en la nomencla-

tura del estatolito de esta especie.

KEY WORDS: /llex coindetiz, statolith, morphology, North-eastern Atlantic.

PALABRAS CLAVE: /llex coindetiz, estatolito, morfología, Atlántico noreste.

INTRODUCTION

Cephalopods play an important role components of the diet of such top pre-

in the trophic web of marine ecosystems dators as marine mammals (XAMPENY

as both predators and prey of many ma- AND FILELLA, 1976; CLARKE, 1980, 1986;

rine species (AMARATUNGA, 1983). These CLARKE, MARTINS AND PRINCE, 1993;

marine molluscs have been cited as GONZÁLEZ, LÓPEZ, GUERRA AND BA-

* Instituto de Investigaciones Marinas (Consejo Superior de Investigaciones Científicas), Eduardo Cabello 6,

36208 Vigo, Spain.

Iberus, 15 (1), 1997

RREIRO, 1994), large teleosteans (BOUXIN

AND LEGENDRE, 1936; BELLO, 1991; GUE-

RRA, SIMÓN AND GONZÁLEZ, 1993;

CLARKE, CLARKE, MARTINS AND SILVA,

1995), sea-birds (RODHOUSE, CLARKE

AND MURRAY 1987; CLARKE, CROAXALL

AND PRINCE, 1991; FURNESS, 1994; CROA-

XALL AND PRINCE, 1996) and cephalo-

pods (RASERO, GONZÁLEZ, CASTRO AND

GUERRA, 1996, RODHOUSE AND NIGMA-

TULLIN, 1996). Beaks, statoliths, chiti-

nous sucker rings and, to a lesser extent,

gladius, were the main structures which

allowed a positive identification to the

taxonomic level of species in studies

about trophic relationships between

cephalopods and other marine animals.

These hard structures remain unaltered

in the stomach contents of their preda-

tors and represent an important source

of information on the trophic relations-

hips where these animals are involved.

Statoliths of cephalopods are small

hard paired structures composed by cal-

cium carbonate in the form of aragonite.

They are situated in fluid-filled cavities

termed statocysts inside the cartilagi-

nous skulls of the cephalopods belon-

ging to the subclass Coleoidea (CLARKE

AND MADDOCK, 1988a). The statolith

and the macula (statoconia system)

constitute the receptor organ for detec-

tion of gravity. This is one of the func-

tions of the statocysts, in which the level

of sophistication is equivalent to the

vertebrate vestibular system (YOUNG,

1960, 1989; STEPHEN AND YOUNG, 1982;

BUDELMANN, 1978, 1988, 1990).

Statoliths are also the structures

most frequently used for studies on age

and growth. The statoliths of many cep-

halopod species show growth incre-

ments, which have been shown to have

a daily periodicity of deposition in seve-

ral species (see JACKSON, 1994).

Since teuthoid statoliths are appa-

rently species-characteristic and have a

greater likelihood of fossilisation than

other cephalopod structures, they have

become very important for identifi-

cation of fossil species (CLARKE AND

Frrch, 1975, 1979). The applications of

image analysis which have been used in

the morphological study of the stato-

132

liths, both recent and fossil, have shed

light on certain phylogenetic relations-

hips among cephalopods (CLARKE AND

MADDOCK, 1988a; 1988b). As was done

with shape analysis of fish otoliths

(CAMPANA AND CASSELMAN, 1993), the

statolith morphology of cephalopods

has been used also for stock discrimina-

tion (BORGES, 1995).

As ontogenetic changes do exist in

the statolith (MORRIS AND ALDRICH,

1984; GUERRA AND SÁNCHEZ, 1985;

CLARKE AND MADDOCK, 1988b; Bkru-

NETTI AND IvANOVIC, 1991), some

remains on the reliability of this appro-

ach about this subject. From descrip-

tions based solely on one statolith from

one specimen, the result of the analysis

of shape can be altered (LOMBARTE,

SÁNCHEZ AND MORALES-NÍN, 1995) and

a correct prey identification could also

be uncertain.

The aim of this study was to deter-

mine the changes in the statolith shape

for the ommastrephid squid lllex coinde-

ti1 (Vérany, 1839) during its ontogenetic

growth.

MATERIALS AND METHODS

341 specimens of Illex coindetii were

collected in the North-eastern Atlantic

(Fig. 1) from November 1991 to October

1992. Fishing was carried out at depths

ranging from 100 and 350 m over the

Galician continental shelf. The animals

were sexed, measured to the nearest

mm mantle length (ML), weighed (to 0.1

g) and assigned a maturity stage accor-

ding to LIPINSKI (1979). The squid ran-

ged from 48 to 379 mm ML. The stato-

liths were removed from the head and

preserved in 96% ethanol. Statolith ma-

jor axis and maximum width were re-

corded. Then, the statoliths were measu-

red using an eyepiece graticule. Terms

used in descriptions were assigned fo-

llowing the nomenclature established by

CLARKE (1978).

Measurements were made using an

image analysis system (IAS); the equip-

ment used is reviewed by Macy (1995).

The description of the each develop-

GONZÁLEZ AND GUERRA: Growth of /llex coindetiz statolith

$ Vigo

Miño River

, Burela.

Celeiro

-» z

8 W

Figure 1. /llex coindetiz. Fishing area where the samples were obtained.

Figura 1. Illex coindetii. Area de pesca donde se obtuvieron las muestras.

mental stage of the statolith was made

based on microphotographs.

RESULTS

Statolith morphology: Statolith

maximum length ranged from 0.47

(female of 48 mm ML) to 1.66 mm

(female of 360 mm ML). No significant

differences (p<0.05) between maximum

length and maximum width of male and

female statoliths of equivalent ML were

found at any stage of development.

Therefore, sex does not affect the growth

of the statolith in /llex coindetii.

Figure 2 shows a diagrammatic picture

of the different parts of an adult Illex coin-

detii statolith in posterior and anterior

view. Terms used in descriptions and the

measurements made are also shown.

Some characteristic features were obser-

ved in these statoliths: a) there is conti-

nuity between the dorsal dome and the

lateral dome. This feature can be obser-

ved between the superior and inferior

lobes of the lateral dome as well; b) the

posterior dome groove is very patent; c)

the rostrum is short and an anterior rostral

lobe does not exist; d) the wing is very

broad; e) the medial fissure is well deve-

loped and also has a small posterior inden-

tation; £) the dorsal spur is very clear.

Statolith development: Although

the above description refers to a late

developmental stage of the statolith in

an adult animal, the way to reach this

definitive stage is quite complex. Thus,

the statolith has become increasingly

complex and passed through different

stages which implies growth in different

planes. These stages can be described as

follows (Fig. 3):

Stage I: Statoliths of immature

animals ranging from 50 to 80 mm ML.

The medial fissure is not yet visible but

will be situated under the dorsal dome.

This area will be the surface where the

wing will connect with the body of the

183

Iberus, 15 (1), 1997

Dorsal dome

Dorsal indentation

Medial fissure == 2

Wing

Ventral indentation

Rostral angle

Rostrum

Superior lobe

Lateral lobe

Inferior lobe

Posterior dome groove

Rostrum

Dorsal dome

/ Posterior indentation

Wing

Maximum length

Maximum width

Figure 2. Diagrammatic picture of the different parts of an adult /llex coindetiz statolith.

Figura 2. Diferentes partes del estatolito de un Vlex coindetii adulto.

statolith in a later developmental stage.

Another characteristic of the stage 1 sta-

tolith is the wide rostral angle (>140").

The primordium of the rostrum is also

visible in the ventral zone of the stato-

lith. It is virtually impossible to differen-

tiate the dorsal dome, the superior and

inferior lobes of the lateral dome, being

round the general shape.

Stage II: This developmental stage

includes the statoliths of animals with ML

between 90 and 130 mm. The main cha-

racteristic of this stage is the growth of the

rostrum, and the change in shape of the

statolith, which is enlarging the ventral

direction. Small dark zones in the ventral

zone can be distinguished. This crystalli-

sation will form the wing of the statolith.

The wing formation is observed in two

directions, from the rostrum to the dorsal

plane of the statolith and from the dorsal

dome to the ventral plane. There is a slight

differentiation between superior and

medium lateral dome.

Stage III: This stage is remarkably dif-

ferent from the preceding one. It appears

in submature and mature squid ranging

from 130 to 200 mm ML. The enlargement

of the rostrum continues and the forma-

tion of the wing spans from the ventral to

the dorsal zone. The medial fissure is

small. A zone devoid of crystallisation

called the foramen appears for the first

134

time. The foramen runs parallel to the

rostrum and will disappear gradually. The

rostral angle is getting narrow and at this

stage 1t forms almost a 90” angle.

Stage IV: Mature animals between

200 and 250 mm ML. The developmen-

tal stage is close to definitive conditions.

The foramen is partially or totally occlu-

ded and the wing is formed along the

entire statolith. As the statolith grows,

the lobes become more distinct. Practi-

cally, the rostral angle is 90".

Stage V: This stage describes the sta-

toliths of mature specimens bigger than

250 mm ML. There are only minor

changes in morphology from stage IV.

Crystallisation in the wing is stronger

and the foramen is totally occluded. The

formation of the wing emphasises the

medial fissure.

Figure 4 illustrates the relationship

between the mantle length and the

maximum length and width of the lllex

coindetii statolith.

DISCUSSION

This paper gives a description of dif-

ferent developmental stages of Illex coin-

detii statoliths, based on observations of

the statolith growth. There are impor-

tant changes in shape of the statolith

GONZÁLEZ AND GUERRA: Growth of /llex coindetíz statolith

Figure 3. Stages of development for the statolith of //lex coindetiz. A: Stage 1, maximum statolith

length (MSL)= 0.67 mm and maximum statolith width (MSW)= 0.40 mm; B: Stage II, MSL =0.90

mm and MSW= 0.63 mm; C: Stage III, MSL= 1.13 mm and MSW= 0.80 mm; D: Stage IV, MSL=

1.20 mm and MSW= 0.80 mm; E: Stage V, MSL= 1.53 mm and MSW= 1.03 mm.

Figura 3. Estadios de desarrollo del estatolito de Ylex coindetii. A: Estadío 1, longitud máxima del estato-

lito (LME)= 0,67 mm y anchura máxima del estatolito (AME)= 0,40 mm; B: Estadío 1H, LME= 0,90

mm y AME= 0,63 mm; C: Estadío 11, LME 1,13 mm y AME= 0,80 mm; D: Estadío IV, LME= 1,20

mm y AME= 0,80 mm, E: Estadío V., LME= 1,53 mm y AME= 1,03 mm.

through the life cycle of Illex coindeti.

Considering the statolith of ommastrep-

hids, it is important to note the difficulty

in differentiating between the dorsal

dome and the superior and inferior

lobes described by CLARKE (1978) for

teuthoids. This observation agrees with

SÁNCHEZ (1981) for Illex coindetii speci-

mens from Mediterranean waters and

ARKHIPKIN (1990) and BRUNETTI AND

IVANOVIC (1991) for Illex argentinus. It

was also observed that the rostral angle

for statoliths of Illex coindetii juveniles is

clearly higher than 140? and progressi-

vely it is getting narrower until it

reaches 90? in mature animals.

Stages I and Il correspond to speci-

mens ranging from three to five months

of age. The description coincides with

stages I and II defined for Illex argenti-

nus (BRUNETTI AND IVANOVIC, 1991) and

the “definitive stage” noted by Morris

and ALDRICH (1984). The shape of the

statolith is enlarged to the ventral side

and it grows in the ventral zone. The

formation of the wing converges from

two different directions, from the dorsal

zone to the ventral one and vice-versa.

Stage III of the statolith of lllex coin-

detii corresponds to submature and

mature animals between five and eight

months of age ranging from 130 to 200

185

Iberus, 15 (1), 1997

Statolith measurements (mm)

—»

Mantle length (mm)

O Maximum length (mm)

+ Maximum width (mm)

Figure 4. /llex coindetii. Relationship between mantle length and maximum length and width .

Figura 4. Mex coindetii. Relación entre la longitud del manto y la anchura y longitud máximas.

mm ML. The union of the wing with the

dorsal dome of the statolith was obser-

ved in this stage, which agrees with the

observations Of MORRIS AND ALDRICH

(1984) for the “juvenile stage” of Illex

illecebrosus and stage III of ARKHIPKIN

(1990) and BRUNETTI AND IVANOVIC

(1991) for Illex argentinus. From this

stage onwards, the growth of the stato-

lith slows down; this could be related

with the process of maturation of the

animals as showed in the Figure 4. In

stage III appears for the first time the

foramen, which was noted as a characte-

ristic feature of ommastrephids. This is a

lack of crystallisation that runs parallel

to the rostrum. It is formed when both

planes of the wing grow, connecting

over the medial part of the statolith,

leaving one distinct zone between this

point and the body of the statolith.

Stages IV and V are very similar in

shape. However, a main difference can

be found when the foramen disappears

136

completely in stage V. Stage IV of Illex

coindetii corresponded to animals of

between 200 and 250 mm ML and ages

ranging from eight to ten months.

Finally, the statoliths of animals bigger

than 250 mm ML and older than ten

months are included in the stage V.

These statoliths are similar to those des-

cribed by MORRIS AND ALDRICH (1984)

for the “advanced stage” in statoliths of

Illex illecebrosus, to the stage VI observed

by BRUNETTI AND IVANOVIC (1991) for

Illex argentinus and the statolith descri-

bed by SÁNCHEZ (1981) for an adult /llex

coindetii specimen from the Mediterra-

nean Sea.

On the whole, it can be concluded that

the shape of the statolith changes gra-

dually from the juvenile stage until it

reaches a definitive stage of development.

There are three features to be noticed

during the growth of the lllex coindetii sta-

tolith: a) the variation of the rostral angle,

getting narrow progressively, from an

GONZÁLEZ AND GUERRA: Growth of /llex coindetíi statolith

angle wider than 120” in statoliths of juve-

niles to a 90” angle in mature animals; b)

the growth of the statolith slows down

when the animals reach sexual maturity;

c) the rounded shape of the statolith,

which makes difficult the differentiation

between the dorsal dome and the lateral

lobes; for this reason it is proposed to

extend the term dorsal dome to the supe-

rior lobe. Therefore, the inferior lobe of

the lateral dome should be called simply

lateral dome.

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-—.

La SocieDAD ESPAÑOLA DE IMALACOLOGÍA

Junta directiva desde el 18 de octubre de 1996

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Secretario Luis Murillo Guillén

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Francisco Javier Rocha Valdés

Gonzalo Rodríguez Casero

Jesús Souza Troncoso

José Templado González

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50 Iberus

E Revista de la

SOCIEDAD ESPAÑOLA DE MALACOLOGÍA

COMITÉ DE REDACCIÓN

EDITOR Ce

Ángel Guerra Sierra a Instituto de Investigaciones Marinas, CSIC, Vigo, España

EDITORES ADJUNTOS *

Eugenia M* Martínez Cueto-Felgueroso Universidad de Oviedo, Oviedo, España

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

Gonzalo Rodríguez Casero Universidad de Oviedo, Oviedo, España

ComiTÉ EDITORIAL

Kepa Altonaga Sustacha Universidad del País Vasco, Bilbao, España

Eduardo Angulo Pinedo Universidad del País Vasco, Bilbao, España

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

Sigurd v. Boletzky Loboratoire Arago, Bonyuls-surMer, Francia

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Edmund Gittenberger Notionaal Natuurhistorisch Museum, Leiden, Holanda

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

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notas breves. /berus edita un volumen anual que se compone de dos o más números.

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SOCIEDAD ESPAÑOLA

DE MALACOLOGÍA

Vol. 15 (2) Oviedo, diciembre 1997

Dep. Leg. B-43072-81

ISSN 0212-3010

Diseño y maquetación: Gonzalo Rodríguez

Impresión: LOREDO, S. L. - Gijón

PREFACE

This volume of /berus comprises a miscellany of various papers and posters presented

at the Twelfth International Malacological Congress held at Vigo (Spain) from 3 Septem-

ber to 8 September, 1996. This congress was organised by Dr. Ángel Guerra and Dr. Fran-

cisco Rocha, members of the Instituto de Investigaciones Marinas (IM) on behalf of Unitas

Malacologica and under the auspices of the Consejo Superior de Investigaciones Científi-

cas (CSIC), the Sociedad Española de Malacología (SEM) and the Cephalopod Interna-

tional Advisory Council (CIAC).

Based at the Cultutal Center of the Caixavigo and the adjacent Casa das Artes do Con-

cello de Vigo, the congress was attended by four houndred and twenty one specialists on

molluscs from different research fields of fifty seven countries. The framework of the Vigo

Congress was composed by five symposia, three free lectures and four workshops. These

ranged over all Classes of Mollusca, the marine, freswater and land environments, evolu-

tion and fossil records, phylogeny and systematics, ecology, medical and applied malaco-

logy, the functional morphology of cephalopods, endemisms in the marine realm, and data

bases. In addition to the communications most of the time was devoted to informal mee-

tings and discussions.

The congress was sponsored by over fourteen Governamental and Official Institutions

and companies. However, it would not be possible without the enthusiasm and the deter-

mination of the Local Organising Committee compossed mainly by people working in the

research group of Ecofisiología de Cefalópodos (1IM), the Universities of Vigo and San-

tiago de Compostela and the Sociedad Española de Malacología (SEM). Their forsight,

kindness and hard work made a very successful congress. We thank all of them and parti-

cularly to the director and the manager of IIM, Dr. Ricardo Pérez Martín and Mr. Luis

Ansorena, respectively. Our acknowdlegment to Dr. Emilio Rolán, President of the SEM

and co-editor of the book of abstracts!, Mrs. María Teresa Fernández, the person in charge

for all the secretariat affaires of the congress, and to all the organisers of the different ses-

sions, who wisely managed the symposia, workshops and free lectures. We are very grate-

ful to all the referees for their expert assistance reviewing these manuscripts. Delay in publi-

cation of this volume is basically our responsability, but we wish to point out correspon-

dence with some of the authors was not easy.

In the oppening address to the Vigo Congress, Dr. Winston E Ponder argued that mala-

cology is not doing justice to the importance of molluscs. Molluscs are a large phylum of

large bodied, well-known animals with a superb fossil record, excellent model organisms

in evolution, genetics, physiology and ecology, economically important (fisheries and cul-

tures), and as agricultural pests and carriers of disease. Analysis of several experts revealed,

however, that few papers on molluscs are given at international meetings or published in

mainstream journals, and instead, most appear in malacological meetings and journals, and

much of it is narrowly focussed and trivial, indicating an inward focus and conservatism.

If malacology is to improve its perception by the scientific community, the basic research

must be focussed on areas of greatest interest, the study of molluscs must contribute to

major areas of scientific inquiry and social and economic concern. Our discipline also needs

a strong and effective voice in the traditional (books and journals) and the new (electro-

nic) media. This, and other issues in non-molluscan specialised journals which now are in

press, must be regarded as an effort of the Vigo Congress organisers to reverse the trend of

margination of molluscan studies in mainstream biology.

This volume has been published under the inestimable collaboration of the associated

editors of Iberus D. Gonzalo Rodríguez Casero and Dr. Eugenia Martínez Cueto-Felgue-

roso. Nine papers are included in this volume. These articles embrace very different aspects

of the malacology and comprise distincts groups of molluscs that belongs to several ecosys-

tems. Thus, we find in this book studies which run from the biogeography and demo-

graphic response of the snail populations to enviromental conditions to the study of mollus-

can evolution, going through inmunology and morphological researches.

Finally, we would like to thank the Spanish Ministry of Education and Science, the

Education Ministry of the Galician Government, the manager of the 5% Centenary of the

Universidad de Santiago de Compostela and the Chancellor of the Universidad de Vigo

for their support to organise the “Iwelfth International Malacological Congress and spe-

cially the Council of Unitas Malacologica for providing funds for publication of this volume

of Iberus.

Angel Guerra and Francisco Rocha

| GUERRA, A., E. ROLÁN AND F. ROCHA (EDS.), 1995. Abstracts of the Twelfth International Malacological

Congress. Vigo, 3th-8th September, 1995. Unitas Malacologica and Instituto de Investigaciones Marinas (CSIC),

Vigo, Spain. 530 pp.

O Sociedad Española de Malacología — ——_———T— lIberus, 15 (2): 1-11, 1997

Phagocytosis by haemocytes from the Lesser Octopus Ele-

done cirrhosa

Fagocitosis en hemocitos del pulpo blanco Eledone cirrhosa

Shelagh K. MALHAM”, Norman W. RUNHAM' and Christopher J. SECOMBES”

Recibido el 8-1-1996. Aceptado el 11-IV-1996

ABSTRACT

Haemocytes from Eledone cirrhosa phagocytose formalized bacteria (Vibrio anguillarum).

The phagocytic capabilities of E. cirrhosa haemocytes are affected by several factors, inclu-

ding the haemocyte culture medium, temperature, duration of the assay, and the bacterial

pre-incubation conditions such as haemolymph concentration, temperature and the duration

of pre-incubation.

Haemocytes will phagocytose in the absence of haemolymph. With a 30min incubation

period the number of phagocytosing haemocytes increases as the pre-opsonization concen-

tration and incubation temperature increase. However after 2 hours at 15 or 20*C the num-

ber of haemocytes phagocytosing unopsonized bacteria is equivalent to the number engul-

fing 100% haemolymph opsonized bacteria.

RESUMEN

Los hemocitos de Eledone cirrhosa fagocitan bacterias formalizadas (Vibrio anguillarum). La

capacidad de fagocitar en estas células se ve afectada por varios factores, incluyendo el

medio de cultivo de los hemocitos, temperatura, duración del experimento, y las condicio-

nes de preincubación de las bacterias, tales como concentración de hemolinfa y tempera-

tura y duración de la preincubación. Los hemocitos fagocitan en ausencia de hemolinfa.

Con un periodo de incubación de 30 minutos, el numero de hemocitos que fagocitan se

incrementa cuando lo hacen la concentración de preopsonización y la temperatura de incu-

bación. Sin embargo, tras dos horas a 15 ó 20*C, el número de hemocitos que fagocitan

bacterias no opsonizadas es equivalente al de hemocitos que fagocitan bacterias tratadas

con hemolinfa al 100%.

KEY WORDS: Eledone cirrhosa, haemocytes, phagocytosis, opsonization.

PALABRAS CLAVE: Eledone cirrhosa, hemocitos, fagocitosis, opsonización.

INTRODUCTION

In vivo and in vitro investigations into blood cells or haemocytes are avidly pha-

the cellular activities of molluscs have de- gocytic and capable of recognising non-

monstrated that, in a number of cases, the self (reviewed by MILLAR AND RATCLIFEFE,

“University of Wales, School of Biological Sciences, Brambell Building, North Wales, Bangor, Gwynedd LL57

2UW, United Kingdom.

” Department of Zoology, University of Aberdeen, Tillydrone Ave, Aberdeen, Scotland AB9 2T'N, U.K.

Iberus, 15 (2), 1997

1994). The process of phagocytosis invol-

ves a number of recognizable stages,

which include attraction, attachment, in-

gestion and killing of foreign organisms,

and is influenced by a number of factors

(reviews by RATCLIFFE, ROWLEY, FITZGE-

RALD AND RHODES, 1985; MILLAR AND

RATCLIFFE, 1994). Variables which have

been shown to affect phagocytic rates in

molluscs include incubation temperature

(FOLEY AND CHENG, 1975), time and pH

(ABDUL-SALAM AND MICHELSON, 1980),

the size of the particle presented for pha-

gocytosis and the nature of the particles

(reviewed by BAYNE, 1983). Though pha-

gocytosis will take place in the absence of

opsonizing agents (RENWRANTZ AND

STAHMER, 1983; TUAN AND YOSHINO

1987; FRYER, HULL AND BAYNE, 1989), se-

veral experiments have shown that solu-

ble humoral factors or opsonins may be

instrumental in non-self recognition

(PROWSE AND TAIT, 1969) and, or enhan-

cement of phagocytosis (reviews by JEN-

KIN, 1976; RATCLIFFE ET AL., 1985).

The haemocyte culture medium has

been shown to influence phagocytosis

with, in the case of the Asian clam, Cor-

bicula fluminea, the presence of divalent

cations being necessary for both

opsonin-independent and opsonin-

dependent phagocytosis (TUAN AND

YOSHINO, 1987). The process of opsoni-

zation also appears to be influenced by

several other factors. FRYER AND BAYNE

(1989), using Biomphalaria glabrata,

showed that for this mollusc opsoniza-

tion is a time-dependent process.

Further, TrIPP (1992), working with Mer-

cenaria mercenaria demonstrated that at

low temperatures, opsonization caused

enhanced phagocytic rates.

The octopus Eledone cirrhosa is

benthic in habit, ranges in depth from

sub-littoral to 770 m and encounters

temperatures between 5 and 15”C

(BOYLE, 1983). The animal has a closed

circulatory system and if wounded pre-

vents blood loss by local vasoconstric-

tion of the area surrounding the wound.

The blood of the octopod does not clot

and further blood loss is prevented by

allowing seepage of blood through the

wou1.d until blood cells eventually plug

the wound (WELLs, 1978, 1983; BAYNE,

1983). If the animal loses a large amount

of blood a dilution of the respiratory

pigment (haemocyanin) occurs which

takes up to 2 hours to be reversed

(WELLS AND WELLS, 1993). There

appears to be only one main type of

blood cell or haemocyte in E. cirrhosa.

The haemocyte matures in the white

body, or haematopoetic organ, of the

animal and is released into the closed

circulatory system (COWDEN AND

Curtis, 1974, 1981). Few cephalopod

defense mechanisms have been elucida-

ted (FORD, 1992). It is known that E. cirr-

hosa haemocytes will phagocytose eryth-

rocytes only in the presence of hae-

molymph in vitro (STUART, 1968). Also in

vivo studies (STUART, 1968; BAYNE, 1973)

using different octopods, demonstrate

that it is mainly fixed phagocytes in

certain organs which clear injected

foreign particles, with haemocytes only

removing a small fraction of them.

This paper investigates whether hae-

mocytes from E. cirrhosa are capable of

phagocytosing dead bacteria in vitro and

whether temperature, time and hae-

molymph concentrations influence pha-

gocytosis. Additional experiments were

also performed to determine whether

bacterial pre-incubation (opsonization)

at different temperatures, times and

haemolymph concentrations affected

phagocytic rates.

MATERIALS AND METHODS

Animals: Octopuses, Eledone cirrhosa

(Lamarck) were obtained from crab pots

around the North Wales coast. The

animals were brought into the aquarium

at the University of Bangor and maintai-

ned in natural seawater at 10-12*C.

After 48 h the animals were weighed,

marked using a panjet and assigned to a

particular tank. Five octopuses per tank

were chosen at random for each set of

experiments.

Haemolymph: Blood was withdrawn

from the branchial blood vessel of each

octopus as described by MALHAM, SECOM-

MALHAM E7 AL.: Phagocytosis by haemocytes from Eledone cirrhosa

BES AND RUNHAM (1995). The blood was

centrifuged at 4*C for 5 min at 800g to

remove the haemocytes. The resulting hae-

molymph from a number of individuals

was pooled and frozen at -20*C. Before

use the haemolymph was thawed and

diluted to a final concentration of 0. 1, 1

or 10% in Sterile Octopus Saline (SOS)

(NaCl, 2.367 g/100 ml; Glucose, 1 g/100

ml; CaCL, 0.116 g/100 ml; KH2PO;, 0.0056

g/100 ml; KCl, 0.1089 g/100 ml;

MgSO4-H20, 0.503 g/100 ml; MgCL, 0.419

g/100 ml).

Haemocytes: From each animal 1 ml

blood samples were withdrawn into 10

ml ofice cold Marine Anticoagulant (NaCl,

2.63 g / 100 ml; Glucose, 1.8 g/100 mi; Tri- *

Sodium Citrate, 0.088 g/ml; Citric Acid,

0.055 g/100 ml) containing ethylene gly-

col-bis(b-aminoethylether) N, N, N”, N', -

tetraacetic acid (EGTA) (0.029 g/100 ml).

After a blood count the haemocytes were

centrifuged at 800 g for 5 min at 4*C, and

washed by resuspension in Octopus Rin-

ger (NaCl, 2.433 g/100 ml; Glucose, 1.4

g/100 ml; EGTA, 0. 015 g/100 ml; KCl,

0.082 g/100 ml; KH2PO),, 0.004 g/100 ml)

containing CaCL (0.0142 g/100 ml), MgCl

(0.0524 g/100 ml) and MgSO: (0.0629

g/100 ml). A final haemocyte count was

made before the haemocytes were washed

for a second time and resuspended in SOS

at 1 x 10 haemocytes/ ml.

Bacteria: Vibrio anguillarum (MT275)

were obtained from the Scottish Office,

Agriculture and Fisheries Department,

Marine Laboratory, Aberdeen. Formali-

zed V. anguillarum were counted, was-

hed twice by resuspension in SOS and

centrifuged at 13000 g for 10 min before

resuspension at 8 x 10% cells/ml in the

required treatments.

Transmission electron microscope

(T.E.M.) preparation: Five hundred yl of

blood was withdrawn from the bran-

chial blood vessel of the octopus and

mixed directly with 500 yl of washed

bacteria. After 2h incubation at 15*C the

blood was centrifuged and the hae-

molymph removed. The pelleted hae-

mocytes were fixed for 24 h at 4*C in

2.5% glutaraldehyde (in 0.1M sodium

cacodylate buffer at pH 7.4). The hae-

mocytes were washed in 0.1M sodium

cacodylate buffer and secondarily fixed

for 2 h at room temperature in 1%

osmium tetroxide before staining en bloc

with 2% uranyl acetate over night. The

pellet was then dehydrated through

ethanol and propylene oxide and

embedded in Spurr resin. Cut sections

(50 nm) were mounted on 100 mesh pio-

loform copper coated grids and stained

with lead citrate. Sections were viewed

in a GEC Corinth 500 at 60 KV.

Phagocytosis assay: Two phagocyto-

sis experiments were performed to

determine the effect of haemolymph

concentration, temperature and time on

haemocyte phagocytosis. Five animals

were used for each experiment. The first

experiment involved incubating hae-

mocytes in 16 well tissue culture slides

(Nunc) for 2 h at different temperatures,

but utilizing one pre-incubation tempe-

rature and time for the bacteria. The

second experiment involved haemocyte

incubations of 30 min only and utilized

different temperatures, times and hae-

molymph concentrations for bacterial

pre-incubations.

For the first experiment 50 ml of the

haemocyte suspension in SOS was put

into each of the 16 well chambers of a

tissue culture slide. Fifty microliters of

either SOS or haemolymph diluted in

SOS was added in duplicate, at half

hour intervals, to selected wells. Bacte-

ria were resuspended in either SOS or

100% haemolymph for 2 h at 15*C and

washed twice before use. Fifty microli- .

ters of either SOS treated or hae-

molymph treated bacteria immediately

followed the haemolymph additions,

again in duplicate. Each well of the

tissue culture slide therefore contained:

50 ml of haemocytes in SOS, 50 ml of

either SOS or haemolymph diluted in

SOS to 0.1, 1 or 10% concentration (final

concentrations of 0.03, 0.33 or 3.33% res-

pectively) and 50 ml of bacteria resus-

pended in SOS after treatment. The

assays were run at four temperatures (5,

10, 15 and 20%C). After 2 h the tissue

Iberus, 15 (2), 1997

culture slides were rinsed in SOS to

remove unattached bacteria and the

slide fixed by immersion in methanol

for 3-5 min.

The second experiment involved the

addition of 50 ml of haemocytes in SOS

at 1 x 10% haemocytes/ml, followed by

50 ml of haemolymph diluted in SOS at

0, 0. 1, 1 or 10% concentrations and 50

ml of the different bacterial preparations

added in duplicate to the tissue culture

slides. The bacteria were washed and

resuspended in haemolymph at concen-

trations of 0, 0.1, 1, 10 or 100%, using

Phosphate Buffered Saline pH 7.0 (PBS,

Gibco, without Ca** and Mg?*) as the

diluent. Bacteria were incubated for 1,

10, 60 or 120 min at 5, 10, 15 or 20*C,

before being washed twice and used in

the assay. The slides were incubated at

temperatures of 5, 10, 15 or 20*C. After

30 min the tissue slides were rinsed with

SOS and the experiment stopped by

immersion of the slide in methanol as

previously.

All slides were then stained in

Giemsa (Sigma), rinsed in Gurr Buffer

(BDH pH 6.8) and air dried before

mounting using DPX.

Statistical analysis: Analysis was

performed by random counting of 200

haemocytes in each well. The haemocy-

tes were counted under oil using a com-

pound binocular microscope at 800x

magnification. All slides were numbe-

red and randomly selected to reduce

observer bias. The number of haemocy-

tes which had phagocytosed bacteria

was expressed as a percentage of the

haemocytes counted in each of the

duplicate wells. The results for each of

the duplicate wells were averaged and

analysis of variance (ANOVA) perfor-

med for the 2 experiments using the 5

replicates. In each case P values of < 0.05

were taken as being significant. The

replicate means were calculated and

Tukey's pairwise comparison was per-

formed for each experiment using the

calculated confidence interval esti-

mation (CI estimation). The CI estimate

allows 2 separate means to be statisti-

cally compared (Rick, 1988).

RESULTS

Phagocytosis of the formalized

Gram negative bacterium, V. anguilla-

rum, by E. cirrhosa haemocytes occurs

both in the presence and absence of hae-

molymph. Collected haemocytes were

incubated with bacteria for 2 h before fi-

xation for T.E.M. Sections clearly indi-

cate that E. cirrhosa haemocytes pha-

gocytose and degrade bacteria (Fig. 1).

From Analysis of variance a number

of significant conclusions were obtai-

ned. Phagocytosis by haemocytes follo-

wing pre-incubation of the bacteria in

100% haemolymph was significantly

greater than phagocytosis following

SOS treatment (F= 594.85, P<0.0001)

(Fig. 2). Highly significant values were

also obtained for the effect of incubation

temperature (F= 155.09, P<0.0001), and

also for the duration of the assay (F=

178.9, P<0.0001). The concentrations of

haemolymph used in the assay medium

did not have a significant effect (E= 0.32,

P= 0.814) indicating that the rate of pha-

gocytosis was statistically equivalent in

assays containing 0, 0.1, 1 or 10% hae-

molymph.

Cross-wise comparisons of the per-

centage of haemocytes phagocytosing

opsonized and unopsonized bacteria,

temperature and assay duration were

also highly significant, (P<0.0001),

whereas cross-wise comparisons invol-

ving haemolymph concentration in the

assay medium, confirmed that the hae-

molymph concentrations, in SOS, did

not affect phagocytic rates. Hae-

molymph concentration was therefore

not considered in further analysis, and

results at each temperature and time

were pooled.

Phagocytosis of bacteria pre-incuba-

ted in SOS was affected by temperature

and time (Fig. 2A). At all temperatures

the number of haemocytes engulfing

bacteria increased over time. At 20*C

there appeared to be fewer haemocytes

phagocvtosing than at 15%C, however

statistically there was no difference

between the means at the 2 temperatu-

res. At 109C there was a rapid increase

in the number of haemocytes phagocy-

MALHAM ET AL.: Phagocytosis by haemocytes from Eledone cirrhosa

IS TAE

Figure 1. Transmission electron micrograph of an Eledone cirrhosa haemocyte (H) having engulfed

a bacterium (Vibrio anguillarum) (B). Scale bar 10 pm.

Figura 1. Microfotografía de un hemocito (H) de Eledone cirrhosa tras haber tragado una bacteria

(Vibrio anguillarum) (B). Escala 10 ym.

tosing bacteria during the first 30 min

followed by a slower rate of increase up

to 2 h. At both 5 and 10*C significantly

lower phagocytic rates were observed

than at 15 and 20*C over the 2 h period.

Fig. 2B shows the mean number of hae-

mocytes phagocytosing bacteria, pre-

incubated in 100% haemolymph, over

time. The haemocyte phagocytic rate

again increased over the 2 h period but

there were far smaller differences

between the incubation temperatures.

The phagocytic rates were again lower

at 5%C than at the other temperatures.

The maximum increase in phagocytosis

at all temperatures occurred within the

first 30 min.

As with the first experiment, the dif-

ferent concentrations of haemolymph in

SOS (at 0, 0.1, 1 or 10%) used in the se-

cond assay were found to have little ef-

fect, so were removed from the pair wise

comparison with no appreciable percen-

tage error increase (0.027%) and the re-

sults pooled at each pre-incubation tem-

perature and time. To simplify the pair-

wise comparison the assay temperature

was not included as a main factor, but

was added as an interacting factor. The

results from the simplified model show

that there were large statistically signifi-

cant differences (F= 1083.35, P<0.0001)

between the haemolymph pre-incuba-

tion concentrations. The pre-incubation

temperatures (F= 61.32, P<0.0001), and

the pre-incubation times (F= 725.24,

P<0.0001) were similarly significantly

different. Pre-incubation of the bacteria

in PBS alone at different temperatures

and time periods caused no significant

difference in the phagocytic rate (Fig. 3).

Bacteria pre-incubated in 0.1% hae-

molymph in PBS at all pre-incubation

temperatures and times were phagocyto-

sed at a significantly lower rate than in

PBS alone. Pre-incubation of the bacteria

Iberus, 15 (2), 1997

[o]

[>]

8 60

S 50

[5]

£=,

E 40

p0)

E 30

——5

EY) —— 10

A 15

10 —= 20

0 0,5 1 1,5 2

Time (Hours)

3 60

3 50

(

E 40

(0)

a 30

——5

20 —==— 10

—= 20

0 0,5 1 1,5 2

Time (Hours)

Figure 2. A: phagocytosis of non-opsonized formalized Vibrio anguillarum at 4 temperatures over a

2h haemocyte incubation period. The bacteria were pre-treated with SOS for 2h at 15*C. Tukeys

CI estimate= 9.52. B: phagocytosis of opsonized formalized Vibrio anguillarum at 4 temperatures

over a 2h haemocyte incubation period. The bacteria were pre-treated with 100% haemolymph for

2 h at 15”C. Tukeys CI estimate = 9.52.

Figura 2. A: fagocitosis de bacterias formalizadas Vibrio anguillarum ro opsonizadas a cuantro tem-

peraturas sobre un periodo de incubación de'hemocitos de dos horas. Las bacterias fueron pretratadas con

SOS durante 2 horas a 15*C. Estimación CI de Tukeys= 9,52. B: fagocitosis de Vibrio anguillarum for-

malizado y opsonizado a cuatro temperaturas sobre un periodo de incubación de hemocitos de 2 horas.

Las bacterias fueron pretratadas con hemolinfa al 100% durante 2 horas a 15*C. Estimación del inter-

valo de confianza de Tukeys= 9,52.

in 1% haemolymph showed initially the valent to the values determined in PBS

same lowered phagocytic rate as for alone. However, at 10 min following pre-

0.1% pre-incubation. However, pre-incu- incubation at 15 and 20%C more hae-

bation of the bacteria in 1% haemolymph mocytes were observed phagocytosing

for 10 min at 20C caused an enhanced bacteria than at 5 or 10*C, or at 1 min at

phagocytic rate which also occurred at all temperatures. The enhanced pha-

all temperatures at 60 and 120 min. Bac- gocytic rate observed using 10% hae-

teria pre-incubated in 10% haemolymph molymph is statistically equivalent to

for 1 min at 5, 10, 15 and 20*C and for 10 the enhanced rate observed at 1% con-

min at 5 and 10*C were statistically equi- centration.

MALHAM ET AL.: Phagocytosis by haemocytes from Eledone cirrhosa

Mb |

AN pl il

] | 0.1% Haemolymph

19)

(e)

MI pl

3315

25 10

”n o

ES 2

(e)

IN)

AN 40

Si N M NN

IN]

! 00% | 100% Haemolymph | ECAARAN 20

i ] a

Cross Tabulated Mean

Percent Phagocytosis

10% Haemolymph

Pre-Incubation Time

Figure 3. Phagocytosis of formalized Vibrio anguillarum. The haemocytes were incubated at diffe-

rent temperatures for 30 min only. The bacteria were pre-incubated in 0% haemolymph (.e., PBS

only), 0.1% haemolymph, 1% haemolymph, 10% haemolymph and 100% haemolymph concen-

trations. The bacterial pre-incubation temperatures were 5, 10, 15 and 20*C and the pre-incuba-

tion times were 1, 10, 60 and 120 min. Tukeys CI estimate = 3.1.

Figura 3. Fagocitosis de Vibrio anguillarum formalizado. Los hemocitos fueron incubados a diferentes

temperaturas durante sólo 30 minutos. Las bacterias fueron preincubadas en concentraciones de hemo-

linfa del 0% (1.e., sólo PBS), 0,1%, 1%, 10% y 100%. Las temperaturas de preincubación de las bac-

terias fueron 5, 10, 15 y 20*C y los tiempos de preincubación de 1, 10, 60 y 120 minutos. Estimación

del intervalo de confianza de Tukeys= 3,1.

DISCUSSION

The results presented here demons-

trate that E. cirrhosa haemocytes are

capable of recognizing and ingesting the

formalized bacterium Vibrio anguillarum.

V. anguillarum is a Gram negative com-

mensal marine opportunist and was

chosen as the experimental bacterium

because it has been isolated from, and

used in previous studies on wound

healing in E. cirrhosa (BULLOCK, PoL-

GLASE AND PHILLIPS, 1987). This bacte-

rium has also been implicated in

causing cephalopod infections when the

animals are held in captivity and is a

common contributory cause of death at

high aquarium temperatures (LEIBOVITZ,

MEYERS AND ELSTON, 1977; HANLON,

FORSYTHE, COOPER, DINUZZO, FOLSE

AND KELLY, 1984; FORD, ALEXANDER,

COOPER AND HANLON, 1986; HANLON

AND FORSYTHE, 1990).

STUART (1968) found that E. cirrhosa

haemocytes required haemolymph for

in vitro phagocytosis of erythrocytes.

The data presented in this paper

demonstrate that the presence of hae-

molymph is not necessary for ingestion

of bacteria. However, this bacterium is

smaller with far less surface area than

an erythrocyte and as such maybe more

easily phagocytosed. It was found by

TYSON AND JENKIN (1974) that haemocy-

tes from a crayfish (Parachaeraps bicarina-

tus) phagocytosed bacteria in the

absence of haemolymph, but erythrocy-

tes were not phagocytosed unless they

were pre-treated with haemolymph

(McKay, JENKIN AND ROWLEY, 1969).

Further JENKIN (1976), suggested that

the concentration of certain recognition

Iberus, 15 (2), 1997

molecules on the crayfish haemocyte

surface was not sufficient to bind eryth-

rocytes, but was sufficient to bind bacte-

ria, and a similar explanation could

apply to E. cirrhosa haemocytes. Another

possibility was demonstrated by BAYNE,

MOORE, CAREFOOT AND THOMPSON,

(1979), who showed that haemocytes

from Mytilus californianus had a greater

affinity for yeast cells than human

erythrocytes, and suggested that pha-

gocytosis of foreign particles was selec-

tive. Results from other molluscan

species also demonstrate that surface

antigenicity of the respective test parti-

cles has an effect on phagocytosis by

haemocytes (TRIPP AND KENT, 1967;

ANDERSON AND GOOD, 1976).

Tripp (1966), using the bivalve M.

mercenaria, concluded that haemolymph

pre-treatment of erythrocytes caused

increased phagocytosis. The same expe-

riment showed however that if untrea-

ted erythrocytes were incubated with

haemocytes for longer periods of time,

the same levels of phagocytosis were

achieved. With E. cirrhosa haemocytes at

15 and 20*—C the phagocytic rate is

higher at 30 min for 100% haemolymph

treated bacteria compared to SOS

treated bacteria, but after 2 h there was

no difference in phagocytic rates

between the 2 treatments. The data pre-

sented here also indicate that a higher

percentage of haemocytes phagocytosed

haemolymph treated bacteria at 5 and

10*C over 2 h than SOS treated bacteria.

Tripp (1992) also showed that the hae-

mocytes of M. mercenaria were avidly

phagocytic in the absence of hae-

molymph, however at low temperatu-

res, in the presence of haemolymph

there was increased phagocytosis of

yeast. ABDUL-SALAM AND MICHELSON

(1980), working with Biomphalaria gla-

brata, also demonstrated that tempera-

ture has an effect on haemocyte pha-

gocytosis. A phagocytic activity peak

was evident at 30C with inhibition of

phagocytosis below 15%C. Low tempera-

ture inhibition (4%C) of phagocytic rates

has also been demonstrated for the hae-

mocytes from the hard clam M. mercena-

ria with maximum rates occurring at 22

and 37%C (FOLEY AND CHENG, 1975).

With SOS treated bacteria, E. cirrhosa

haemocytes demonstrate an activity

peak with about 70% of haemocytes

phagocytosing after 2 h at 15 and 20*C.

At 5*C only 14% of haemocytes contai-

ned bacteria, whereas if the bacteria

were initially pre-incubated in hae-

molymph before addition to the assay

the phagocytic rate at 5%C increased to

around 47%.

The results presented above indicate

that the amount of haemolymph present

in the bacterial pre-incubation medium

has a dramatic effect on the number of

haemocytes subsequently engulfing

these bacteria within a 30 min period.

Haemolymph concentrations of 0.1 and

1% in PBS, resulted in lower numbers of

haemocytes phagocytosing compared to

PBS alone. This inhibition changes to

enhanced phagocytosis, at all higher

pre-incubation concentrations. Further

comparisons demonstrate that the tem-

perature of the pre-incubation medium

and particularly the duration of incuba-

tion are also important factors. The ob-

served trends indicate that increasing

the pre-incubation temperature decrea-

ses the pre-incubation time needed for

enhanced phagocytosis to occur. FRYER

ET AL. (1989), working on B. glabrata, si-

milarly demonstrated that phagocytosis

was inhibited after short pre-incubation

periods, whereas longer pre-incubation

periods of 1 h resulted in enhanced le-

vels. It was suggested by the authors

that initial non-specific adsorption of a

variety of plasma components (opso-

nins) occurred onto, in their case, the ye-

ast surface. Longer exposure to the

plasma allowed more of the opsonins to

bind to the yeast surface. The results

from the data presented here for the dif-

ferent pre-incubation haemolymph con-

centrations and durations of exposure

seem to support this hypothesis. In ad-

dition it is possible that if the tempera-

ture is increased further more of the

available plasma components would ad-

here onto the surface of the bacterium.

When haemocytes from E. cirrhosa

were resuspended in SOS, as stated

above, there is phagocytosis of the for-

MALHAM E7 AL.: Phagocytosis by haemocytes from Eledone cirrhosa

malized bacterium V. anguillarum. In

buffers containing either EDTA or

EGTA, no phagocytosis of the same bac-

terium was evident (Malham, unpublis-

hed data). SOS contains Ca?”* and Mg?*

and it appears likely that the presence of

these divalent ions has an effect on pha-

gocytosis. FRYER AND ADEMA (1993) sho-

wed that manipulated haemocytes from

B. glabrata retained some phagocytic ac-

tivity, but that addition of excess Ca?*

and Mg?* to the haemocytes before the

addition of the target particles enhanced

their phagocytic rates. E. cirrhosa hae-

mocytes were initially drawn into an an-

ticoagulant buffer containing EGTA and

washed in Octopus Ringer, also contai-

ning EGTA, before resuspension in

EGTA-free-SOS, all of which could alter

haemocyte behaviour and affect pha-

gocytosis. Corbicula fluminea haemocytes

(TUAN ET AL., 1987) also required extra-

cellular Ca?* or Mg?* for both opsonin-

dependent and independent phagocyto-

sis. The authors suggest that the opso-

nin possibly exists as a divalent

cation-macromolecular complex due to

the loss of enhanced phagocytosis after

dialysis against EDTA and EGTA. Furt-

her, Mytilus edulis haemocytes phagocy-

tosed yeast cells with high efficiency

when calcium ions were present in the

suspension medium, and gave similar

results when haemolymph alone was

added, but almost no phagocytosis was

recorded with haemocytes in buffered

saline (RENWRANTZ AND STAHMER,

1983). When V. anguillarum was resus-

pended in SOS, E. cirrhosa haemolymph

diluted in SOS, or in PBS alone, there

was no change in the haemocyte pha-

gocytic rate. However, when V. anguilla-

rum was resuspended in haemolymph

diluted in PBS (2 1% haemolymph con-

centration) or in haemolymph alone, en-

hanced phagocytosis was observed.

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218.

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O Sociedad Española de Malacología ——__—_—_—_—_—_——— Iberus, 15 (2): 13-24, 1997

New data on the morphology and the distribution of Bul;-

mulus corneus Sowerby, 1833 (Gastropoda: Pulmonata:

Orthalicidae) in Nicaragua

Nuevos datos sobre la morfología y la distribución de Bulimulus cor-

neus Sowerby, 1833 (Gastropoda: Pulmonata: Orthalicidae) en Nica-

ragua

Antonio Mijail PÉREZ*! and Adolfo LÓPEZ*

Recibido el 8-[-1996. Aceptado el 22-IV-1996

ABSTRACT

Aspects related to the morphology and distribution of Bulimulus corneus Sowerby, 1833 in

Nicaragua are presented. Regarding morphology, a complete redescription of the shell

and the first description of the genitalia are included. The number of records have been

largelly increased; from three localities mentioned in the literature to 53. The previous figu-

res have allowed us to draw a preliminary distribution map of the species in Nicaragua,

and discuss the presence of the closely related species Bulimulus unicolor Sowerby, 1833

in the country.

RESUMEN

Se presentan aspectos relacionados con la morfología y la distribución de Bulimulus cor-

neus Sowerby, 1833 en Nicaragua. En relación con la morfología, se presenta una redes-

cripción de la concha y la primera descripción del aparato genital. El número de registros

de la especie en el país ha sido notablemente incrementado de tres a 53 localidades. Las

cifras anteriores nos han permitido confeccionar un mapa preliminar de distribución para

la especie en Nicaragua, así como discutir la presencia de Bulimulus unicolor Sowerby,

1833, una especie muy relacionada, en el país.

KEY WORDS: Bulimulus corneus, Orthalicidae, morphology, distribution, Nicaragua.

PALABRAS CLAVE: Bulimulus corneus, Orthalicidae, morfología, distribución, Nicaragua.

INTRODUCTION

According to BREURE (1979), the distribution of Bulimulus corneus

genus Bulimulus Leach, 1814 contains 88 Sowerby, 1833 as from SW Mexico to the

species, distributed over the Antilles, central zone of Costa Rica, there being

Central America and northern South apparently no records outside of these

America. MARTENS (1890-1901) gave the limits. In Nicaragua, previous reports

* Universidad Centroamericana, Apartado A-90, Managua, Nicaragua.

' Dirección postal temporal: Universidad del País Vasco, Facultad de Ciencias, Departamento de Biología

Animal y Genética, Laboratorio de Zoología, Apartado 644, 48080 Bilbao, España.

Iberus, 15 (2), 1997

DMAX

Figure 1. Bulimulus corneus. Shell measurements.

Abbreviations. LONG: length; DMAX: maximum diameter; LEC: height of the body whorl;

DME: minimum diameter; LAB: aperture length; AAB: aperture width; LEP: spire length; AEP:

spire width.

Figura 1. Bulimulus corneus. Medidas de la concha.

Abreviaturas. LONG: longitud; DMAX: diámetro máximo; LEC: altura de la vuelta principal;

DME: diámetro mínimo; LAB: altura de la abertura; AAB: anchura de la abertura; LEP: altura de

la espira; AEP: anchura de la espira.

have been from Realejo (Chinandega),

San Juan Castillo (sic), El Toro rapids,

RAAN (Autonomous Region of the

North Atlantic) (MARTENS, 1890-1901)

and Bluefields (FLUCK, 1900).

Martens stated that this species is

closely related to Bulimulus unicolor So-

werby, 1833, and this was confirmed by

PiLSBRY (1897). None of the authors re-

cognized TATE's (1870) reports of B. uni-

color from Granada, Mesapa and San Ni-

colas, on the Pacific slope of Nicaragua.

The internal and external morpho-

logy of B. corneus, shell measurements

and data on distribution were recently

presented for the first time in an abstrac-

ted version (PÉREZ AND LóÓPEz, 1995),

and are here given in detail. New distri-

bution data gathered in the last few

months are also presented, together

with a commentary on the presence of

B. unicolor in Nicaragua.

MATERIALS AND METHODS

All specimens were hand-collected and

live specimens were relaxed in menthol

and fixed in 70% alcohol. All individuals

considered for the study were fully- grown

adult specimens. All localities reported are

additions to those previously mentioned

in the literature. The list of localities is gi-

ven in Table 1. The distribution map was

made using the UTM cartographic met-

hod with a grid size of 100 Km?. When

more than one locality occur on the same

UTM 10 Km? quadrat, only the one that

appears first in the listis mapped. The ab-

breviations w. l. n. and Bib. means wit-

hout lot number and bibliographic loca-

lity respectively.

The variables measured in the shells

are (Fig. 1): 1. length (LONG) 2. maximum

diameter (DMAX,) 3. height of body whorl

(LEC), 4. minimum diameter (DME), 5.

PÉREZ AND LÓPEZ: Morphology and distribution of Bulimulus corneus in Nicaragua

aperture length (LAB), 6. aperture width

(AAB), 7. spire length (LEP), 8. spire width

(AEP). All measurements were made in

adult specimens.

We calculated various descriptive

statistics for the measured variables, in or-

der to give a morphological description

of the samples. We also used a Principal

Component Analysis (PCA) to explore the

variability among populations.

RESULTS AND DISCUSSION

Description: Shell (Fig. 2): Shell thin,

spirally striate, corneus to brown, so-

mewhat translucent, showing through the

dark bands that stipple the mantle. Pro-

file bulimoid-conic. Apex obtuse; proto-

conch typically bulimoid with sculpture

of punctures in an irregular decussate pat-

tern; whorls 5.5 to 6. Aperture ovate, mar-

gin thin, sharp, umbilicus narrow. Mea-

surements taken on the shell are presen-

ted in Tables I and II.

Genitalia (Fig. 3): Penis with wide she-

ath, dilated in its central part, and reaching

to more than one half of the phallus. Epip-

hallus approximately half as wide as pe-

nis. Flagellum thinner, approximately one

half the phallus length. Sperm conduct

thickened at mid-center, ending at globose

spermatheca distally. Vagina more or less

fusiform, slightly longer than the penis,

and ?/3 the width.

We have found that shell dimensions

are quite variable within (Table I) and bet-

ween populations (Table II), as also men-

tioned by PILSBRY (1897). For this reason,

and because of the small total sample size

(n= 44) studied from all populations (11)

we have not considered the taxonomical

implications of the variability. However,

it should be mentioned that shell length

(LONG) and height of body whorl (LEC)

display the highest variances of all varia-

bles considered (Table II). Shell length is

always one of the variables on which des-

criptions are based. We recommend cau-

tion in the use of either variables for a ta-

xonomic characterization of the species.

It must be pointed out that THOMPSON

(1967) invalidated various subspecies of

the closely-related Bulimulus unicolor So-

werby, 1833, believing them to be varia-

tions related to climatic conditions. In this

paper he considered shell length (LONG),

maximum diameter (DMAX) and two ot-

her variables.

In the PCA made from conchological

variables it is possible to see the marked

scatter of the specimens (Fig. 4). Within

the plot, there is a segregation of six indi-

viduals from the populations of El Gua-

yabo, Granada (1), Xiloá, Managua (2),

and Las Lajas, Rivas (3). It is interesting

to notice that the other specimen from Ri-

vas is located within the cloud of points.

The only specimen considered from the

Nicaraguan Atlantic slope (6), can be ob-

served between the cloud of points and

the six individuals previously mentioned.

Another three specimens from Ocotal

(9) segregate towards the lower right cor-

ner of the scatterplot. These specimens, as

the previous six, have conchological fea-

tures very much like the ones from other

populations (see Table 1), although the

ones from Ocotal have larger sizes.

In Table III, it can be seen the contri-

bution made by each principal compo-

nent to total variance. Components I

(70.77%) and Il (18.15%), comprise the

major quantity of total variance (88.92%).

The absence of negative signs (Table IV)

among the eigen values obtained for

component l, also with the larger contri-

bution (70.77%), allow us to presume

that it is related to size and Il is related to

shape. Thus, differences among popula-

tions would be apparently due to size

rather than shape; and it is known that

size is usually influenced by ecological

factors, and consequently is highly corre-

lated with local environmental condi-

tions (BEROVIDES, 1988).

In their genitalia (see Figure 3), the

individuals from the populations of

UCA Campus (Managua Department)

and Ocotal (Nueva Segovias Depart-

ment) share the same external morpho-

logy and show only small differences in

size of the structures. However, a more

detailed anatomical analysis, including

an analysis of the internal anatomy of

the genital ducts (v. g. penis), of the

Ocotal population is required when

fresh material is available.

Iberus, 15 (2), 1997

PÉREZ AND LÓPEZ: Morphology and distribution of Bulimulus corneus in Nicaragua

Figure 3. Bulimulus corneus. Genitalia. A: specimen from Ocotal; B: specimen from Campus UCA;

pr: prostate; p: penis; f: flagellum; a: atrio; (sp: spermatheca)= bc: bursa copulatrix; s. ov: spermo-

viduct; sp. d: spermathecal duct; ep: epiphallus; v: vagina. Scale bars 1 mm.

Figura 3. Bulimulus corneus. Aparato genital. A: ejemplar de Ocotal; B: ejemplar del Campus UCA;

pr: próstata; p: pene; f: flagelo; a: atrio; (sp: espermateca)= bc: bolsa copulatriz; s. 0v: espermoviducto; sp.

d: conducto espermático; ep: epifalo; v: vagina. Escalas 1 mm.

Distribution: Fifty three localities

for B. corneus have been added to those

previous recorded. They distributed

over 12 Departments in the three natural

regions that comprise the country (Fig.

5, Table V).

CONCLUSIONS

The distribution map gives a clear

idea of B. corneus distribution in Nicara-

gua. As pointed out by JACOBSON (1968),

a fairly continuos distribution can be

seen among the samples, suggesting that

absence in other areas is due to lack of

sampling, and that B. corneus is wides-

pread in the country.

B. corneus has a very wide ecological

tolerance, ocurring from low altitude to

more than 2000 m. The species inhabits a

remarkable number of different microha-

bitats, including soil with herbs, soil

with litter, tree trunks, logs, stones, walls

of ruined houses, etc. The wide geograp-

hical distribution of the species, can pro-

(Left page). Figure 2. Bulimulus corneus. Shell morphology. A: Las Canoas (length 12.8 mm, dia-

meter 7.0 mm); B: Campus UCA (length 10.6 mm, diameter 6.7 mm), C: Ocotal (length 19.9

mm, diameter 11.35 mm).

(Página izquierda). Figura 2. Bulimulus corneus. Morfología de la concha. A: Las Canoas (1 longitud

12,8 mm, diámetro 7,0 mm); B: Campus UCA (longitud 10,6 mm, diámetro 6,7 mm); C: Ocotal

(longitud 19,9 mm, diámetro 11,35 mm).

Iberus, 15 (2), 1997

Table I. Variables measured considering each sample separately (X= average, S= standard deviation).

Abbreviations as in Figure 1.

Tabla I. Variables medidas considerando los ejemplares de cada muestra independientemente (X= media,

S= desviación standard). Abreviaturas como en la Figura 1.

LOCALITIES VARIABLES

LONG DMAX LEC DME LAB AAB LEP AEP

Xiloá [n= 6)

XK 10.93 6.70 A 6.13 SS] 3.87 4.38 4.85

Mín 10.10 5.80 DES) 5.30 Doll 2.95 2.9 ARS

Máx 11.70 O 8.6 6.5 6.0 4.7 5.8 5.4

S 0.62 0.61 137 0.46 0.33 0.63 1.29 0.35

Apoyo (n= 10)

X 11.62 8.19 8.82 6.31 DL EZ 3.6 4.49

Mín 10.0 6.3 7.8 DS) 4.6 2.4 2.4 37

Máx 13.5 8.0 10.0 YES 6.0 3.9 4.75 DS

S 1.04 052 0.76 0.48 0.42 0.42 0.64 0.45

Asososca (n= 4)

X 1237 a 9.27 7.0 DS 3.6 3.82 4.9

Mín 12.0 6.5 8.4 DA 45 ZE 399 4.3

Máx 13% 7.8 9.6 7.1 7 3.8 4.1 5.0

S 0.59 0.17 0.42 0.1 0.35 0.21 0.41 0.16

Las Canoas [n= 5)

XK 18202 SIS 9.46 6.6 9 3.62 4.33 AED

Mín 1285 6.8 9.0 6.3 AZ SZ 4.2 4.7

Máx 13.8 DL, 10.0 EZ 5.8 4.3 4.6 DS

S 052 057 0.42 0.35 0.22 0.42 0.17 0.26

El Guayabo (n= 5)

XK 10.65 DS DIS 5.95 IDO 4.85 IS 5.05

Mín 10.1 6.0 DS 5.8 Del 4.8 4.6 4.7

Máx 1912 6.3 6.0 6.1 6.0 4.9 6.0 5.4

S 0.78 0.2 035 0.22 0.64 0.22 0.99 0.5

Las Lajas (n= 2)

X 14.25 YES 6.82 6.7 6.35 ASIS SY 5.8

Mín 12.9 6.5 6.3 6.0 Dl 3.8 EZ DES)

Máx 1576 8.1 SS 7.4 7.0 4.5 00 6.1

S 1:91 1.18 0.74 0.99 0.92 0.5 0.18 0.65

Tepeyac (n= 2)

X 11.25 Del 8.75 5.85 4.8 SD) SS) 45

Mín 11.0 6.7 8.6 0) 4.7 2 SS) 4.4

Máx 155 YES 8.9 6.7 4.9 3.9 3.4 4.6

S 0.35 0.42 0.21 1162 0.14 0.5 (0) 0.14

Ocotal (n= 4)

XK 1712 9.52 237 8.72 ES, 4.92 5.40 6.16

Mín 13.6 8.0 10.5 8.0 7.4 4.2 3.8 ZO

Máx 19:9 11S5 11212 10.0 8.4 EZ 6.9 AZ

S ZO, AO) 1.54 0.87 0.48 0.5 1.34 0.80

Campus UCA (n= 3)

XK 11.56 AZ 8.78 (3 5.03 3.46 SSZ 4.47

Mín 10.6 6.7 8.3 6.0 4.9 3.0 2.8 4.1

Máx 12.8 7.8 9.55 2 5. 1 3.8 3.8 4.9

S 1.12 0.55 0.63 0.64 0.12 0.42 0.50 0.41

Laurel Galán (n= 3)

XK 12.45 7.6 8.95 6.7 6.02 IEA 7 4.6

Mín 15144 AO 8.35 6.0 4.9 3.0 2.8 4.1

Máx 18389 8.1 9.5 YES 6.35 3.6 4.0 5.0

S 0.98 0.55 05% 0.65 0.45 0.12 0.29 0.46

PÉREZ AND LÓPEZ: Morphology and distribution of Bulimulus corneus in Nicaragua

Table IL. Variables measured considering all samples pooled. (X= average; Min: minimum value;

Max: maximum value; S= standard deviation). Abbreviations as in Figure 1.

Tabla 11. Variables medidas considerando los ejemplares de todas las muestras agrupadas. (X= media;

Min: valor mínimo; Max: valor máximo; S= desviación standard). Abreviaturas como en la Figura 1.

Variables Xx Min Max S

LONG 12.40 10 19.90 2.02

DMAX JESZ Date INESS 0.99

LEC 8.78 4.1 14.20 1.89

DME 6.59 DO) 10.00 0.89

LAB 5.64 4.5 8.40 0.87

AAB SINO) DIA DENO) 0.67

LEP AZ ZA 6.90 0.97

AEP 4.86 37 7.20 0.64

Axis ll

Axis |

Figure 4. Axis I and II of the Principal Component Analysis. Each number represents a sample. 1:

El Guayabo; 2: Xiloá; 3: Las Lajas; 4: Asososca; 5: Campus UCA; 6: Loma del Mico; 7: Laurel

galán; 8: Tepeyac; 9: Ocotal; 10: Apoyo; 11: La Ceiba.

Figura 4. Ejes 1 y 11 del Análisis de Componentes Principales. Los números corresponden a las muestras.

1: El Guayabo; 2: Xiloá; 3: Las Lajas; 4: Asososca; 5: Campus UCA; 6: Loma del Mico; 7: Laurel galán;

8: Tepeyac; 9: Ocotal; 10: Apoyo; 11: La Ceiba.

Iberus, 15 (2), 1997

Table III. Percentage of variance explained by each one of the principal components. Abbreviations

as in Figure 1.

Tabla III. Porcentaje de varianza explicado por cada uno de los componentes principales. Abreviaturas

como en la Figura 1.

Component Percent of cumulative number Variance percentage

LONG 70.77 ANOTA

DMAX SMS 88.92

LEC 4.72 93.64

DME EDS, 95.88

LAB 2.06 SISI

AAB 0.89 98.84

LEP 0.71 SEDO

100.00

AEP 0.44

Table IV. Eigen values obtained with Principal Component Analysis, considering axes L, Il and III.

Abbreviarions as in Figure 1.

Tabla IV. Valores propios obtenidos con el Análisis de Componentes Principales, considerando los ejes l,

11 y 1. Abreviaturas como en la Figura 1.

Components

LONG 03993

Variables

DMAX 0.3726

LEC 0.3072

DME 0.3872

LAB 0.3902

AAB 0.2860

LEP 0.2861

Íl 111

0.1351 0.2987

0.3116 -0.0867

-0.4996 0.0703

-0.2181 -0.2486

0.0308 -0.0825

0.4631 0.7271

O3S339 0.4999

0.2948 0.2258

AEP 0.3756

bably be explained by the numerous mi-

crohabitats that it is capable of filling.

Considering at the same time the

morphological variability and the ecolo-

gical range of this species, WOLDA's

(1970) stament comes to mind, that va-

riation should be understood in terms of

the possibilities of survival in natural po-

pulations, and not only as a biologically

isolated fact about the ecology of species.

Regarding the presence of B. unicolor .

in Nicaragua, Tate's reports were not re-

cognized by MARTENS (1890-1901), or by

PiLsBRY (1897), and may have had their

origin in the marked variability of B. cor-

neus. We think that only B. corneus is

found in Nicaragua.

PiLsBRY (1897), mentioned Greytown

(RAAN: Autonomous Region of the

North Atlantic), and later Fluck (1900)

gave Bluefields (RAAS: Autonomous Re-

gion of the South Atlantic) as localities

for B. unicolor in Nicaragua. More re-

cently, BREURE (1979) quoted Perico Is-

land in the Bay of Panamá as the only lo-

cality in Central America.

In the last two years we have collec-

ted specimens from Bluefields, one of

PÉREZ AND LÓPEZ: Morphology and distribution of Bulimulus corneus in Nicaragua

160

130

Figure 5. Bulimulus corneus. Distribution in Nicaragua, in UT'M notation of 100 Km?.

Figura 5. Bulimulus corneus. Distribucion en Nicaragua, en notación UTM de 100 Knr.

the localities for B. unicolor, and two ot-

her nearby localities (Las Delicias and

La Fonseca) (see Table IV). This material

agrees well with the description of B.

corneus.

We are therefore led to think either

that in these localities both species live

sympatrically, and we have not so far co-

llected B. unicolor, or that B. unicolor does

not occur there at all.

In view of the fact that in the revision

of the subfamily BREURE (1979) did not

include distribution data for B. unicolor

in Nicaragua, and taking into account

the morphological variability that

THOMPSON (1967) has shown in both B.

unicolor, and B. corneus, as mentioned by

Pilsbry and also studied by us, we have

decided for the moment to accept that

the latter species is the only one that oc-

curs in Nicaragua.

ACKNOWLEDGEMENTS

We are grateful to Dra. Ana Isabel

Puente (University of Basque Country,

Faculty of Sciences, Department of

Animal Biology and Genetics, P. O. Box

644, 48080 Bilbao) for critical revision

and advice about the paper. Also to Lic.

Zamira Guevara (Central America Uni-

versity, Department of Ecology and

Natural Resources, P. O. Box. 90,

Managua, Nicaragua) who collected and

processed part of the studied material

with us. The great encouragement to

our work, given by Ing. Luis Fiallos,

Dean of the Faculty of Agricultural

Sciences, is also to be mentioned and

gratefully acknowledged. The sugges-

tions made by two anonymous revie-

wers contributed to largely improve the

manuscript's scientific quality.

Iberus, 15 (2), 1997

Table V. List of new localities for B. corneus Sw., in Nicaragua. S= number of shells, Sa= specimens

in alcohol. The samples used for statistical purposes are marked with an asterisk. Bibliographic loca-

lities are considered as Bib under “Lot number”. w. l. n. means: “without lot number”.

Tabla V. Lista de nuevas localidades para B. corneus Sw., en Nicaragua. S= número de conchas, Sa=

especímenes en alcohol. Las muestras usadas con propósitos estadísticos aparecen marcadas con con un aste-

risco. Las localidades bibliográficas aparecen con la abreviatura Bib en “Lot number”; w. l. n. significa:

“sin número de lote”.

Code Lot Localities

number

Coordinates

Geographical

BOACO Department

1 92:19 "El Sácal

CARAZO Department

2 90:18 La Baronesa

CHINANDEGA Department

3 Bib Realejo

CHONTALES Department

4 92:14 Punta Mayal

5 88:30 Nueva Guinea

ESTELI Department

6 93:09 Estanzuela

GRANADA Department

7) 91:26 Tepeyac

8 92:01 Aguas Calientes, Cocibolca

9 w.Ln Isleta de Ken

10. 92:12 Laguna Blanca

11. 95:72 El Guayabo

12 ER933lO Apoyo

LEON Department

13 90:05 Asososca

14 90:06 Laguna Monte, Galán

15 88:26 Salinas Grandes

MANAGUA Department

16 88:21 Villa Carmen

17 88:28 Las Sierritas

18 90:07 Xiloá

19 SES Asososca

ZOO Las Mercedes, Lago Xolotlán

ZII S SY Apoyeque

22 94:47 San Francisco Libre

ZION Los Placeres, Km 63

24 95:04 Km 66.8 Carr. Matagalpa

12*33'10” N, 85"33'30" W

12%10'N, 8717 W

123927" N, 83918” W

11%52'20" N, 85*26' W

11%43' N, 84*57' W

13%14'04” N, 862216" W

11%52.5' N, 85"59.5' W

11%52' N, 85"55'40" W

11%51'41” N, 85"53'40" W

11%46'15 N, 85%57'45" W

1158" N, 8559 W

1155 N, 85%57'45" W

12%26' N, 86"40' W

12%16'12” N, 86"30'4” W

1219 N, 8616' W

123 N, 8616' W

12%14' N, 86"46' W

121811” N, 85198" W

12%9'30" N, 8610' W

12%15' N, 8621 W

12*30'12” N, 86%17'40" W

12*33' N, 86"3'41” W

12%29'06" N, 86%04'17" W

UTM

16PFJ 58

16PFJ 72

16PDJ 78

16PFJ 71

16PGH 79

16PEK 73

16PFJ 01

16PFJ 11

16PFJ 10

16PFJ 11

16PFJ 01

16PEJ 37

16PE) 47

16PEJ 25

16PEJ 74

16PEJ 73

16PE) 74

16PEJ 74

16PEJ 84

16PEJ 74

16PEJ 78

16PFJ 08

Material

examined

125

25(*)

Ss(*)

105+55a

As")

155

10S+4Sa

ós(*]

5Sa

1Sa

PÉREZ AND LÓPEZ: Morphology and distribution of Bulimulus corneus in Nicaragua

Table V. Continuation. *

Tabla V. Continuación.

Code Lot Localities

number

25 95:06 Km42.5 Carr. Matagalpa

26 95:07 Las Canoas

LOSA Las Canoas

28 95:26 Mateare

ZII Carr. Tipitapa-Masaya, Km 43

SOI La Polvosa

31 95:28 Sierra de San Andrés

SS: S Nandayosi

33 95:32 Nandayosi, cerca del rio

34 95:34 LosFilos de Guajachillo

SIA 38 San Bartolo

SOI El Conchital

37 95:40 Hacienda “El Apante”

38 95:44 Hacienda “El Callao”

39 95:45 Samaria

AOS El Tamarindo

41 95:59 El Platanal

42 95:64 Carr.Sur, Km 15.5, INCAE

ASI 72 Campus UCA

MATAGALPA Department

44 92:83 Ciudad Darío

AS 2:88 Fuentepura

NUEVA SEGOVIAS Department

d6 94:59 Ocotal

RIVAS Department

AT Z3 9 Río Las Lajas

RIO SAN JUAN Department

48 92:24 La Toboba

49 92:25 El Castillo

50 w.ln San Carlos

51 92:26 Laurel Galán

Coordinates

Geographical

12%21'02” N, 8602'58" W

12%19'00” N, 80%00'07” W

121905" N, 855922" W

121410" N, 86%25'48" W

120948” N, 86%00'07" W

121324” N, 86"24'59” W

12%10'28” N, 86*24'35"” W

12%06'43” N, 86"31'14” W

12%06'54” N, 86"30'16" W

12%08'33" N, 86"24'33 W

11%54'58' N, 86"33'05" W

11%54'18” N, 86"33'43" W

11%57'48” N, 862915" W

12%01'42” N, 8620'07" W

MESAUSAN, IRSA A

12%29'37" N, 86"05'01” W

12%27'06" N, 86%05'02” W

10%03'13” N, 86"18'33" W

12%07'30" N, 861613" W

12%43'50" N, 861153" W

1258" N, 8655" W

13%36'35" N, 862818” W

112 N, 85 W

11%8' N, 84*57'50" W

11720” N, 8424 W

12720" N, 86"46' W

12%7' N, 86"46' W

RAAS (AUTONOMOUS REGION OF THE SOUTH ATLANTIC)

52 93:23C Las Delicias

53 93:23B La Fonseca

54 94:19 Loma del Mico

55 Bib Bluefields

12%16'12” N, 8352/50" W

12%15'36" N, 83"58'48" W

12%4'24” N, 83%47'42” W

12% N, 834418" W

RAAN (AUTONOMOUS REGION OF THE NORTH ATLANTIC)

Bib

San Juan Castillo

14%24' N, 8354'54" W

UTM

16PFJ 06

16PFJ 16

16PFJ 06

16PEJ 65

16PEJ 64

16PEJ 53

16PEJ 42

16PEJ 41

16PEJ 52

16PEJ 72

16PEJ 42

16PEJ 98

16PEJ 97

16PEJ 73

16PEJ 74

16PEK 90

16PFK 13

16PEL 51

16PFH 35

16PGH 72

16PGH 81

16PGH 43

17PJP

17PJR

Material

examined

155

108

118

325

135

ÁS

155

145

15S

3Sa(*)

135

55")

251")

351")

Iberus, 15 (2), 1997

REFERENCES

BEROVIDES, V., 1988. Orden y diversidad en el

mundo viviente. Editorial Cintífico- Técnica,

La Habana. 108 pp.

BREURE, A.S. H., 1979. Systematics, phylogeny

and zoogeography of Bulimulinae. Zoologis-

che Verhandelingen, 168: 1-205.

FLuck, W. H., 1900. Shell collecting in the Mos-

quito Coast. The Nautilus, 14: 94.

JACOBSON, M. K., 1968. On a collection of te-

rrestrial mollusks from Nicaragua. Nautilus,

81: 114-120.

MARTENS, E. V., 1890-1901. Biologia Centrali-

Americana. Land and Freshwater Mollusca. Tay-

lor and Francis, London. 706 pp.

PÉREZ, A. M. AND LÓFPEZ, A., 1995. New data on

the morphology and the distribution of Bu-

limulus corneus Sowerby, 1833 (Gastropoda:

Pulmonata: Orthalicidae). In Guerra, A., Ro-

lán, E. and Rocha, F. (Eds.): Abstracts of the

Twelfth International Malacological Congress,

Vigo. Unitas Malacologica and Instituto de In-

vestigaciones Marinas, Vigo: 396-398.

PiLsBRY, H. A., 1897. Manual of Conchology. 2nd

series, vol. 11, pp. 65-144.

TATE, R., 1870. On the land and freshwater mo-

llusca of Nicaragua. American Journal of Con-

chology, 5: 151-162.

THOMPSON, F. G., 1967. The land and freshwa-

ter snails of Campeche. Bulletin Florida State

Museum, 11(4): 221-256.

WOoLDA, H., 1970. Ecological variation and its

implications for the dynamics of populations

of the landsnail Cepaea nemoralis. Proc. Adv.

Study Inst. Dynamics Numbers Popul.: 98-108.

O Sociedad Española de Malacología —————— Iberus, 15 (2): 25-34, 1997

Phenological patterns and life history tactics of Helicoidea

(Gastropoda, Pulmonata) snails from Northern Greece

Patrones fenológicos y estrategias de vida en Helicoidea (Gastropoda,

Pulmonata) del Norte de Grecia

Maria LAZARIDOU-DIMITRIADOU and Stefanos SGARDELIS*

Recibido el 8-[-1996. Aceptado el 31-VIL1996

ABSTRACT

The present study mainly concerns with the differences in the biological cycles and strategies

adopted by different Helicoidea snail species. In Northern Greece the climatic conditions are

not very uniform. Some snails breed during the same period as in Northern Europe but most

breed in autumn, as species from Southern Europe do. Breeding may take place in all sea-

sons except in winter, and seems to be species-specific. Long-lived snails of big size differ

from shortlived species of small size as to the time of their breeding period. The climatic con-

ditions affect the time of the breeding season and their whole life cycle and phenologies. En-

vironmental variables in Northern Greece are strongly seasonal and thus Helicoidea snails

exhibit predictable oscillations in their activity patterns, which can be interpreted by the de-

mographic response of the populations. Terrestrial snails seem to follow two different pheno-

logic curve types: the semelparous and short-lived species populations show a more stable

phenological pattern than the biennial and pluriennial ones, who mature after the first year of

their lives, being more plastic trying to face the climatic differences from one year to another.

RESUMEN

El presente estudio trata de las diferencias en los ciclos biológicos y a las estrategias adopta-

das por diferentes especies de Helicoidea. En el norte de Grecia, las condiciones climáticas

no son muy uniformes. Algunas especies crían durante el mismo periodo en que lo hacen en

el N de Europa, pero la mayoría lo hacen en otoño, como sucede en especies del S del con-

tinente. La cría puede tener lugar durante casi todas las estaciones, excepto el invierno, y el

periodo parece ser específico para cada especie. Las especies longevas y de gran talla di-

fieren de las de pequeño tamaño y vida más corta en la duración de su ciclo de cría. Las

condiciones climáticas afectan al momento de la temporada de cría y a todo su ciclo vital y

fenología. Las variables ambientales son fuertemente estacionales, así que aparecen osci-

laciones predecibles en los patrones de actividad, que pueden ser interpretadas por la res-

puesta demográfica de las poblaciones. Las babosas parecen seguir dos tipos de curvas fe-

nológicas distintas. Las especies semelpáricas y de corta vida muestran un patrón fenológico

más estable que el de especies bianuales y prurianuales, que maduran tras el primer año de

vida y son más flexibles a la hora de enfrentarse a las diferencias climáticas interanvales.

KEY WORDS: Biological cycle, phenology, Northern Greece, Helicoidea, Helix, Eobania, Helicella, Monacha,

Bradybaena.

PALABRAS CLAVE: ciclo biológico, fenología, N Grecia, Helicoidea, Helix, Eobania, Helicella, Monacha, Bradybaena.

* Departments of Zoology and Ecology, School of Biology, Aristotle University of Thessaloniki, 54006

Thessaloniki, Macedonia, Greece.

Iberus, 15 (2), 1997

INTRODUCTION

Greece has a Mediterranean climate

which is differentiated mainly along a

northern-southern gradient. In Northern

Greece climate is transient from

Mediterranean (mostly coastal areas) to

temperate (inland areas). A typical cha-

racteristic of this climate type is the

coincidence of high temperatures and

low precipitations during summer (las-

ting from June to October). The wet sea-

son is divided by a cold winter which is

milder in the coastal areas. Drought is a

strong agent controlling population dy-

namics and activity of most soil inverte-

brates as it imposes a pause in most

physiological activities. Low temperatu-

res during winter are also important for

population dynamics and activity as

they impose hibernation in some of the

invertebrates e. g. terrestrial gastropods.

So observed discontinuities in popula-

tion development during the transition

from the favourable to the unfavourable

seasons and vice-versa may be attribu-

ted to environmental thresholds.

Although the association between

climate and life history phenomena is

self evident, it can vary among terres-

trial molluscs, even between popula-

tions of the same species. Phenology

reflects certain aspects of the demo-

graphy of a population, that is the

timing of its life cycle characteristics in a

given environment. The classification of

phenological patterns into categories or

types (WOLDA, 1988) is better by using

phenological models (VAN STRAALEN,

1982; STAMOU, ASIKIDIS, ARGYROPOULOU

AND SGARDELIS, 1993). Using a phenolo-

gical model, complex phenograms can

be classified into types considering their

skewness, curtosis, phase and period.

The aim of the present study was to

find out whether terrestrial gastropods

adopt a general phenological pattern if

* they are differentiated according to their

origin, or their biotopes (inland and

coastal areas) or life spans. The present

study is a part of an extensive research

done on the distribution and ecology of

Helicoidea gastropods in Northern

Greece (LAZARIDOU-DIMITRIADOU, 1981,

1995; LAZARIDOU-DIMITRIADOU AND

KATTOULAS, 1981, 1985, 1991; STAIKOU,

LAZARIDOU-DIMITRIADOU AND FARMA-

KIS, 1988; HATZUOANNOU, ELEUTHERIA-

DIS AND LAZARIDOU-DIMITRIADOU, 1989;

STAIKOU, LAZARIDOU-DIMITRIADOU AND

PANA, 1990; STAIKOU AND LAZARIDOU-

DIMITRIADOU, 1990, 1991).

MATERIALS AND METHODS

Data used derived from monthly

quantitative samplings of Helicoidea

snails from different parts of Northern

Greece. The following species were

studied: Family Bradybaenidae, Brady-

baena fruticum (Múller, 1774); Family

Helicidae, Cepaea vindobonensis (Férus-

sac, 1821), Eobania vermiculata (Muller,

1774), Helix lucorum Linnaeus, 1758,

Helicella (Xerothracia) pappi (Schút, 1962),

Helix figulina (Rossmássler, 1839), Helix

pomatia rhodopensis Kobelt, 1906, Theba

pisana (Múller, 1774); Family Hygromii-

dae, Cernuella virgata (Da Costa, 1778),

Monacha cartusiana (Múller, 1774), Xero-

lenta obvia (Menke, 1828), Xeropicta

arenosa Ziegler, 1836, Xerotricha conspur-

cata (Draparnaud, 1801). Sampling

lasted two or four years depending on

the species and their life span (LAZARI-

DOU-DIMITRIADOU, 1981, 1995; LAZARI-

DOU-DIMITRIADOU AND KATTOULAS,

1981, 1985, 1991; STAIKOU ET AL., 1988;

STAIKOU ET AL., 1990; STAIKOU AND

LAZARIDOU-DIMITRIADOU, 1990, 1991).

Details regarding the sites and the sam-

pling procedures are given in previous

studies on these species (LAZARIDOU-

DIMITRIADOU, 1981, 1995; LAZARIDOU-

DIMITRIADOU AND KATTOULAS, 1985,

1991; STAIKOU ET AL., 1988). Ombrother-

mic data for Northern Greece from 1980

to 1990 are given in Figure 1. Data were

provided by Mahairas P., Professor of

Climatology from the Aristotle Univer-

sity of Thessaloniki.

Fischer's exact test for independence

in 2 x 2 contingency tables (ZAR, 1984)

was used for comparisons between the

different categories, e. g. snails with

autumnal and vernal-estival reproduc-

tive periods, long-lived (> 3 years) and

LAZARIDOU-DIMITRIADOU AND SGARDELIS: Phenology of Helicoidea in N Greece

JF MA M

Precipitation (mm)

Prec. (mm)

A SON D

Months

Figure 1. Ombrothermic diagram from Northern Greece. Temperature: open squares; Precipitation:

solid rhombus.

Figura 1. Diagrama ombrotérmico del Norte de Grecia. Temperatura: cuadrados abiertos; Precipitación:

rombos sólidos.

short-lived snail species (up to 3 years),

large (largest shell diameter > 25 mm)

and small sized snails.

The phenological pattern of Helicoi-

dea species was studied by using the

phenological model applied for the

study of microarthropods (STaAMOU ET

AL., 1993). In short, in this model when

asymmetries and discontinuities are dis-

played the scales of the time-axis were

adjusted. Changing time scales results

in the definition of a new variable

termed ecological time (ET), which is a

function of a standard clock time

(STAMOU ET AL., 1993). This technique is

based upon the following considera-

tions: 1) the timing of a population in

the field is determined by the sequence

of demographic events and/or beha-

vioural characteristics (i. e. migratory),

and 2) the rate of the demographic

events depends on the fluctuations of

environmental variables.

In this model it is assumed that the

phenology of any population inhabiting

a seasonal environment can be descri-

bed by a symmetric periodic curve:

F(ET) = EXP(a-+b x COS(27 x (ET-9/T)) (1)

where the independent variable ET,

termed Ecological time, is a function of

standard clock time (ST), ET = f (ST). In

the course of standard time, ecological

time is going faster during periods of

sharp changes in abundance and slower

during periods of abundance stability.

Thus, the proposed equation describing

the length of the Ecological time unit

(AET) as a function of Standard time ST

is:

AET= f (ST) = EXP(or+b1 x COS(27 x

(ST-p1)/T1)) (2) *

The model has eight parameters of

which the period T and the phase q of

the phenogram, as well as period T1 and

the phase q1 of the function relating ET

to ST, are the most important. The

period T and the phase q of the pheno-

grams are expressed formally in ET

units. For convenience they could be

expressed in ST units (as T' and q”) by

using equation (2) for the calculation of

Iberus, 15 (2), 1997

the Standard time T” or q” which corres-

ponds to T or € units in the Ecological

time-scale (see STAMOU ET AL., 1993: fig.

1). For the comparison of phenograms

two more parameters can be derived: a)

an estimation of the sharpness (curtosis)

of the phenogram C=(R2-R1)/T”, where

(R2-R1) is the time interval around the

phase q”, during which the abundance is

above overall mean, and b) an esti-

mation of the skewness of the pheno-

gram S= (0'-Qm)/T' where qmis the time

(in ST units) when the abundance of the

population is at minimum. Thus, pheno-

grams displaying a peak at p'-qm= T'/2

(half period) is symmetric (S= 0.5), phe-

nograms displaying a peak soon after

the minimum (S< 0.5) are positively

skewed, and phenograms with S> 0.5

are negatively skewed. The model was

fitted on log-transformed census data.

For ETi= f (STi) given as a time vector

and for a given set of q and T, the para-

meters a and b were estimated by least-

square regression.

RESULTS

The model fitted to census data for

snail populations sampled at monthly

intervals from different areas are shown

in Figure 2. The values of the most

important parameters of the fitted phe-

nological model are given in Table 1.

In all but two examined cases the

phenological pattern was strongly sea-

sonal (Fig. 2), with a more or less 1 year

periodicity apart from Monacha cartu-

siana which seems to display a six

months periodicity at least during 1984

(Table I). Abundance of Bradybaena fruti-

cum and Eobania vermiculata fluctuated

almost randomly throughout the year.

Population densities of the different

species do not exhibit phase synchroni-

zation. Even different populations of the

same species do not always exhibit a

peak density at the same time of the

year. Furthermore even the same popu-

lation displays a phase instability

during successive years of study. For

instance Xerolenta obvia from Paleokas-

tro peaked either in January or in April

(q, in Table I). X. obvia from Karvali

peaked in July. Helix lucorum displayed

a similar interannual instability. In both

cases the shift in phase seems to be asso-

ciated with an unexpected change of the

weather, an extended dryness (STAIKOU

ET AL., 1988: fig. 2) which provoked an

overall decline of the population density

(Fig. 2).

The phase expressed as months after

minimum population densities (qm)

(Table I) is a measure of the rate of

population increase from absolute

minimum to peak densities. Qm might

have lower values when the time

needed for a species to mature is short.

Minimum (Qm values were estimated for

X. obvia in Paleokastro, the 1st and the

third year of study, indicating a rapid

population growth which occurs in a

period of about 3 months (Fig. 2). In

Karvali, where X. obvia gets mature in 2

years, Qm value was larger (Table 1), as

was the case with M. cartusiana in 1985.

Helicella (Xerothracia) pappi snails that

need 2-3 years to mature exhibited

larger Qm values, and H. lucorum snails

- which need 3-4 years exhibited the

largest value of all except in 1983.

Species that mature in one year,

exhibit a very rapid population growth

just after the adverse period, which is

winter, and consequently they are posi-

tively skewed. Species that mature in 2-

3 years are negatively skewed (Table Il,

Fig. 3). In this case the phenological

pattern may be positively skewed or

symmetrical but the population remains

active for longer periods and the pheno-

logical pattern is platycurtic (Table II,

Fig. 3). Whereas species that mature in

one year usually display a leptocurtic

phenology (Table Il, Fig. 3). The only

species population that showed both a

negatively and a positively skewed phe-

nological pattern was H. lucorum.

The positively skewed phenograms

are leptocurtic when they concern snail

species populations that mature in one

year whereas they may be slightly platy-

curtic when they live in regions where

favourable conditions last longer as is

the case of Xeropicta arenosa Ziegler in

Edessa. Pluriennial snail species popula-

LAZARIDOU-DIMITRIADOU AND SGARDELIS: Phenology of Helicoidea in N Greece

”,* Helix lucorum

1) Edessa

Population density (ind./m2)

JFMAMJJASONDJFMAMJJASONDJFMAMJJASONDJFM

83 84

14 A

a o Monacha cartusiana

42 o

E A AN Edessa

E 10 / Ñ p N

= (o) ¡MORO NS

Dos 7 Ñ Y 3 :

a / A ; Y

5 Wa 4 ES

sS. U ó e ás NS e

5 ó > a) D

(e) o

A S

a o

5] 2

[aY

0 AA]

MJJASONDJ]FMAMJJASONDJFMAMJIJJ

6 Xeropicta arenosa

Edessa

Population density (ind./m2)

JEMAMJJASONDJIJFMAMJJASONDJFMAMJJASONDIJFMA

83 84

Months

] Xerolenta obvia

os Paleokastro ¿$

Lo 1

40 " Á Mn

¿ LO

y X | :

o |

204 |! eN l No

E so S DA i je]

Ea] ES PS ES A , o

HAL

ONDJFMAMJJASONDJFMAMJJASONDJFMAMIJAS

180

160 o

Xerolenta obvia

140 / V .

/ MN Karvali

120 / EN

' Va

10) P y

80 i X .

í ye dl

60 la A >

90 A ,

, »

20 Ñ JÓ

$ le

O AAXA>->>—- -á ->>--- 91

mM y y AÑES OON D ñ F M A M y J A S

89 90

Helicella (Xerothracia) pappi

IN Philippi

MAMJJASONDJFMAMJJASONDJFMAMJJASONDJ

Months

Figure 2. Abundance variations of Helicoidea snails from Northern Greece. Line: model estimates;

asterisks: observations.

Figura 2. Variaciones de la abundancia de caracoles helícidos del Norte de Grecia. Línea: estimaciones

del modelo; asteriscos: observaciones.

tions usually exhibit platycurtic pheno-

logical patterns, too, since their adults

diapause and hide in the soil or under

plants and do not emerge massively.

Negatively skewed and symmetric phe-

nograms are usually platycurtic.

It seems that there are no negatively

skewed-leptocurtic species except for M.

cartusiana that is negatively skewed and

slightly leptocurtic.

Maximum activity duration is about

the same for different populations of the

same species or for the same population

during successive years of study (Table

[, MA column).

In Northern Greece long-lived

species are of big size and short-lived

are of small size (x?= 9.983, P= 0.001).

Additionally, short-lived species breed

in autumn whereas long-lived species

may breed in autumn or vernal-estival

period (x?= 4.261, P= 0.039) (Table III).

In Northern Greece bigger helicids may

breed during the vernal-estival or

Iberus, 15 (2), 1997

Table I. The estimated parameters of the fitted model in standard time units (months).

Tabla 1. Parámetros estimados del modelo ajustado en unidades de tiempo standard (meses).

Phase ej Phase (qm Maximum

Species Place Year E (Months after (Months after activity period R? Das ps

January) minimum dens!y) MA (Monthts) dpi

Helicella pappi Philippi 1987 12.6 2.9 5.9 7.1 OIOSIEZS

1988 10.5 3.4 6.8 6.9 0.95

Xerolenta obvia Karvali 1990 12.6 6.7 4.7 4.4 0.79 2

Paleokastro 1988 14.2 0.3 29 42 0.84 ]

1989 10.8 3.5 30) 7.3 0.77

1990 10.4 0.2 2.8 2.9 0.91

Xeropicta arenosa Potidea 1979 12.7 6.1 )) 5.1 0.87 |

1980 11.2 6.2 38) 6.9 0.88

Edessa 1984 12,5 6.6 4.9 6.7 0.69 ]

1985 11223 7.8 5.8 6.9 0.68

Monacha cartusiana Edessa 1984 6.5 2.4 37 3.2 0.53 l

1985 13.0 -1.1 6.2 4.3 0.45 2

Helix lucorum 1983 12.5 4.] 5.0 8.1 0.94 34

1984 12.0 8.4 7.9 8.5 0.84

1985 12.0 8.9 8.5 8.2 0.86

Table IL. Skewness (negatively skewed <0.5; positively skewed >0.5) and curtosis (leptocurtic <0.5;

platycurtic >0.5) from the phenograms of Helicoidea snails from Northern Greece). Abbreviations:

Phil: Philippi; Pal: Paleokastro area; Kar: Karvali; Pot: Potidea; E: Edessa (the paranthesis means slightly).

Tabla 11. Desviación (desviado negativamente <0, 5; desviado positivamente >0,5) y curtosis (leptocúrtico

<0,5; platicúrico >0,5) en los fenogramas de caracoles helícidos del Norte de Grecia. Abreviaturas: Phil:

Philippi; Pal: Paleokastro area; Kar: Karvali; Pot: Potidea; E: Edessa (los paréntesis significan ligeramente).

Species Place and Year deta o Symmetric Leptocurtic Platycurtic Mesocurtic ada

Helicella pappi Phil. 1987 (+) + 23

1988 + >

Xerolenta obvia Paleo. 1988 + . 1

1989 (+) +

1990 + +

Karv. 1990 + + (+) 2

Xeropicta arenosa Potid. 1979 + (+) ]

1980 + +

Edes. 1984 + + l

1985 + +

Monacha cartusiana Edes. 1983 + (+) 2

1984 + + 1

1985 + (+) + 2

Helix lucorum Edes. 1983 + + 3-4

1984 + +

1985 +

autumnal period and small ones breed of the reproductive period is very short

mainly in autumn (x= 4.261, P= 0.039) whereas in the inland areas it is variable

(Table III). In coastal areas the duration according to the species.

LAZARIDOU-DIMITRIADOU AND SGARDELIS: Phenology of Helicoidea in N Greece

ps mM

Right

O.4

Xik] xapl

80 Platy-

hi2

hpf2

a ni3

60 a Left

Skewness

4ES

mc3

Kurtosis

20 Lepto-

Figure 3. Ordination of phenograms into skewness(S)- curtosis (C) plane (Skewness: negatively ske-

wed< 0.5; positively skewed> 0.5. Curtosis: leptocurtic< 0.5; platycurtic> 0.5) of Helicoidea snails

from Northern Greece. Numbers denote 1st, 2d or 3d generation. Abbreviations, xap: Xeropicta are-

nosa Potidea; xae: Xeropicta arenosa Edessa; hl: Helix lucorum; hpf: Helicella pappi Philippi; mc:

Monacha cartusiana; xip: Xerolenta obvia Paleokastro.

Figura 3. Ordenación de fenogramas respecto al plano desviación (S)- curtosis (C) (Desviación: negati-

va< 0,5; positiva> 0,5. Curtosis: leptocúrtico< 0,5; platicúrtico> 0,5) de babosas de la familia

Helicoidea del Norte de Grecia. Los número denotan las primera, segunda y tercera generaciones.

Abreviaturas, xap: Xeropicta arenosa Potidea; xae: Xeropicta arenosa Edessa; hl: Helix lucorum; hpf

Helicella pappi Philippi; mec: Monacha cartusiana; x1p: Xerolenta obvia Paleokastro.

DISCUSSION

In Northern Greece the climatic con-

ditions are not uniform (HATZIOANNOU

ET AL., 1989). The ombrothermic

diagram for Northern Greece (Fig. 1)

from 1980 to 1990 shows that the dry

season is from June to October, whilst

the wet season is divided by a cold

winter period. Breeding may take place

almost in all seasons except during

winter. In Northern Greece, snails

mainly breed from April to the end of

autumn. The strong seasonality of the

climate imposes a seasonal pattern of

breeding. Consequently, there are two

main breeding periods: the autumnal

breeding period starts with the first

rainfalls and stops with low temperatu-

res (LAZARIDOU-DIMITRIADOU, 1981;

STAIKOU AND LAZARIDOU-DIMITRIADOU,

1991) and the vernal-estival breeding

period which starts when the mean

monthly temperature rises over 10%C

and stops when the arid period starts

(LAZARIDOU-DIMITRIADOU, 1981; LAZA-

RIDOU-DIMITRIADOU AND KATTOULAS,

1985; STAIKOU ET AL., 1988). Most of the

land snails though, mainly short-lived

and small snails, breed during the

autumnal period (Table III) as Helicoi-

dea species from Southern Europe do

(CHATFIELD, 1968; REAL AND REAL-

TESsTUD, 1983; HELLER, 1982; SACCHI,

Iberus, 15 (2), 1997

Table II. Life cycle characteristics from Helicoidea snails from Northern Greece. Abbreviations:

D: largest shell diameter. Ehinos is found in Rhodope area, Edessa and Thessaloniki in North

Central Macedonia, Philippi and Karvali near Kavala, and Paleokastro and Potidea in Chalkidiki.

Abbreviations, Y. years up to maturity; SL: short lived < 3 years; LL: long lived > 3 years; SS: small

sized D < 25 mm; LS: large sized D > 25 mm; V: vernal-estival reproductive period; A: automnal

reproductive period.

Tabla III. Características del ciclo de vida de los caracoles helícidos del Norte de Grecia. Abreviaturas: D:

mayor diámetro de la concha. Ehinos se encuentra en el área de Rhodope, Edessa y Thessaloniki al Norte

de Macedonia, Phillipi y Karvali cerca de Kavala, y Paleokastro y Potidea en Chalkidiki. Abreviaturas,

Y: años hasta la madurez; SL: vida corta < 3 años; LL: vida larga > 3 años; SS: pequeña talla D < 25

mm; LS: gran talla D > 25 mm, V: periodo reproductivo estival; A: periodo reproductivo otoñal.

Species Locality Longitude

Helix pomatia rhodopensis Ehinos 249 58' 34"

Helix lucorum Edessa DINA

Monacha cartusiana

(rarely )

Bradybaena fruticum Edessa

Cepaea vindobonensis Edessa

Helix figulina Thessaloniki 22% 57' 29”

Theba pisana Thessaloniki

Xerotricha conspurcata Thessaloniki

Eobania vermiculata Thessaloniki

Helicella (Xerothracia) papi Philippi 249 15' 48”

Xerolenta obvia Paleokastro MUDA

Karvali 2423011

Xeropicta arenosa Potidea DNA

Edessa IAS

Cernuella virgata Potidea ESSE

Latitude Y SL LL SS LS A

419 16' 50” 3 + +

409 47 47" 3-4 + +

2 + + +

1 + +

2 + +

419 24' 26” 2 + + +

2 + + +

| + + +

2 + + +

419 7'26' 2-3 + + +

409 24' 59" | + + +

409 59 44” 2 + + +

4091124” 1 + + +

407 47' 47” 1 + + +

401124" | + + +

1971; BONAVITA AND BONAVITA, 1962;

DEBLOCK AND HOESTLANDT, 1967) and

only some breed during the vernal

period as land snails from Northern

Europe do (POLARD, 1975; WOLDA AND

KREULEN, 1973). M. cartusiana, which is

of Northern origin, breeds in both

seasons (STAIKOU AND LAZARIDOU-DIMI-

TRIADOU, 1990). Estival breeding period

happens in places with a wet climate

during summer months (STAIKOU ET AL,,

1988). In areas where different species

coexist (as in Logos area in Edessa) alt-

hough the climatic conditions are the

same the species do not breed during

the same period provoking less antago-

nistic intraspecific reactions to their hat-

chings (STAIKOU ET AL., 1988). Semelpa-

rous species with an r-strategy synchro-

nize their breeding period with the

favourable period which is October-mid

November (LAZARIDOU-DIMITRIADOU,

1981, 1995; LAZARIDOU-DIMITRIADOU

AND KATTOULAS, 1985).

There is also a marked difference

between the snail species living along

the sea shore and the inland ones. Cou-

pling and laying of eggs is more or less

synchronous for the populations living

along the seashore and do not last more

than a week each. On the contrary bree-

ding lasts about a month for the inland

species (LAZARIDOU-DIMITRIADOU, 1981;

STAIKOU AND LAZARIDOU-DIMITRIADOU,

1991).

The climatic conditions under which

the land molluscs live do not only affect

the time of the breeding season but also

their whole life cycle and phenologies.

X. obvia needs two years to mature in

coastal or semi-coastal areas and one

year on the mountains (e. g. Paleokastro,

LAZARIDOU-DIMITRIADOU AND SGARDELIS: Phenology of Helicoidea in N Greece

Central Chalkidiki, unpublished data).

Similarly, M. cartusiana which is of nort-

hern origin needs two years to mature

in the south instead of one, because it

has to face the summer aestivation, alt-

hough 15% of its population in Edessa

tends to be semelparous (STAIKOU AND

LAZARIDOU-DIMITRIADOU, 1990), as in

Northern Europe (CHATFIELD, 1968).

Species like B. fruticum and E. vermi-

culata which are iteroparous and univol-

tine species, living for a few years, exhi-

bit no seasonal trend of abundance but

fluctuate almost randomly during the

year. These show rather stable patterns

despite the oscillations of environmental

parameters. All the rest of the studied

Helicoidea species can encounter seaso-

nality by adjusting the timing of their vi-

tal activities. They respond to the ad-

verse period of the year, which is winter

time for Northern Greece, by displaying

asymmetric (positively or negatively

skewed phenograms) seasonal patterns

of abundance variation. These patterns

reflect a differential response of the spe-

cies to tolerance against stress which se-

ems not to be region-specific but species-

specific. The annual or biennial r-strate-

gists (X. arenosa, X. obvia) show a

positively skewed phenological pattern

due to the rapid development of juveni-

les soon after the adverse winter period.

These juveniles have been hatched in au-

tumn but stayed buried in the soil du-

ring winter. Their growth stops before

summer dryness during which the geni-

talia and the gonad maturation take

place and the snails are ready to lay eggs

before their death in autumn. On the

contrary, populations of pluriennial spe-

cies are characterized by low abundance

during the onset of the adverse period

and a negatively skewed or symmetric

phenciogical pattern. It seems that the

growth rate of juveniles is much slower

than that of annual species. The only

species population that showed both a

negatively and a positively skewed phe-

nological pattern was H. lucorum. Howe-

ver, a positively skewed pattern in 1983,

that is a rapid growth of population den-

sity was probable just after the adverse

period since a massive emergence from

hibernation of adult snails took place be-

cause of good weather conditions which

started earlier than usually (STAIKOU ET

AL., 1988: fig. 2). Additionally, this spe-

cies has a low net reproductive rate (Ro

= 0.9) and a low annual turnover ratio

(P/B = 1.24), the snails live up to 12 ye-

ars and they mature after the third year

of their lives (STAIKOU ET AL., 1988).

Negatively skewed and leptocurtic

phenograms could not be characteristic

of a snail species since it would mean

that this population would grow slowly

and steadily during the favourable pe-

riod of the year and would introduce ra-

pidly growing immature snails just be-

fore the adverse period. This would be

possible only if immature snails were

more resistant and tolerant to the stress.

Such a phenology has not also been re-

corded in acari or collembola (SrTamou

ET AL., 1993; SGARDELIS, SARKAR, ASIKI-

DIS, CANCELA DA FONCECA AND STAMOU,

1993). However, M. cartustana is a case

of negatively skewed and slightly lepto-

curtic phenology. In this population 15%

of juveniles mature in one year as it

happens in Northern Europe. So, in 1984

the dry season (summer time) was inte-

rrupted by a wet period (STAIKOU ET AL,,

1988: fig. 2) and this 15% of juveniles,

which had already matured, managed

to lay eggs before the majority of the

snails which were mature and ready to

copulate and lay eggs in autumn. This

population, though, comes from Nort-

hern Europe and it is found in the sout-

hern limits of its distribution.

To sum up, environmental variables

in Northern Greece are strongly seaso-

nal and thus Helicoidea snails exhibit

predictable oscillations in their activity

patterns, which can be interpreted by

the demographic response of the popu-

lations as it has been found with soil

microarthropods (STAMOU ET AL., 1993;

SGARDELIS ET AL., 1993). The semelpa-

rous and short-lived snail species popu-

lations show a more stable phenological

pattern than the biennial and plurien-

nial ones, who mature after the first year

of their life, and they are more plastic

trying to face the climatic differences

from one year to the other.

Iberus, 15 (2), 1997

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Iberus, 15 (2): 35-50, 1997

Fragmented knowledge on West-European and Iberian

Caudofoveata and Solenogastres

Conocimiento fragmentado de los Solenogastros y Caudofoveados

de Europa occidental y Península Ibérica

Luitfried von SALVINI-PLAWEN*

Recibido el 8-1-1996. Aceptado el 4-X-1996

ABSTRACT

A basic problem in our knowledge of the aplacophoran molluscs, viz. the Caudofoveata

and the Solenogastres, is the poor availability of faunistic samplings. This lacunarity even

concerns the European waters; in the present contribution, particular attention is paid to

the gap in the records along the French and Iberian shelf regions. This is underlined by

presenting an updated geographic distribution of eight caudofoveate and thirteen soleno-

gastre species. Benthos investigators are called upon to focus more intensively on sam-

pling the smaller marine fauna from mobile bottoms of the West-European shelf regions.

RESUMEN

Un problema esencial para el conocimiento de los Caudofoveados y Solenogastros

(moluscos aplacóforos) es la insignificante disponibilidad de material recogido en diferen-

tes muestreos faunísticos. Esta carencia todavía afecta al Atlántico europeo y particular-

mente concierne a la falta de muestras en la plataforma continental de Francia y de la

Península Ibérica. Esta situación se pone en evidencia con la recopilación actualizada de

la distribución geográfica de ocho especies de Caudofoveados y trece de Solenogastros.

Se hace una invitación especial a los investigadores del bentos para que intensifiquen su

atención por la pequeña fauna marina de sustratos blandos en la plataforma occidental

europea.

KEY WORDS: Caudofoveata, Solenogastres, Aplacophora, new records, distribution, Europe.

PALABRAS CLAVE: Caudofoveados, Solenogastros, Aplacophora, nuevas citas, distribución, Europa.

INTRODUCTION

This contribution is restricted to a

very simple, but momentous problem:

the dearth of faunistic information with

all its consequences in both classes of

aplacophorous molluscs, the Caudofo-

veata (formerly Chaetodermomorpha)

and the Solenogastres (formerly Neome-

niomorpha).

Members of the Caudofoveata and

the Solenogastres live predominantly in

marine offshore habitats below 50 meters

depth and are in general not really rare

members of benthic biotopes (see SAL-

VINI-PLAWEN 1990). The Caudofoveata

(average size 2-15 mm) are micro-omni-

vores burrowing within muddy sedi-

* Institut fúr Zoologie, Universitat Wien, Althanstraffe 14, A-1090 Wien IX, Austria.

Iberus, 15 (2), 1997

ments, whereas the Cnidaria-vorous So-

lenogastres (average size 2-20 mm) are

bound to clay, secondary hard bottoms or

enidarian colonies. Our scarce know-

ledge about species diversity, biology

and zoogeography of the representatives

of both groups is in part due to the intri-

cate, high-effort and expensive sampling

methods for benthic meiofauna (ships,

cable winches, benthic sledge-dredges).

Due to these technical and financial diffi-

culties, investigation of marine meio-

fauna is generally restricted to the “home

turf” of marine biological stations or fis-

hery institutes as typified by Ply-

mouth/UK or Naples/Italy. For more

than a century, this has resulted in an un-

balanced biogeographic and systematic

knowledge of small fauna, even in Euro-

pean waters, restricting, the informative

data predominantly to animals of the

North and Mediterranean Seas. Surpri-

singly, there are only poor records from

the shelf region off France and the Iberian

peninsula (delimited in Figure 1 by the

200 m isobath).

Therefore, a special invitation is ad-

dressed to all Spanish, Portuguese, and

French colleagues who perform benthic

offshore investigations to include in their

projects the sampling of meiofauna from

mobile bottoms. It is only with the help

of such cooperative collecting work that

examination, determination and research

on small benthic animals such as Caudo-

foveata and Soleno-gastres can be satis-

factorily carried out. This cooperation is

vital to adequately enlarge our know-

ledge about the organization, biology

and biogeography of these primitive mo-

lluscan groups that still bear calcareous

bodies instead of a shell. Furthermore,

this knowledge is essential to also under-

stand Mollusca in general.

Finally, it must be underlined once

more that the determination of members

of both classes requires detailed examina-

tions (see SALVIN-PLAWEN, 1975 versus

ODHNER, 1921, for Caudofoveata; serial

sections for most Solenogastres).

The nature of the problem becomes

more evident when one examines the con-

crete documentation. With respect to the

presently-known caudofoveate and sole-

nogastre fauna in Northeast-Atlantic (Eu-

ropean) waters, there is a distinct gradient

between the Scandinavian-British records

(cf. SALVINI-PLAWEN, 1975; SEAWARD, 1982,

1991) and the West-European reports. Ex-

cept for a recent collection from four sites

off the coast of Galicia (in connection with

the project “Fauna Ibérica”; in prepara-

tion), other knowledge of aplacophoran

representatives from French and Iberian

shelf regions is restricted to a few random

findings. On the other hand, investigations

and records of both Caudofoveata and So-

lenogastres are again available from the

Mediterranean Sea. For Europe asa whole,

this results in an almost “bipolar” pattern

of intraspecific distribution. Clearly, in

contrast to the regular sampling in Scan-

dinavian, British and Mediterranean seas

(cf. SALVINI-PLAWEN, 1972, 1975, 1977a, b,

1988; SEAWARD, 1991), no purposeful offs-

hore samplings have been conducted on

the Iberian and French shelf in the past to

obtain at least an overview of West-Euro-

pe's small benthic fauna including Cau-

dofoveata and Solenogastres (the French

BIOGAS and POLYGAS samplings lie be-

yond the European shelf region).

The overall knowledge on Caudofo-

veata and Solenogastres has significantly

increased during the last decades, but is

still very poor when compared with ot-

her Mollusca (for an organisational and

structural overview see SALVINI-PLAWEN,

1985b, and SCHELTEMA, TSCHERKASSKY

AND KUZIRIAN, 1994, respectively; their

phylogenetic status is analysed in SAL-

VINI-PLAWEN AND STEINER, 1996). The

poor, random information on their occu-

rrence in West-European waters also ne-

gatively affects our knowledge on the full

range of organisation (systematics, com-

parative anatomy) as well as on biologi-

cal conditions and circumstances.

The above-mentioned “bipolarity” in

intraspecific distribution, with the inter-

vening West-European gap, becomes ob-

vious when considering all aplacophoran

representatives known from both nort-

hern and southern waters; these are do-

cumented below. Other species with an

up to now purely Mediterranean or

North-European distribution have a po-

tential West-European occurrence (Lusi-

SALVIN-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres

: Of, Gibraltar

Figure 1. Section of the West-European Atlantic demonstrating the ofÉshore shelf region down to

the 200 m isobath (followed by the continental decline to 3000 m depth).

Figura 1. Sección del Atlántico europeo mostrando la zona de la plataforma continental, hasta la isóba-

ta de 200 m (seguida de la zona del talud continental hasta los 3000 m de profundidad).

tanic region and / or Bay of Biscay ); exam-

ples given below are the caudofoveates

Psilodens tenuis and Chaetoderma strigis-

CAUDOFOVEATA

The Caudofoveata burrow within

mobile bottoms and have adapted a ver-

miform body with reduced pedal sole

quamatum as well as the solenogastres Bi-

serramenia psammobionta and Anamenia

gorgonophila.

(midventral fusion of the lateral mantle

rims). Generally, sampling in muddy bio-

topes (sledge-dredges, grabs) success-

Iberus, 15 (2), 1997

fully yields specimens. This group (for-

merly Chatodermomorpha) was separa-

ted from the Solenogastres and elevated

to class rank due to the paraphyletic sta-

tus of its aplacophorous organisation (see

SALVINI-PLAWEN AND STEINER, 1996). It

includes 98 named species classified into

three families (Limifossoridae, Prochae-

todermatidae, Chaetodermatidae). Se-

venteen European representatives have

been described so far, six of which occur

in the Mediterranean including three en-

demic species (cf. SALVINI-PLAWEN, 1990).

All species of the West-European

shelf region along with the Iberian waters

of the Mediterranean will be documen-

ted. Thus, among the Prochaetodermati-

dae, the deep-sea species Prochaetoderma

yongei Scheltema, P. clenchi (Scheltema),

and P. (Chevroderma) turnerae (Scheltema)

from the BIOGAS-cruises are not consi-

dered. These three species also inhabit

the basin of the Bay of Biscay (2* 10' - 92

W> at depths of 1175-2006 m (P. yongei),

1913-2430 m (P. clenchi) and 2124-4760 m

(P. turnerae) (see SCHELTEMA, 1985; for ta-

xonomy cf. SALVINI-PLAWEN, 1992).

Besides Falcidens aequabilis, other spe-

cies of Falcidens (Chaetodermatidae) are

likewise of biogeographical interest: at le-

ast among the known species which are

provided with a slender, tail-like poste-

rior body, each appears to inhabit a well-

defined, non-overlapping geographic re-

gion. Thus, E gutturosus (Kowalevsky,

1901) is Mediterranean (endemic), while

F. crossotus Salvini-Plawen has a Scandi-

navian-British distribution. A third “tai-

led” species, Falcidens vasconiensis Sal-

vini-Plawen, comes from the Gulf of Gas-

cogne (SALVINI-PLAWEN, 1996), and

future (not yet recorded) Lusitanic repre-

sentatives may well belong to yet another

species.

Family LIMIFOSSORIDAE

Scutopus ventrolineatus Salvini-Plawen, 1968

Known distribution (Figure 2A): Scan-

dinavian coast (Skagerrak to Tromso0),

North Sea, West-Scotland, Irish Sea,

southern Bay of Biscaya, Alborán Sea

(off Vélez-Málaga), Gulf of Lion (off

Banyuls, off Marseille), SE Africa (off

Durban); 40-1248 m.

Remarks: The occurrence of this very

slender and often coiled species has

been summarised in SALVINI-PLAWEN

(1975, 1977b). Supplementary records

come from the North Sea (Hartley, 1984)

and from off Barcelona / Catalonia with

the cruises RETRO 1 (41? 08' 07” N, 022

04” 32” E, 510 m) and ESPERMA 89 (41?

0437" N, 019 5933" E, 600'm) carried

out by Luis Dantart; a recent finding

comes from off Vélez-Málaga (4* 03” W)

at 400 m. This species is of special inte-

rest insofar as it has also been recorded

from off Southeast Africa, which indica-

tes a distribution along all East-Atlantic.

Scutopus robustus Salvini-Plawen, 1970

Known distribution (Fig. 2B): Off the

Norwegian coast with larger gaps from

Oslofjord to North of Trondheimsfjord,

scattered in the Western Mediterra-

nean Sea to 9? East; 50-3542 m.

Remarks: There are no additional

records referring to this slender, up to

10 mm species beyond the occurrence

summarised in SALVINI-PLAWEN (1975,

1977a).

Psilodens tenuis Salvini-Plawen, 1977

Known distribution (Fig. 2C): Lusita-

nic Atlantic S of Cap Sao Vicente; 2500

Remarks: There is a single record

only, as communicated in SALVINI-

PLAWEN (1977a).

SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres

Family PROCHAETODERMATIDAE

Prochaetoderma raduliferum (Kowalevsky, 1901)

Chaetoderma radulifera Kowalevsky, 1901, Archs. Zool. exp. gén., sér. 3, 9: 264.

Known distribution: Endemic in the

Mediterranean Sea, known from the Sea

of Marmara in the East to off the Alge-

rian coast in the West; 30-2415 m.

Remarks: To date, this species is

known only from the Mediterra-nean

Sea (see map Abb. 5 in SALVINI-PLAWEN,

1977b). Unlike most other members of

the Prochaetodermatidae, P. raduliferum

is not a true deep-sea species. As is

demonstrated by Adriatic and lonic

samplings (30-215 m) as well as by

records from off Banyuls (60-275 m)

summarised in SALVINI-PLAWEN (1977b),

it is quite regularly found on muddy

offshore bottoms. In accordance with

this, there are new records from off the

West coast of Malta at 120-160 m

(MrirsuD, 1996; the specimen photograp-

hed by Mirsub Fig. 2, however, is a

broken Falcidens gutturosus, see below),

from off Barcelona /Catalonia by Luis

Dantart (four stations at 41? 04' 37”-41?

OIM AN AOS ASS OLA SE SOS

680 m), and most recently from off

Vélez-Málaga to off Málaga (80-300 m).

Family CHAETODERMATIDAE

Falcidens gutturosus (Kowalevsky, 1901)

Chaetoderma gutturosum Kowalevsky, 1901, Archs. Zool. exp. gén., sér. 3, 9: 281.

Known distribution: Endemic in the

Mediterranean Sea, known from off

Palestine and from the Sea of Marmara

in the East to off Málaga in the West; 40-

866 m.

Remarks: Falcidens gutturosus is a

fairly common species characterised

by a slender, tail-like posterior body

with an orange-red terminal tassle.

Beyond the already known, purely

Mediterranean distribution (see map

Abb. 2 in SALVINI-PLAWEN, 1977b),

there are new samplings from off the

West coast of Malta at 120-160 m

(Mirsub, 1994, 1996), by L. Dantart

from off Barcelona (see SALVINI-

PLAWEN, 1996) and by A. Zenetos from

the Gulf of Korinth as well as from the

Gulf of Petalión (56 m; Greece); most

recently, specimens were recorded

from off Vélez Málaga (40 m) and off

Málaga (211 m).

Falcidens vasconiensis Salvini-Plawen, 1996

Known distribution: Gulf of Gas-

cogne; 141-170 m.

Remaks: Up to present there is a single

record only from off the Cap Breton in

the southeastern Bay or Biscaya (SALVINI-

PLAWEN, 1996). Its distribution throug-

hout the shelf region of the Gulf of Gas-

cogne is to be expected.

Falcidens aequabilis Salvini-Plawen, 1972

Known distribution: Endemic in the

Mediterranean Sea, ranging from the

Aegean Sea to the western Mediterra-

nean deep-sea bottom as far as the

Greenwich meridian; 132-3542 m.

Remarks: This species appears to in-

habit deeper and/or far offshore bot-

toms. Because of the technical effort in-

volved, it is consequently less fre-

quently recorded than F. gutturosus or

Iberus, 15 (2), 1997

Prochaetoderma raduliferum, but is well-

documented from the West-Mediterra-

nean deep-sea (Campagne Polymede, cf.

SALVINIEPLAWEN, 1977a and 1977b: map

Abb. 2). There is a new record by Luis

Dantart from off Barcelona / Catalonia

(RETROTATAOI406AN 02 03 OLE

350-426 m).

Chaetoderma (?) strigisquamatum Salvini-Plawen, 1977

Known distribution (Fig. 2C): Basin of

Alborán (W-Mediterranean Sea); 1491

Remarks: There is a single record of this

chaetodermatid species, whose generic

classification needs confirmation (radula

SOLENOGASTRES

The Solenogastres (= “those with a

belly furrow”) include narrowed apla-

cophoran molluscs that still bear a pedal

groove to glide upon (formerly Neome-

niomorpha). A total of some 190 species

has been described which, in accordance

with integumentary characters, is grou-

ped into four orders (Pholidoskepia, Ne-

omeniamorpha, Sterrofustia, Cavi-

belonia). To date, 46 species are known

in European waters, of which 26 (inclu-

ding 19 “endemics”) are represented in

the Mediterranean Sea. However, little

is known about their biogeography:

most species have been recorded only

once, and inaccurate descriptions (see e.

g. Wirenia, below) additionally contri-

bute to difficulties in classification, re-

sulting in insufficient faunistic informa-

tion. Thus, the poorly described Nemato-

menia (?) corallophila (Kowalevsky, 1881),

recorded from off La Calle / Algeria at

73-183 m (37? N, 8* 30” E) as living epi-

zoically on Corallium rubrum (Linné),

apparatus; cf. SALVINI-PLAWEN, 1977a). Ba-

sed on the known distribution of Caudo-

foveata in the Mediterranean Sea in ge-

neral (SALVINI-PLAWEN, 1977b), a Lusita-

nic occurrence of this species might be

expected.

could only be recognised in the future

and re-described if it is rediscovered

again on a red coral (its alledged finding

in the Bay of Rosas/Costa Brava is a

mistranslation by Mars, 1965, from Ma-

LUQUER, 1917).

Most species documented here, toget-

her with a few others, belong to the small

number of representatives found several

times. Records of these findings are very

much tied to sampling methods and

habitat. For example, the well-known Neo-

menta carinata was never recorded by

means of sledge-dredges (muddy

bottoms), as predominantly used by the

author and his group. In another example,

those Solenogastres living on thecapho-

ran Hydrozoa or Octocorallia (e. g. Nema-

tomenia and Anamenia, below), are more

often sampled from secondary hard

bottoms (e. g. with Agassiz-trawls) or by

workers studying cnidarians. Thus, all

these circumstances help explain our frag-

mented biogeographic knowledge.

Order PHOLIDOSKEPIA

Family DONDERSIDAE

Nematomenta flavens (Pruvot, 1890)

Dondersia flavens Pruvot, 1890, Archs. Zool. ex. gén., sér. 2, 8: XXIL

Known distribution (Fig. 2D): Off

Banyuls - Costa Brava, Shetland Islands;

45-167 m.

Remarks: This slender, up to 4 cm

long species has a showy lemon-

yellow colour. It is not rare along the

SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres

rocky (French and Spanish) Catalan

coast, feeding upon Hydrozoa-The-

caphora at 45-90 m (Pruvor, 1891;

MALUQUER, 1917; Mars, 1965). Ano-

ther record refers to the Shetland

Islands at 167 m, epizoic upon Lafoea

dumosa SARS; the anatomical examina-

tion revealed the presence of a vesti-

gial radula and the necessity for a

family reclassification (SALVINI-PLA-

WEN, 1978: 39-40).

Recently, the Irish Sea Survey

(MACKIE, OLIVER AND REES, 1995: 192)

collected eleven samples of N. banyu-

lensis (see below); however, there is no

information about the exact methos of

determination applied. The sampled

material had been fixed in formalin

and then preserved in alcohol (MACKIE

ET AL., 1995: 15-16); the specific body-

colour of both N. banyulensis (red) and

N. flavens (yellow) is no longer visible

after such treatment. Therefore, it

might well be that some of the N. ban-

yulensis-records in reality belong to the

externally very similar N. flavens. Only

an accurate histological determination

(serial sections) can provide the exact

specific classification of these speci-

mens.

Nematomenta banyulensis (Pruvot, 1890)

Dondersia banyulensis Pruvot, 1890, Archs. Zool. ex. gén., sér. 2, 8: XXIL

Nematomenia banyulensis var. norvegica Odhner, 1921, Bergens Mus. Aarb. 1918-19, Naturvid.

reekke 3: 43.

Muyzomenia Simroth, 1893, Zeitschr. wiss. Zool., 56: 324.

Known distribution (Fig. 2E): Off Dal-

matia, Gulfs of Naples and Salerno,

Cóte Vermeille, off Roscoff, Plymouth

Sound to Irish Sea to W-Scotland, off

Northumberland, Trondheimsfjord-Fill

(amfjord; 31-300 m.

Remarks: This well-known species

likewise lives epizoically upon Hydro-

zoa-Thecaphora. Its slender body

reaches a length of up to 3 cm and is red

(as are also two other Mediterranean

species). Its distribution is summarised

in NIERSTRASZ AND STORK (1940) and

more recently in SEAWARD (1982, 1991)

for the British waters. It has also

recently been found several times by the

Irish Sea Survey (MACKIE ET AL., 1995:

192), but compare the above remarks

with N. flavens. Geographically new

records include samples from off Sebe-

nico /Sibenik (Adriatic Sea) at 57 m, 61

m and 67-68 m (see also SALVINI-

PLAWEN, 1986) and from the Fill

(an)fjord = north-eastern Hitra Island off

Trondheimsfjord (Mus. Uppsala).

A comparative examination of Medi-

terranean and Norwegian (syntype)

individuals, particularly with respect to

the mantle scales, revealed no differen-

ces which would vindicate a separation

of the Norwegian specimens (as varia-

tion or subspecies proper).

Stylomenia salvatori Pruvot, 1899

Known distribution: Off Banyuls,

(?)Costa Brava; about 60-80 m.

Remarks: This species had been

found together with Rhopalomenia agla-

opheniae (q. v.) in an aquarium filled

with benthic material from off Banyuls-

sur-Mer; based on the presence of Rh.

aglaopheniae, this indicates an original

depth of about 60-80 m (see PrUVOT,

1891: 721). MALUQUER (1916: 244, 1917:

37-38) reports finding animals similar

to S. salvatori from the Bay of Rosas and

off Llansa (Costa Brava). Even if the

occurrence of this species is to be expec-

ted there, the record needs to be confir-

med because no accurate determination

(histological examination) was perfor-

med.

Iberus, 15 (2), 1997

Family LEPIDOMENIIDAE

Tegulaherpia myodoryata Salvini-Plawen, 1988

“Species D” in Salvini-Plawen, 1968a, Sarsia, 31: 132.

Tegulaherpia celtica Caudwell, Jones and Killeen, 1995, Journ. Conch. (London), 35: 258.

Known distribution (Fig. 2F): Off Li-

vorno, off Banyuls-sur-Mer, southern Bay

of Biscay (North of Asturias, THALASSA-

Stat. W-415)?, Celtic Sea, area around Ber-

gen (Raunefjord, Hjeltefjord), area around

Trondheim (Fill (an)fjord, Trondheimsf-

jord); 75-470 (75-860 / 1150?) m.

Remarks: This Mediterranean species

is likewise native to Northern Europe. In

the course of examining more compre-

hensive Solenogastres material from the

North Atlantic, the already communica-

ted “Species D” (SALVINI-PLAWEN, 1968a)

and T. celtica (CAUDWELL, JONES AND KI-

LLEEN, 1995), according to histological exa-

mination by series sections, turned out to

be conspecific with T. myodoryata from the

Western Mediterranean Sea as described

in SALVIN-PLAWEN (1988). Moreover, se-

veral other Norwegian individuals (Hjel-

tefjord, 200 m, Fill (an)-fjord, Trond-

heimfjord, 470 m) likewise belong to this

species.

A single specimen from the THA-

LASSA-Cruise (Stat. W-415, 439 55' 06” N,

06” 11 18” W; 860-1150 m), forwarded in

1971 by F. Monniot (Paris) to the author,

possibly also represents T. myodoryata,

since the mantle scales fully fit into the

range of shape, outline and size of the myo-

doryata-scales. However, the animal was

useless for histological examination, and

the record from a depth between 860 and

1150 m lends doubt to a conspecificity as

long as no bathymetrically interbridging

and /or additional samples are taken.

Family WIRENIIDAE

Wirenia argentea Odhner, 1921

Aesthoherpia glandulosa Salvini-Plawen, 1985, The Mollusca (Academic Press), 10: 94.

“Species B” in Salvini-Plawen, 1968a, Sarsia, 31: 131.

Aesthoherpia glandulosa Salvini-Plawen and “Species D” Haszprunar, 1986, Zool. Anz., 217: 345-360.

Known distribution (Fig. SA): Area

around Bergen/Norway, Hardanger-

fjord, area of Trondheimsfjord, Adriatic

Sea, Aegean Sea; 95-700 m.

Remarks: A most recent examination of

the hitherto missing type material of

Wirenia argentea Odhner (now in the

Naturhistoriska Rijkmuseet, Stockholm)

revealed that Aesthoherpia glandulosa

Salvini-Plawen is conspecific with it.

Despite some inaccurate and insufficient

presentations by ODHNER (1921: 31-34;

foregut, no radula, and so on), which led

to the description of Aesthoherpia (see

PLAWEN, 1988: 383-384), Wirenia has

nomenclatorial priority. The organisation

of the species and its presently known geo-

graphic distribution are communicated

(as Aesthoherpia glandulosa) in SALVINI-

PLAWEN (1988). Some additional findings

come from recently examined Norwegian

samples: area Northwest of Bergen (Hjel-

tefjord, 280 m; Herdlafjord, 200 m; Man-

gerfjord, 350 m) and Fill (an)fjord-Trond-

heimsfjord (95 m, 185 m, 320 m, 470 m and

490-500 m). The record of “Wirenia argen-

tea” by HARTLEY (1984) from the North Sea

needs specific confirmation.

Family MACELLOMENIDAE

Macellomenta palifera (Pruvot, 1890)

Paramenia palifera Pruvot, 1890, Archs. Zool. exp. gén., sér. 2, 8: XXIIL

SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres

¡IS 7 Ñ N DN

Figure 2. A-C. Caudofoveata. Known European distribution. A: Scutopus ventrolineatus Salvini-

Plawen, 1968; B: Scutopus robustus Salvini-Plawen, 1970; C: Known records of Psilodens tenuis

Salvini-Plawen, 1970 (black circle) and of Chaetoderma(?) strigisquamatum Salvini-Plawen, 1971

(asterisk). D-F: Solenogastres. Known distribution. D: Nematomenia flavens (Pruvot, 1890); E:

Nematomenia banyulensis (Pruvot, 1890); E: Tegulaherpia myodoryata Salvini-Plawen, 1988.

Figura 2. A-C. Caudofoveados. Distribuciones conocidas. A: Scutopus ventrolineatus Salvini-Plawen,

1968; B: Scutopus robustus Salvini-Plawen, 1970; C: Citas conocidas de Psilodens tenuis Salvini-

Plawen, 1970 (círculo negro) y de Chaetoderma(?) strigisquamatum Salvini-Plawen, 1971 (asterisco).

D-F: Solenogastros. Distribuciones conocidas. D: Nematomenia flavens (Pruvot, 1890); E:

Nematomenia banyulensis (Pruvot, 1890); F: Tegulaherpia myodoryata Salvini-Plawen, 1988.

Iberus, 15 (2), 1997

Known distribution (Fig. 3B): Cóte

Vermeille, Irish Sea (?); 80-120 m.

Remarks: This species, with its parti-

cular calcareous mantle-bodies, was ori-

ginally recorded with a single specimen

North of Port Vendres (Cóte Vermeille;

southeastern France) on muddy bottom

at 80 m. Two individuals recently

sampled from the Irish Sea at 80 m and

120 m come very close to M. palifera

(CAUDWELL ET AL., 1995). In view of the

“bipolar” occurrence of other species

(demonstrated herein), there is a high

probability that the species are identi-

cal. However, as the British animals

have not been investigated anatomically

(series sections), true conspecificity

ramains uncertain.

Order NEOMENIAMORPHA

Family NEOMENIIDAE

Neomenta carinata Tullberg, 1875

Solenopus nitidulus Koren and Danielssen, 1877, Arch. Math. Naturvid. (Kristiania), 2: 124.

Solenopus affinis Koren and Danielssen, 1877, Arch. Math. Naturvid. (Kristiania), 2: 127.

Neomenia grandis Thiele, 1894, Zeitschr. wiss. Zool., 58: 223.

Known distribution (Fig. 3C): Nor-

thern Kattegat and Bohuslán (W-Swe-

den), Norwegian coast between Oslo-

fijord and Sognesjóen/Sogne-fjord,

Romsdalsfjord, Trondheimsfjord, South

of Lofoten, Iceland, Shetland Islands,

British Isles, off Roscoff, Costa Brava,

Gulf of Genova, Gulf of Naples, off Mes-

sina; 10-565 m.

Remarks: This up to 3 cm long species

has a stoutish shape and is well docu-

mented along the coast of Scandinavia

and around the British Isles (KOREN AND

DANIELSSEN, 1879; WIRÉN, 1892; ODHNER,

1921; Muus, 1959; SEAWARD, 1982, 1991)

including Strindfjord / Trondheimsfjord

(Mus. Copenhagen) and the Hebrides

(Mus. Leiden). The Mediterranean

records include N. affinis (Koren and

Danielssen) which, according, to certain,

minor anatomical differences (pers. obs.),

can be classified as a subspecies only

(SALVINI-PLAWEN 1986); the same holds

true for N. grandis Thiele (NIERSTRASZ

AND STORK, 1940). A remarkable record

refers to Iceland (KNUDSEN, 1949), a

region in which Neomenia dalyelli (Koren

and Danielssen) is generally found.

Order CAVIBELONIA

Family PARARRHOPALIDAE

Eleutheromenia sierra (Pruvot, 1890)

Paramenta sierra Pruvot, 1890, Archs. Zool. exp. gén., 8: XXUL

Known distribution (Fig. 3D): Costa

Brava, Bretagne; Irish Sea; Trondheim

region; 40-128 m.

Remarks: PRUVOT (1891) typifies the

species from a single specimen (Cap

Creus / Costa Brava; 80 m) and in 1897 he

reports another finding from off Roscoff

at about 40 m (Pruvor, 1897). A single spe-

cimen of typical appearence (lobed dor-

somedian keel) from Stjórn (North or

Trondheim / Norway), despite the geo-

graphical distance, after serial section reve-

aled to be Eleutheromenia sierra. Conse-

quently, the questioned presence or this

species in the southwestern Cartigan Bay

(Irish Sea, 52 m; see HARTLEY 1984,

SEAWARD, 1991: 14) as well as the speci-

mens from nine samples or the Irish Sea

Survey referred to E. sierra (CAUDWELL ET

AL., 1995: 266; MACKIE ET AL., 1995: 192;

not documented in Fig. 3D) are indirectly

confirmed to really belong to this species.

On the other hand, the Pararrhopalii-

dae represent a fairly diverse group of sys-

SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres

Figure 3. Solenogastres. A: Known distribution of Wirenía argentea Odhner, 1921; B: Records of Ma-

cellomenia palifera (Pruvot, 1890) (black circles) and of Meromenia hirondellei Leloup, 1949 (asterisk);

C: Known distribution of Neomenia carinata Tullberg, 1875; D: Records of Eleutheromenia sierra

(Pruvot, 1890), see text; E: Known distribution of Biserramenia psammobionta Salvini-Plawen, 1968;

E: Known European distribution of Rhopalomenia aglaopheniae (Kowalevsky and Marion, 1887).

Figura 3. Solenogastros. A: Distribución conocida de Wirenia argentea Odhner, 1921; B: Citas de Macel-

lomenia palifera (Pruvot, 1890) (círculos negros) y de Meromenia hirondellei Leloup, 1949 (asterisco);

C: Distribución conocida de Neomenia carinata Tullberg, 1875; D: Citas de Eleutheromenia sierra (Pru-

vot, 1890), véase texto; E: Distribución conocida de Biserramenia psammobionta Salvini-Plawen, 1968;

F: Distribución europea de Rhopalomenia aglaopheniae (Kowalevsky y Marion, 1887).

Iberus, 15 (2), 1997

tematically very difficult representatives

(see SALVINI-PLAWEN, 1978); several geo-

graphically close records of Pararrhopa-

liidae may represent different species (or

even genera). Thus, another record from

the Irish Sea (Pruvotina sp. in CAUDWELL

ET AL., 1995: 265-267) clearly does not

belong to E. sierra; the same can be said

about two THALASSA-specimens from

off Galicia and off Asturias (Bay of Biscay)

Family SIMROTHIELLIDAE

Biserramenta psammobionta Salvini-Plawen, 1968

Known distribution (Fig. 3E): Irish Sea,

Plymouth area, Bretagne, Galicia; 8-30 m.

Remarks: In addition to the type mate-

rial from off Roscoff at 8-10 m (SALVINI-

PLAWEN, 1968b; see also MONNIOT, 1965),

several individuals have recently been

recorded in Plymouth Sound at 9-11 m

(50% 20” 43” N, 4? 09 05” W; see also

KIKINGER AND SALVINI-PLAWEN, 1995).

Moreover, a single specimen has been

sampled by lan Killeen during the Irish

Sea Survey from the Cardigan

Bay / Wales (Stat. 46, 52* 19% 12” N, 04? 37"

W) at 30 m, provided by Cathy Caudwell

to the author (see also CAUDWELL ET AL.,

1995). Finally, the 12 Solenogastres refe-

rred to as “Lepidomenia sp.” by Celia Bes-

teiro in her Ph. D. thesis (1986) from

Galicia / Spain (Ría de Ferrol, “Bajo de la

Palma”; 43? 27' 59” N, 08* 16' 23” W; 14

m) also represent Biserramenia psammo-

bionta. They all come from coarse sand or

shell gravel bottoms and at least those

from Roscoff, Plymouth and Galicia are

interstitially living animals (cf. SALVINI-

PLAWEN, 1985a; also OTT AND BOcH-

DANSKY, 1991, for the Plymouth animals).

The histological examination of these

specimens revealed some details beyond

the original description (SALVINI-

PLAWEN, 1968b). First, the characteristic

circular musculature around the spaw-

ning ducts and the posterior mantle

cavity is not yet elaborated in juveniles.

The slender pericardioducts with a tiny

lumen still open from dorsal into the

spawning ducts close to their rostral

ends, these ducts being paired throug-

hout with a wide lumen. Further diffe-

rentiation thus includes a curving elon-

gation of the rostral portion of the spaw-

ning ducts, which results in the adult

spawning ducts bending dorso-poste-

riorly (as described in 1968). Here, the

pericardioducts join their ends not

axially but ventrally, thus causing a

bulgy enlargement or even a slight bend

in the continuous lumen. This bulged

enlargement is the site of sperm storage,

thus functioning as receptacula seminis;

well-defined, set-off seminal pouches

(vesiculae seminales, as described

earlier), however, are not present. The

paired lateral pouch of the ventrorostral

mantle cavity is well differentiated only

in fully-grown individuals and often

merely represents two simply lobulated,

more or less distinct sacculations

opening medially through a short duct

or pore into the pallial space. Rather

than being paired in the sense of two sin-

gular, separate ganglia, the cerebral

ganglia are fused together in the middle

third of their extension. In some speci-

mens a small dorsoterminal sense organ

could be detected at the rear of the body.

Family AMPHIMENIIDAE

Meromenia hirondellei Leloup, 1949

Known distribution (Fig. 3B): Nort-

hern Bay of Biscay; 166 m.

Remarks: A single fragment of this

species had been recorded from the con-

tinental platform in the northern Bay of

Biscay (46” 27” N, 4* 09% 45” W) at 166 m

depth. Due to the unknown organisa-

tion of the anterior body, the generic

classification is uncertain (LELOUP, 1950:

21-23; SALVINI-PLAWEN, 1972: 224-225).

SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres

o y

Figure 4. Solenogastres. A: Known European distribution of Anamenia gorgonophila (Kowalevsky,

1880); an additional record refers to the Azores; B: Known distribution of Dorymenia sarsii (Koren

and Danielssen, 1877).

Figura 4. Solenogastros. A: Distribución europea de Anamenia gorgonophila (Kowalevsky, 1880); una

cita adiccional se refiere a las islas Azores; B: Distribución conocida de Dorymenia sarsii (Koren y Daniels-

sen, 1877).

Family RHOPALOMENIDAE

Rhopalomenia aglaopheniae (Kowalevsky and Marion, 1887)

Rhopalomenia eisigi Thiele, 1894, Zeitschr. wiss. Zool., 58: 269.

Known distribution (Fig. 3F): Off Cap

Matapan/Tainaron (South-Peleponnes),

Gulf of Naples, off Marseille, Cóte Ver-

meille, off Roscoff, British Isles; 50-137 m.

Remarks: This well-known species li-

ves upon Hydrozoa-Thecaphora, almost

exclusively upon Lytocarpia myriophyllum

(Linné). The distribution is compiled in

NIERSTRASZ AND STORK (1940), SALVINI-

PLAWEN (1972) and SEAWARD (1982, 1991).

The identification of several specimens

from off Monrovia / Liberia (THIELE, 1906:

324) needs re-examination and / or confir-

mation.

Family STROPHMENIDAE

Anamenia gorgonophila (Kowalevsky, 1880)

Proneomenia nierstraszi Stork [in Nierstrasz and Stork], 1940, Zoologica (Stuttgart), 99: 57.

Anamenia heathi Leloup, 1947, Bull. Mus. roy. Hist. Nat. Belgique, 23 (26): 1-11.

Known distribution (Fig. 4A): Gulfs of

Naples and Salerno, off La Calle (eas-

ternmost Algeria), off Marseille, Sea of

Alborán, Gorringe-Bank (WSW of Cap

Sao Vicente), Azores; 65-845 m.

Remarks: The records of this species have

beenrevised by SALVINI-PLAWEN (1972). The

investigation of numerous Solenogastres

recorded more recently from the SW of the

Isle of Alborán (see TEMPLADO, GARCÍA-

CARRASCOSA, BARATECH, CAPACCIONI,

JUAN, LÓPEZ-IBOR, SILVESTRE AND MASSÓ,

1986: 101-102) revealed that they in part

belong to A. gorgonophila, and the known

distribution of the species supports the

assumption of its presence in Lusitanian

waters as well. This species lives upon Gor-

gonaria, predominantly upon Paramuricaea

clavata (Risso) = P. chamaeleon (Koch), but

also upon Eunicella spp. and others.

Iberus, 15 (2), 1997

Family PRONEOMENIIDAE

Dorymenia sarsil (Koren and Danielssen, 1877)

Simrothiella sarsi Auctt. (see Opinion 1185 ICZN)

Known distribution (Fig. 4B): Trond-

heimfjord, Sognefjord, Bergen area, Os-

lofjord, Skagerrak, Gorringe Bank (off Cap

Sao Vicente); North Atlantic-Arctic Ocean

outside Troms0?; 183-681 m (1134 m?).

Remarks: The up to 7 cm long, slender

species was redescribed by ODHNER (1921)

and is externally characterised by a dis-

tinct (dorso-)terminal extension of the

body; a photograph is given in SALVINI-

PLAWEN (1968a: Abb. 23). Some recently

examined material from Scandinavian col-

lections extends the known distribution

(see Fig. 4B); the North Atlantic-Arctic spe-

cimens (71 25 N, 15” 41' E, 1134 m; see

ODHNER, 1921, and JAECKEL, 1954) only

doubtfully belong to D. sarsii based on the

geographic and bathymetric distribution.

In addition to the polystichous radula, a

pair of copulatory stylets and the presence

of a dorsoterminal sense organ typical for

the genus, histological investigations un-

derline two particular specific characters

in mature animals: the anterior portion of

the pericardioducts bears small pockets

serving as vesiculae seminales, and the

posterior portion of the spawning ducts

(“lower gametoducts”) in front of their fu-

sion each elaborate a ventral enlargement

or voluminous sacculation (pocket). Both

these characters allow this particular Dory-

menta species to be identified in SCHEL-

TEMA ET AL. (1994: Figs. 22 E and 24 G) as

D. sarsii (Koren and Danielssen). The ge-

ographic distribution of this species is thus

enlarged to the Gorringe Bank of the Ibe-

rian shelf region (36? 50' N, 9* 15 W, 681

m, Cf. SCHELTEMA ET AL., 1994: 18).

Family LEPIDOMENIIDAE

Lepidomenia ? spp.

Lepidomenia hystrix Auctt. non Marion and Kowalevsky, 1886

Lepidomenia (?) swedmarki Salvini-Plawen, 1985, Stygología, 1 (1): 103.

Remarks: Some records of small, mesop-

sammic Solenogastres from off Marseille,

Bretagne and off Belfast / Northern Ireland

have been systematised as Lepidomenia

hystrix Marion and Kowalevsky (see

SALVINI-PLAWEN, 1985a; SEAWARD, 1991).

Referring to the discussion in SALVINI-

ACKNOWLEDGEMENTS

The updating of the geographical

distribution in part comes from the

Scandinavian material in elaboration by

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O Sociedad Española de Malacología —_—_—_—_—_—_—_———— Iberus, 15 (2): 51-66, 1997

Molluscs as evolving constructions: necessary aspects for a

discussion of their phylogeny

Los moluscos como construcciones en evolución: aspectos necesarios

para una discusión sobre su filogenia

Karl EDLINGER* and Wolfgang Friedrich GUTMANN!

Recibido el 8-[X-1995. Aceptado el 19-X-1996

ABSTRACT

A model for the evolution of molluscs is presented. The reconstruction is based on the Frank-

furt Theory which conceives organisms as energy transforming hydraulic units which are

subject to evolutionary transformation according to the constructional principles ruling orga-

nisation. Evolution is reconstructed as a process of constructional transformations in which

all stages are explained as explicitly viable constructions; the irreversible transformation

phases are also rationally explained by referring to constructional properties of the orga-

nismic machines. lt is maintained that the predecessors of molluscs must have been elongate

worm like animals which were internally tethered by muscles in a way that creeping on the

flattened ventral “foot” surface became possible. Only organisms controlling the cross-section

by a densely spaced muscle system could start creeping on hard substrate. The establishment

of the radula is shown to have been dependent on the adhesive creeping movements which

allowed anchorage of the construction to the substrate during rasping. The formation of the

shell elements was rendered possible by concentration of motility to the ventral side while

the dorsal body wall was held undeformed as a precondition for shell formation. Formation

of the muscle grid of the foot on the ventral side of the body and the stabilisation by skeletal

elements of the dorsal side caused a shift of the inner organs into a dorsal hump. From the

model for the primitive molluscs with segmented shells the continuation into the conchiferan

constructions with fused shells and the constructional lineages into the major mollusc cons-

tructions are given in captions parallelizing sequences of visualisations of the constructional

stages with the major alterations.

RESUMEN

Se presenta un modelo sobre la evolución de los moluscos basado en la Teoría de Frankfurt,

que concibe a los organismos como unidades hidráulicas transformadoras de energía suje-

tas a transformaciones evolutivas de acuerdo con los principios constructivos que regulan la

organización. La evolución se reconstruye como un proceso de transformaciones construc-

cionales en el que todos los estados se explican como construcciones explícitamente viables;

las fases de transformación irreversibles también se explican racionalmente refiriéndolas a

propiedades construccionales de las máquinas orgánicas.

Se defiende que los predecesores de los moluscos deben haber sido animales alargados,

tipo gusano, que contaban con una malla muscular interna de tal manera que fuese posible

arrastrarse sobre la superficie aplanada ventral del “pie”. Sólo aquellos organismos que

controlasen su sección mediante un sistema muscular densamente espaciado podían empe-

zar a arrastrarse sobre un sustrato duro.

* Naturhistorisches Museum Wien, 3. Zoologische Abteilung, Burgring 7, A-1014 Wien, Austria.

Prof. Gutmann died on April 1997.

Iberus, 15 (2), 1997

La aparición de la rádula se muestra como un proceso dependiente de los movimientos de

reptación que permitieron el anclaje de la estructura al sustrato durante el raspado. La for-

mación de elementos conchíferos fue posible por la concentración de la motilidad en la cara

ventral mientras que la pared dorsal del cuerpo se mantenía sin deformación, paso previo

para la formación de una concha. La aparición de la malla muscular del pie en la cara ven-

tral del cuerpo y la estabilización de la cara dorsal mediante elementos esqueléticos pro-

vocaron el traslado de los órganos internos hacia una ¡joroba dorsal. A partir del modelo

de moluscos primitivos con conchas segmentadas se presenta el desarrollo hacia construc-

ciones conchíferas con conchas fusionadas y hacia las principales estructuras construccio-

nales de moluscos, mediante encabezamientos de sequencias paralelas de estados de de-

sarrollo, incluyendo las principales alteraciones.

KEY WORDS: Molluscs, hydraulic constructions, phylogenetical reconstruction, metamery, radiation.

PALABRAS CLAVE: moluscos, construcciones hidráulicas, reconstrucción filogenética, metamería, radiación.

INTRODUCTION

Evolution can never be directly obser-

ved. It must be reconstructed in models

describing the sequences of transforma-

tional steps. All stages of evolution neces-

sarily have to be represented by explicitly

viable organismic constructions. The the-

oretical concept and the methodology

which allows such reconstructions is the

Frankfurt Evolution Theory (FET)

(GUTMANN, 1974, 1989; BONIK, GRASSHOFF

AND GUTMANN, 1977; GUTMANN AND

EDLINGER, 1994a, b, c, d; EDLINGER, 1989a,

b; EDLINGER, GUITMANN AND WEINGAR-

TEN, 1991; VOGEL, 1991). In accordance

with the demands and results of other bio-

logical disciplines, this theory conceives of

organisms as energy transforming units

and as autoformative constructions.

Reconstructions of the evolutionary

changes in the sense of the Frankfurt Evo-

lutionary Theory are supported by the

insight into the constructional properties

of all animal soft body systems which

function as hydroskeleton apparatuses

and hydraulic units. Before reconstruc-

tions are attempted the principles and

physical laws governing the organismic

entities must be known.

ORGANISMS: ENERGY TRANS-

FORMING ENTITIES

On the basis of constructional expla-

nations metazoan animals must be des-

cribed and explained as self-sustaining

and energy transforming systems. Basi-

cally, they function as machines. They

are capable of actively acquiring matter

and energy from their environment. By

transforming the chemical energy thus

obtained into mechanical force the wor-

king activity of the body construction is

generated.

Energy transformation in the cons-

-tructions is determined by the structure

of specific macromolecular components,

mainly in muscles and cilia. To become

effective as driving engines the energy

transforming structures have to be inte-

grated into an energy cascade of a mecha-

nically coherent structural whole in which

the forces are transmitted to mechanically

working units. A chain of force transmit-

ting structures connects the energy trans-

forming sites with the working units of

the animal body. This chain must never

be interrupted because interruption would

lead to dysfunction and failure of the orga-

nismic construction. The internal activity

is also dependent on the form determi-

ning structural order and on the effective

suppression of useless motoric deforma-

tions by restraining structures.

After the chemo-mechanical energy

transformation on the macromolecular

level the energy is finally utilised in the

machinery for the generation of form,

locomotion, behaviour, and also for

reproduction.

EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions

The constructional concept of orga-

nisation conceives of organismic units

as hydraulic entities that are composed

of enclosing membranes and flexible

integumental walls. The form of every

living body must be enforced by tethe-

ring and or bandaging structures which

suppress the tendency of all living

(hydraulic) units to assume a spherical

shape. In animal constructions the form

is entirely or mainly determined by the

tensile force of muscles and by the mesh

of connective tissue structures which are

organised in highly ordered arrays. In

the course of evolution all transforma-

tion stages must be shown to comply

with the laws of form-enforcement in

hydraulic units. The internal construc-

tional principles and law-like principles

determine the directionality and the

irreversibility of evolutionary change.

The capability of living construc-

tions to function as energy transforming

apparatuses and the ability to obtain an

input of matter and energy are determi-

ned by the structural order of the energy

transforming and working construction.

All stages of evolutionary transforma-

tion series must never lose their structu-

ral order and can only undergo gradual

transformation in a way that the prece-

ding constructional stages open up the

organismic options of the subsequent

alterations. These alterations lead into

constructional alleys with further trans-

formation sequences.

Evolution which is driven by the

activity of the organisms themselves

and the insuppressible generation of

non-directional variation has to follow

very specific internal principles; the step

by step alterations must be forced on

alleys of ordered constructions and on

sequences of non-fortuitous stages.

Because the principles and laws of

organisation can be elucidated in extant

organisms, the transitional sequences can

be reliably reconstructed. So models can

be formulated which are rational in

respect to the methodology applied. Con-

sequently they provide valid explanations

of the intermediate constructional stages.

In all organismic constructions the

course of evolution, mainly of the gross

morphological level, can be shown to de-

pend on the constructional organisation

outlined above. Consequently all minor

physiologically, cytologically, and histo-

logically based functions must be un-

derstood as subservient and dependent

on the conditions given by the construc-

tion as a whole.

Morphological features and so called

morphological characters in the traditio-

nal sense, at all levels of the organisms,

must be seen as enforced by mechanical

structures and as energy transforming

structures.

Constructional morphology and the

reconstruction of evolutionary transfor-

mation do not allow the employment of

traditional methodologies which prescribe

the dismantling and disintegration of

living constructions into morphological

patterns or an array of distinct characters.

Such a procedure is advocated by cladis-

tic “methodologies”. Selection and deli-

mitation of features automatically des-

troys the coherence of organismic systems

and the basic hydraulic constitution. In

living machines constructional coherence,

the hydraulic properties, the order of form-

enforcing structures, and the cataract of

energy transforming structures of orga-

nisms are not accessible to character analy-

sis and comparison of form as advocated

by traditional morphology. Only cons-

tructional alterations based on the recons-

truction of constructionally and functio-

nally viable stages are of interest.

Superficial aspects of similarity of form

and so called homologies in the sense of

traditional morphology are also of no rele-

vance because constructional explanation

has to follow lines of analysis that are com-

parable to engineering procedures.

Such principles preclude the depic-

tion of fortuitous alterations and arbi-

trary morphoclines and allow the deter-

mination and the reconstruction of the

transformation alleys and the establish-

ment of the sequences of stages in the

course of evolution. All evolutionary

alterations must be shown to be delimi-

ted by the internal constructional pro-

perties of the living machines (Fig. 1).

Thus, the constructional alterations of

evolution have to be reconstructed as

Iberus, 15 (2), 1997

REMOVAL OF

WASTE haa

METABOLIC === A GENE EXPRESSION AND

ACTIVITY GENETIC INTERACTIONS

IONIC COMPOSITION ¿== 2

VOLUME REGULATION

PROPERTIES OF MEMBRANES

(SOLIDITY, PERMEABILITY)

CELL SHAPE AND

ARRANGEMENT OR TISSUES

POLYMERS AND

CROSSLINKING PROTEINS

ENERGY

MECHANICAL PROPERTIES AND ACTIVITIES

(VISCOSITY, STIFFNESS, CONTRACTILITY)

CYTOSKELETAL

PATTERN

RYTHM AND

EXTRACELLULAR MATRIX AND AUTODEFORMATION

ADHESIVE FORCES

PERMANENTLY ACTIVE

ORGANISMIC

CONSTRUCTION

MODULATING

PARTIAL SYSTEMS

MEA

Figure 1. Interrelations between the different levels and parts of organisms and between organisms

and their environment (after BEREITER-HAHN, 1991 and EDLINGER, 1995).

Figura 1. Interrelaciones entre los diferentes niveles y partes de organismos y entre los organismos y su

medio ambiente (tomado de BEREITER-HAHN, 1991 y EDLINGER, 1995).

internally guided and directed and cannot

be understood as adaptational processes

or as ruled by environmental factors in the

sense of Darwinian thinking. Evolution in

the sense of the FET, which is certainly the

most radical variant of post-Darwinian

concepts assumes a new status and is

based on the properties of the evolving

organismic entities. Therefore, all Darwi-

nian tenets are excluded from the recons-

tructions as they are not conducive to the

constructional principles which rule orga-

nismic constructions and evolutionary

transformation.

The theoretical tenets and the met-

hodology of the reconstruction procedu-

res were elaborated by a group of

authors (BONIK ET AL., 1977; GUTMANN,

1974, 1989; EDLINGER, 19894, B, 199la, b,

1992a, b, 1994a, b; EDLINGER ET AL., 1991;

VOGEL, 1991) Numerous models

provide corroborating evidence for the

successful application of the methods

and for the validity of the non-Darwi-

nian evolutionary theory.

On the basis of the just outlined con-

cept ontogenetic and phylogenetic deve-

lopment of molluscs (like that of other

“bauplans” of animals) must be recons-

tructed as transformational sequence of

organismic constructions over viable

intermediate constructional forms. The

sequence of evolutionary alterations

must be ruled by the intrinsic laws and

constructional principles which are res-

ponsible for the continuation of energy

transformation and motoric activity in

all stages.

MOLLUSGCS AS A CASE STUDY

In the following the evolution of the

mollusc constructions is presented as a

case study. The stages of mollusc evolu-

tion have to be described and figured

out as tethered systems which in most

cases allow flattening of the foot. In this

way they can exert sufficient control of

body shape by tethering muscles while

the shell structures serve only as defor-

mation suppressing elements in the

dorsal parts of the coherent whole.

THE DERIVATION OF THE MO-

LLUSCS: THE KEY TRANSFORMA-

TION STEPS

In reconstructions based on cons-

tructional morphology the morphocli-

nes and all aspects of directionality of

evolutionary transformation are

derived from the biomechanical princi-

EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions

ples of the living organisms. Thereby,

the arrow of time is firmly supported

by irreversible constructional explana-

tion. On the basis of the constructional

analysis only the derivation of molluscs

from annelid-like forerunners can be

consistently explained (GUTMANN,

1974; EDLINGER, 1991a).

The basic mollusc construction must

certainly have been a creeping creature

with a flattened foot and a dorsal shell

or a sequence of shell elements. The

problem posed by this basic and only

sketchily figured out construction can

be formulated in the following way.

How can such a constructional

constitution arise in skeleton free

hydraulic forerunners of whatever kind

of primitive metazoan organisation?

Which constructional organisation

might have been the incipient stage for

the emergence of the basic properties?

The key changes can only be explai-

ned if one sets the starting point of the

transformation sequence in an annelid

like segmented soft body construction

which was capable of controlling the

cross-section of the body in a way that

creeping on a flattened ventral side

became possible. Flattening must have

been the first stage of adhesive creeping.

The initiation of creeping on a flatte-

ned ventral side was a realistic option of

annelid-constructions with an internal

tethering by dissepimental and other

muscles which traversed the cross

section of the elongate apparatus.

In conjunction with the development

of the adhesively creeping foot and its

anchorage at the substratum the radula

developed in a well demarcated head

region. The use of the radula as a

rasping organ was dependent on the

concomitant formation of the adhesive

creeping foot apparatus. This apparatus

allowed the radula indirectly to be

pressed against the substratum during

feeding. Simultaneously the shells could

come into existence as dorsal stabilising

structures in the non deformed dorsal

portions of the body, while an intensifi-

cation of motility for peristaltic adhesive

creeping occurred on the ventral side.

This constructional constitution is in

some way still existing in extant flatte-

ned chitons.

After the establishment of this cons-

truction with its segmented sequence of

skeletal structures in polyplacophoran-

like organisms new paths of evolution

were viable. They led to worm-like

constructions in a step by step loss of

the shell. In this way some recent forms

of worm-like molluscs, the Ventroplicata

and the Caudofoveata, are easily deriva-

ble from chiton-like predecessors.

Fusion of the shell-plates occurred in

the other major branch. Fusion of the

segmented shells, in accordance with

specific changes of the soft body led to

the formation of the conchiferan cons-

truction with its typical unified shell.

One consequence of this fusion of

the conchiferan shell was that, in con-

junction with the formation of the fused

shell a narrow waist developed

between the cephalopodium and the

dorsal body portion while the internal

organs were restructured in the frame of

a visceral hump.

The basic conchiferan organisation is

to a certain extent represented in the still

extant Tryblidiaceans which display

clear metamerism of some organs. The

basic Conchiferans provided the cons-

tructional basis for a radiative explosion

that generated most molluscan cons-

truction types of the conchiferan level.

The narrowing waist caused and

even enforced reductions especially of

the of the number of' dorsoventral

muscles, gills, and kidneys and the com-

pensatory enlargement of the few per-

sisting organs.

The major steps of transformation

are described and depicted in the illus-

trations. The complete explanation of

the transitions leading to the mollusc

constructions cannot be presented here.

It was elaborated in a sequence of

papers which may be consulted by the

interested and critical reader. In the

following the major steps are depicted

in the illustrations. The ensuing expla-

nation is formulated as a caption in

respect to the illustration (Fig. 2). It

should be stressed that the pictures are

no typical morphological illustrations.

Iberus, 15 (2), 1997

They present visualisations of the array

of form enforcing structures and have to

be conceived as constructional models.

(1.1.) A worm-like predecessor (GUT-

MANN, 1974; EDLINGER, 1991a) with me-

tameric coelomic pouches, dissepiments,

and other segmental tethering elements.

Flattening is made possible by dorsoven-

tral and other muscles on the dissepiments.

All non-longitudinal muscles are helpful

in controlling the cross-section and allow

flattening of the worm-like body in the in-

cipient stages of creeping on hard subs-

trate.

Metamerism which allowed flattening

of the ventral side of the body is manda-

tory in the precursor stages of molluscs.

Flattening of the body is only possible by

internal tethering structures such as mus-

cle bearing dissepiments and internal mus-

cles transcending the coelomic chambers

which were capable of suppressing the

tendency to assume a circular shape. Lack

of cross sectional tethering as observed in

non-metameric worms would result in a

circular cross-section of the soft body and

preclude even the beginning creeping on

hard substrate.

In an early stage of evolution the

mouth became equipped with chitinous

teeth which were located around the sto-

modeum and in the foregut.

(1.2.) Transition to the basic mollusc

construction required the formation of a

thick ventral muscle lattice forming a flat

and highly deformable foot which was

able to follow the contours of the bottom.

This enabled the organisms to creep ad-

hesively and start rasping at the substrate.

In the course of the incipient stages of

transformation the coelom and the other

inner organs were shifted to the dorsal

side. In the dorsal part of the animals co-

elomic metamerism was retained and even

further demarcated by the newly develo-

ping shells.

(1.3.) As muscular activity in the form

of peristaltic waves travelling over the foot

was concentrated in the ventral portion of

the pre-mollusc constructions the dorsal

parts of the body were held mostly un-

deformed. In this situation serial shell ele-

ments developed as economising struc-

tures; they replaced energetically expen-

sive structures such as muscles and con-

nective tissue structures. The newly

developed skeletal structures served as

stabilising elements in the deformable and

still worm-like hydraulic apparatus. As

the shells formed a sequence of indepen-

dent elements bending movements of the

body were still possible; they are remnants

of the worm-like bending and peristaltic

activity of the predecessors. The dorsally

situated serial plates were mechanically

connected to the foot by strong muscles

which continue to demarcate the prece-

ding metamerism of the annelid-like pre-

decessors.

In accordance with the mechanical re-

quirements for the flattening of the foot

the pairs of muscles were arranged as den-

sely spaced pairs of vertical, transverse,

and oblique bundles under each plate.

These muscles project into the dense mus-

cular grid of the foot. Transversely orien-

ted muscles cross from one side to the ot-

her. In this way the form of the whole ani-

mal is under control of the tethering

structures. Lateral extensions of the shells

and their roof-like position allowed the

formation of lateral grooves. In these gro-

oves the metameric gills were established

as projections of the soft body wall. They

were indispensable for the developing mo-

lluscs because the adhesion of the foot to

the bottom and the covering dorsal shells

reduced the body surface usable for res-

piration. Only the lateral parts of the soft

body continued to be exposed to the su-

rrounding medium. The newly develo-

ping gills were forced into the old meta-

meric constellation because the vessels

had to pass between the pre-existing me-

tameric muscle bundles.

Around the shell a spicule bearing

girdle consisting of connective tissue and

musculature was established. This girdle

served as a protection for the lateral gro-

oves with the gills.

Pari passu with the rearrangement of

the musculature the large coelomic cavi-

ties of the annelid like ancestors were res-

trained to a narrow dorsal strip. While the

posterior part of the coelom was altered

EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions

=/=05/2 y

Figure 2. The evolution of chiton-like molluscs out of annelid-like metameric ancestors. An early

branching into chiton-like and conchiferan constructions occurs.

Figura 2. Evolución de los moluscos con forma de quitón a partir de ancestros metaméricos tipo anélido.

Tiene lugar una pronta separación entre construcciones conchíferas y con forma de quitón.

into the pericardium around the effecti-

vely pumping heart the other parts ser-

ved as cavities containing the gonads.

In conjunction with adhesive creeping

the structures around the mouth were per-

fected into a radula by shifting of the chi-

tinous teeth into a ventral pouch of the fo-

regut. The establishment of the radula

must be seen in conjunction with the cre-

eping mode of locomotion. The scraping,

radula could only become effective when

the body remained firmly attached to the

substrate in a way that exertion of pres-

sure by the radula onto the substrate

would not push the animal body away

from the underlying surface. From this fo-

llows that the radula could only develop

in strict interdependence with the cree-

ping performance of the foot.

(1.4.) Reconstructed predecessors of

Conchiferan and Polyplacophoran cons-

tructions. In the anterior portion of the

body the coelom was gradually reduced.

The gut formed lateral pouches in the form

of the midgut-glands, which, besides their

SÍ

Iberus, 15 (2), 1997

Figure 3. Evolutionary transformation of chiton-like predecessor into Placophora, Caudofoveata

and Ventroplicata.

Figura 3. Transformación evolutiva del predecesor tipo quitón en Placophora, Caudofoveata y

Ventroplicata.

digestive function, served as newly formed

fluid filled entities in the frame of the form

enforcing structures and as a substantial

part of the filling of the inner spaces of the

body. The gut pouches were also helpful

in holding the gut in its position. As the

coelomic space was restrained by the mus-

cle construction and receded to narrower

dorsal cavities glandular kidney structu-

res had to develop as extensions of the

metanephridia. They collected excretory

material from the extra coelomic spaces

mainly in the muscle grid. Such an alte-

ration of the metanephridial excretory or-

gans was necessitated by the enlargement

of the extracoelomic fluid fillings of the

muscular grid. The coelom inevitably lost

its function as the sole fluid filling unit of

the body. There can be no doubt that the

evolutionary transformation of the mus-

cle apparatus and the restructuring, of the

excretory system were coupled and mu-

tually dependent.

(2.1.) The Polyplacophoran like tran-

sitional stage. The model represents an

early organisational constellation in mo-

lluscan evolution. These forms retained

the metameric organisation of muscle

system, shell arrangement, gills, and

kidneys (Fig. 3).

(2.1.1.) The fully developed Polypla-

cophoran constructions represent a side

EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions

Figure 4. The radiation of conchiferan constructions resulting in diverging organismic types which

are capable of occupying different environmental conditions.

Figura 4. Radiación de construcciones conchíferas que resulta en tipos de organismos divergentes capaces

de ocupar diferentes condiciones ambientales.

branch of primitive molluscs. Flattening of

the shells became more pronounced. The

serial shell plates continue to preserve the

metameric arrangement of the dorsoven-

tral musculature. In the course of flattening,

the circulatory and the gill systems were

restructured in a way allowing the non-

metameric array of the gills and decou-

pling of internal metamerism and gill-po-

sition. This is unique in Polyplacophorans

and not representative for the transition to

the molluscs with fused shells.

(2.1.2.) Evolution of the Caudofovea-

tan construction is part of a radiative

differentiation of the polyplacophoran

organization. This lineage must have

started from polyplacophoran forms

(EDLINGER, 1989b) which underwent re-

duction of the shells. Concomitantly

with shell reduction the lattice-like mus-

cle foot was only retained in the anterior

portion of the worm like body. The gut

was also considerably altered; in the

rear portion the intestinal canal was su-

rrounded by the midgut-glands, which

held the gut in its position. In these bu-

rrowing constructions only one poste-

rior pair of gills was retained in the hind

part of the body. The resulting worm-

like form developed independently of

the Ventroplicatan constructions.

(2.1.3.) Evolution of the Ventroplicatan

construction is also a branch of Polypla-

cophoran radiation: loss of shell structu-

res in conjunction with an enlargement of

the girdle resulted in the formation of a

worm-like body. The foot-complex per-

sisted as a narrow groove running along

the ventral side of the whole body length.

The groove remains internally tethered

by the dorsoventral muscle bundles which

are certainly rudiments of the serial shell

related retractors. In contrast to the bu-

rrowing Caudofoveata the Ventroplicata

are capable of climbing in “tree-like” en-

vironments. To corroborate the derivation

of the worm-like forms from Polyplacop-

horans it should be kept in mind that some

extant Polyplacophorans as Cryptoplax

show an observable tendency to reduce the

shells.

(2.2.) In the evolutionary alley to the

Conchiferan constructions (Fig. 4) the Mo-

noplacophorans demarcate a strategic

' intermediate stage.

Iberus, 15 (2), 1997

(2.2.1.2.)

Figure 5. Transformation of monoplacophoran-like ancestors into scaphopods by extreme dorso-

ventral elongation.

Figura 5. Transformación de ancestros tipo monoplacóforo en escafópodos mediante una elongación dor-

soventral extrema.

The unified shell was formed by fu-

sion of the segmental skeletal elements.

This process ensued in the deepening of

the groove between the cephalopodium

and the shell-covered visceral hump. The

waist became more pronounced and the-

reby the cephalopodium was rendered

more flexible in respect to the shell. In the

Monoplacophora the metameric soft

body organization of muscles, gills,

nephridia and portions of the coelom is

still retained. Metamerism is obviously

fading from the rostral portion of the

body (GUTMANN, 1974; EDLINGER, 1991a).

Noticeable is the concentration of the an-

cestral structures, coelom, sac bearing

nephridia, and gills in the rear part of the

body. This situation is indicative of the

transition to the other conchiferan cons-

tructions which sprang from a radiative

divergence of constructions.

(2.2.1.) The fully developed Conchife-

ran Constructions.

In all conchiferan constructions with

fused shells the separation of the two

body-portions, the cephalopodium and

the dorsal hump, by a waist allows free

movement of the foot and the undeformed

posture of the visceral sack with the stiff

shell frame far from the substrate. The for-

mation of the waist enforced the reduction

of the number of gill-pairs in all derived

conchiferan constructions.

(2.2.1.1.) Neopilinida.

The Neopilinida are flattened recent

Monoplacophoran constructions with clear

remnants of metamerism in the arrange-

ment of muscles, nephridia and gills. So

they must be early representatives of the

transition phase to the Conchiferans. Ho-

wever, clear indications of reduction of

metamerism from the anterior portion of

the body become evident; the anterior gills

are reduced and the muscle bundles fused

(GUTMANN, 1974; EDLINGER, 1991a).

(2.2.1.2.) Evolution of Scaphopod cons-

tructions.

Scaphopods are derived from a Mo-

noplacophoran stage with a posterior slit

in the shell for the expulsion of faeces. A

dorsoventral elongation of the construc-

tion caused the reduction of the number

of muscles and gill pairs and the forma-

tion of a tube-like shell. The foot changes

its form by a rearrangement of the inter-

nal muscle-lattice and becomes a burro-

wing organ. The posterior slit of the shell

became an elongate hole. This hole enabled

the animals to ventilate their mantle cavity

when they penetrated into the substrate.

The water is sucked in from the anterior

region and expelled through the poste-

rior hole. The total reduction of the gills

is enforced by the narrowing of the man-

tle cavity when the elongate organisms

developed (EDLINGER, 1991b) (Fig. 5).

EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions

(2.2.1.3.)

Figure 6. The evolution of the gastropod construction by torsion. Torsion occurs after the forma-

tion of a waist between the cephalopodium and the visceral hump and reduction of most of doso-

ventral muscles.

Figura 6. La evolución de la estructura de un gasterópodo mediante torsión. Ésta ocurre tras la forma-

ción de un estrechamiento entre el cefalopodio y el asa visceral, y la reducción de la mayoría de los mús-

culos dorsoventrales.

(2.2.1.3.) Evolution of Gastropods (ED- ceral hump after a very narrow waist had

LINGER, 1988a, 1988b, 1989a). formed and most of the gill pairs had been

The most significant event of gastro- reduced. Torsion led to an anterior posi-

pod evolution was the torsion of the vis- tion of the partially reduced mantle ca-

Iberus, 15 (2), 1997

(2.2.1.4.)

(2.2.1.4.2.)

Figure 7. The evolution of cephalopods characterized by the appearance of gas-filled spaces at the

top of the shell and by a radical rearrangement of the musculature of the cephalopodium with

strong muscular arms and suckers.

Figura 7. Evolución de los cefalópodos, caracterizada por la aparición de cámaras llenas de gas en la parte

alta de la concha y una redistribución radical de la musculatura del cefalopodio con fuertes ramas mus-

culares y ventosas.

vity with the remaining gills and the for-

mation of the chiasma of the lateral nerve

cords. The precondition for torsion lies in

a step by step reduction of dorsoventral

muscle bundles of a Monoplacophoran

ancestor and by narrowing of the waist

between the cephalopodium and the vis-

ceral hump. This process is connected with

a step by step spiralling up of the shell. The

persistence of only one pair of crossing

obliquely transversal muscles could cause

the torsion. Totally bilaterally organised

symmetric Bellerophontacean shells are

representative of this process (Fig. 6).

(2.2.1.3.1) Radiation of Gastropod cons-

tructions: All gastropod lineages are de-

rived from a Bellerophontacean stage with

a bilateral and helical shell, which posses-

sed slits or a series of holes in the poste-

rior part of the shell for the expulsion of

faeces. Only one pair of parallel dorso-

ventral muscles persisted. Asymmetry

could arise by the partial or total reduction

of one of the dorsoventral muscles and

the change of the planispiral shell of Be-

llerophon-like ancestors to an asymme-

tric helical form. In most cases the gills be-

came also unequal or one gill was entirely

reduced (EDLINGER, 1988a, 1988b, 19894a).

Flattening and secondary reduction of

the coil can result in the formation of a cup-

like shell. Flattening must cause a change

in the muscle arrangement. The dorso-

ventral muscles and their scares on the

inner of the shells were enlarged to form

EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions

Figure 8. The evolution of bivalves by a rearrangement of the dorsoventral muscles and a division

of the shell to a bivalve one.

Figura 8. Evolución de los bivalvos por medio de una redistribución de los músculos dorsoventrales y una

división de la concha hacia una concha bivalva.

horseshoe-like insertions. The gills can

persist as in Fissurella. In this case the re-

tractor muscle will form a homogeneous

horse-shoe-like entity. If the gills are also

reduced and substituted by secondary

gill-like structures the muscles are splitted

up into a Monoplacophoran-like situation

of Patella (EDLINGER, 1988a, 1988b, 1989a;

HASZPRUNAR, 1988).

(2.2.1.4.) Evolution of Cephalopod

constructions (BONIK, GRASSHOFE, GUT-

MANN AND KLEIN-RÓDDER, 1977). The de-

velopment of gas-filled chambers in the

apical part of the shell rendered the orga-

nisms more buoyant. The foot with its de-

formability and the adhesive capabilities

was altered into a system of muscular

arms with suckers which could grasp prey.

Retraction of the soft-body into the shell

inevitably caused expulsion of a water-

current. In the course of evolution im-

provement of this mechanisms was alte-

red into effective jet propulsion when the

hind part of the foot was modified to form

a narrow funnel through which the water

expelled from the mantle cavity was con-

centrated to enhance the jet effect. This

constructional situation can be observed

in all cephalopods (Fig. 7). An early bi-

furcation or diphyletic development gave

rise to the branches of endocochlean and

exocochlean Cephalopod construction

(2.2.1.4.1, 2.2.1.4.2).

(2.2.1.5.) Evolution of Bivalve Cons-

tructions (VOGEL AND GUTMANN, 1980).

The origin of Bivalves can be understood

as a rearrangement of the dorsoventral

muscles in a high chambered Monopla-

cophoran ancestor. A considerable por-

tion of the dorsoventrally and obliquely

arranged muscles in the foot were shif-

ted into a horizontal position connecting

not the shell and the foot as in the former

stages but the two laterally bent down

plates of an evolving bivalved shell.

Contraction of the muscles brought the

flanks of the shells together in protective

behaviour. Most of the vertical (dorso-

ventral) portions of the musculature re-

tained their original retractile function in

respect to the foot but the number of re-

tractors was reduced. Consequently

most of the gills disappeared with one

pair remaining.

The remaining pair of gills are utilised

as filterfeeding devices because lateral

cephalic lobes bridged the gap between the

mouth and the gills and formed conveyer

belts for the transport of food from the

gills to the mouth (Fig. 8).

Iberus, 15 (2), 1997

DISCUSSION AND CONCLUSION

Itis very obvious that in the perspec-

tive of constructional morphology and

organismic evolution character-analysis

and the establishment of sequences of

organismic forms in the sense of homo-

logies are inconclusive and arbitrary.

They pose problems but do not provide

answers because description of form and

character analysis do not lead to cons-

tructional insight or an understanding of

form determination. Constructional alte-

ration in evolution can not reasonably be

derived from genetic or other molecular

features (GHISELIN, 1988; WAGELE, 1994;

WAGELE AND WETZEL, 1994; EDLINGER,

1995). The understanding of organisms

as constructions leaves no doubt that the

constructional configuration determines

evolutionary change of the living machi-

nery and prescribes the sequence of

constructional stages and of irreversible

steps. There is little freedom for contin-

gency in the order of the “bauplan” con-

figurations and no encouragement for

simple description and form compari-

son. Nothing useful can come from such

traditional approaches which are blind

to causal aspects and insensitive for ex-

planatory principles.

When organisms are conceived as

energy transforming constructions evo-

lution must be ruled by constructional

principles and not by subjective “gestalt”

properties or subservient molecular me-

chanisms. As could be shown in the fore-

going context strict obeisance of the cons-

tructional principles allows the rejection

of constructionally impossible or impro-

bable alternatives. From the methodology

applied and the mode of reconstruction of

the evolutionary transformations just ad-

vocated follows that traditional phyloge-

netic concepts, form sequences, and cla-

dograms based on usual procedures are

not considered valid. Therefore, traditio-

nal hypotheses are bypassed and left out

of consideration when the criteria of the

Frankfurt-Theory are applied. If some-

body feels the need to adhere to the idea

that Plathelminths, non-segmented

worms, or Nemerteans might be the pre-

cursors of mollusc he or she should pre-

sent a continuous model with a strict ex-

planation of the intermediate construc-

tional steps.

The genes and other molecular me-

chanisms which are subservient in relation

to the living constructions and their struc-

tures have to comply with the construc-

tional requirements and must follow cons-

tructional modifications of the machinery

in the course of structural reorganisation.

Taken as separate features molecular and

physiological mechanisms are not useful

for the elucidation of constructional change

in evolution. Therefore, constructional

morphology neglects all traditional sug-

gestions as to the affinity of organismic

groups based on form similarities in the

sense of homologies and on subservient

and constructionally dependent molecu-

lar and physiological properties.

Itis not possible to give a list of all the

published sequences of forms which were

figured out to represent what all the aut-

hors tried to suggest as stages of mollusk

evolution (SCHELTEMA, 1978; GÓTTING,

1980a, 1980b; HAas, 1981; BANDEL, 1983;

LAUTERBACH, 1983a, b; HASZPRUNAR, 1988,

1992a, b). Many contributions were for-

mulated by paleontologists (RUNNEGAR

AND POJETA, 1974; RUNNEGAR, POJETA,

NOEL, TAYLOR, TAYLOR AND MCCLUNG,

1975; MAREK AND YOCHELSON, 1976; Yo-

CHELSON, FLOWER AND WEBERS, 1973; Po-

JETAJR., 1987) who started from one or the

other fossilstructure. Most morphoclines

presented since the last century are highly

contradictory, however, all of them have

in common an access that arbitrarily selects

some features of skeletal or soft body struc-

tures.

Organisms as constructionally res-

trained and not deliberately transforma-

ble entities are not even expected. In none

of the published morphoclines is the di-

rectionality of evolutionary transforma-

tion elaborated or explained as irreversi-

ble. Organisms are misconceived as sha-

ped and freely modifiable pieces of art.

Fruitful discussion can only start when

alternative models with continuous ex-

planations and well supported transfor-

mational polarities are given. In the pre-

sent situation the discussion of all the

morphoclines would require the sacrifi-

EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions

cium intellectus. Also in the field of ra-

diation of the phylum Mollusca recons-

tructions have to be done on the consistent

basement of constructional laws and so-

lid criteria of directionality of evolutio-

REFERENCES

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O Sociedad Española de Malacología

Iberus, 15 (2): 67-74, 1997

Mollusc fauna of the medium high mountain ranges of the

Hungarian Holocene: a zoogeographical research

Fauna de moluscos de media y alta montaña del Holoceno de Hun-

gría: una investigación zoogeográfica

Levente FÚKOH*

Recibido el 30-X-1995. Aceptado el 15-11-1997

ABSTRACT

An attempt is made to complete the zoogeographical studies of the mollusc fauna of the

medium-high mountain ranges of the Hungarian Holocene by analysing twenty-eight chrono-

logically and biostratigraphically known faunae. Eighty four species are classified in nine fau-

nalcentres and four biozones (Vallonia costata, Clausillidae, Granaria frumentum and Helici-

gonia faustina — Acicula polita). A brief discussion is made on the abundance of several

species of each faunal-centre. The picture drawn from the fauna agrees with the geographical

position and geomorphological conditions of Hungary (Carpatian Basin, Central-Europe)

RESUMEN

Se pretende completar los estudios zoogeográficos de la fauna de moluscos de media y alta

montaña del Holoceno de Hungría mediante el análisis de ventiocho faunas conocidas tanto

cronológica como estratigráficamente. Ochenta y cuatro especies fueron clasificadas en

nueve “centros faunísticos” y cuatro biozonas (Vallonia costata, Clausillidae, Granaria fru-

mentum and Helicigonia faustina — Acicula polita). Se hace una breve discusión sobre la

abundancia de varias especies en cada “centro faunístico”. La representación que se obtiene

de la fauna coincide con la posición geográfica y las condiciones geomorfológicas de Hun-

gría (planicie de los Cárpatos, Europa central).

KEY WORDS: Molluscs, Zoogeography, Holocene, Hungary.

PALABRAS CLAVE: Moluscos, Zoogeografía, Holoceno, Hungría.

INTRODUCTION

The paleoecological and biostratigrap-

hical studies of the mollusc fauna of the

medium-high mountain ranges of the Hun-

garian Holocene (Fig. 1) has been comple-

ted in the last few years. (FUKOH, 1991,

1992a, 1992b, 1993a, 1993b). Although

during these estudies, zoogeographical

examinations were carried out for certain

faunae (FUKOH, 1983, 1989; BABA AND

FUKOH, 1984), a comprehensive view is

lacking. The aim of this paper is to complete

the lack of information about this subject.

The analysis of twenty-two chronolo-

gically and biostratigraphically known

faunae (FUKOH, KROLOPP AND SÚMEGI,

1995) is completed in this work (Fig. 2), in-

creasing the number of species previously

cited from 81 (Table 1) to 84 (Table III).

* Mátra Museum, H-3200 Gyóngyós, Kossuth u. 40, Hungary.

Iberus, 15 (2), 1997

MEM(12%)

MA(4%)

AMe(1%)

PM(30%)

Figure 1. Map showing the position of the localities of the Biozones stratotypes and the zoogeo-

graphical distribution of the Holocene mollusc fauna in the Hungarian medium high mountain

area. A: Búkk Mts.; B: Aggtelek karst; C: Bakony Mts. Faunal centres, PM: Ponto-mediterranean;

SA: Siberian-Asiatic; MEM: Middle-European-Mountain; HM: Holomediterranean; AM: Adriato-

Mediterranean; EM: European-Mountain; MA: Middle-Asiatic; CS: Caspi-Sarmatian; AMe:

Atlanto-Mediterranean. Biozones, 1: Vallonía costata biozone; 2: Clausiliidae biozone; 3: Granaria

frumentum biozone; 4: Helicigona faustina — Acicula polita biozone.

Figura 1. Mapa mostrando la posición de las localidades de los estratotipos de las biozonas y la distribu-

ción zoogeográfica de la fauna de moluscos de media y alta montaña del Holoceno de Hungría. A: Montes

Búkk; B: karst de Aggtelek; C: Montes Bakony. “Centros faunísticos”, PM: ponto-mediterráneo; SA: sibe-

riano asiático; MEM: media montaña europea; HM: holomediterráneo; AM: adriatico-mediterráneo;

EM: montaña europea; MA: medio asiático; CS: caspiano-samartiano; AMe: atlanto-mediterráneo.

Biozonas, 1: biozona de Vallonia costata; 2: biozona de Clausiliidae; 3: biozona de Granaria frumen-

tum; 4: biozona de Helicigona faustina — Acicula polita.

MATERIAL AND METHODS distribution of the species. This metho-

dology was chosen because, according to

the preliminary examinations and calcu-

lations, it is more suitable when resear-

ching fossil materials.

The methodology used to carry out the

classification presented in this paper is the

same employed to situate zoogeographi-

cally the species of the Middle-European

faunae. These methods can be divided into

two main groups as follows:

1. Methods based on recent distribu-

tion of the species (KERNEY, CAMERON AND

JUNGBLUTH, 1983; FLASAR, 1971; KÓRNIG,

1983; ALEXANDROWITZ, 1983, 1984; FRANK,

1988, 1990, 1992a, 1992b)

2. Derivative method (BABA, 1982).

This method is based on recent and fossil

RESULTS AND CONCLUSSIONS

Table II shows the distribution in num-

ber of the 84 mollusc species found in the

studied area by faunal-centres and biozo-

nes. Figure 3 illustrates the zoogeograp-

hical distribution in number of the Holo-

cene and recent mollusc species.

FEÚKOH: Mollusc fauna of the medium high mountain ranges of the Hungarian Holocene

Boreal Atlantic Sub-Boreal Sub-Atlantic

8 7 6 5 4 3 2 1 0

1000 radiocarbon year BP. (1950)

Vallonia costata Clausiliidae Granaria frumentum Helicigona faustina-

zone zone zone Acicula polita zone

Muflon 1/9-6

Mutflon 1/5-2

| Muflon 1/1

Muflon 11/3-2

Kajla-bérc 3

Kolyuk 11/17-12 Kajla-bérc 2-1

| Kólyuk 11/10-1

Rejtek 1/11

Rejtek 1/II

Szentléleki-v.

Horvatí-h. 4-3

Szalajka 11

Szalajka 5

Szalajka 3a

Szalajka 3b-1

Monosbél

Petényi H5 Petényi H3

Kis-kohát 4

Csunya-v 1/2

Csunya-v 111

Nagyoldal 6

Nagyoldal 5-4

Szentgál

Háromágú

Rigó-h

Baradla

Figure 2. Biostratigraphic identification of significant terrestric Holocene localities of Hungary (twenty-

eight chronologically and biostratigraphically known faunae). Biúikk Mts.: Muflon”-cave, “Kajla-

bérc”-cave, “Kólyuk IT”-cave, “Rejtek ”-cave, “Horvéti”-hole, “Petényi”-cave, “Csunya”-valley rock

shelter, “Csunya”-valley I=rock shelter, “Háromágu”-cave, “Kajla-bérc”-cave, “Szalajka”-valley-tra-

vertine, “Monosbél”-travertine, “Szentléleki”-valley-rock shelter, “Kálmán-rét”-shaft-cave, “Kiskóhát”-

shaft-cave. Aggtelek karst: “Baradla”-cave, “Nagy-oldal”-shaft-cave. Bakony Mts: Szentgál, Mecsek-

hill, “K6-lik”-cave, “Rigo”-hole. Lines represent the biostratigraphic and chronologic extension and/or

connection of localities (shown in Figure 1).

Figura 2. Identificación bivestratigráfica de las localidades terrestres más significantes del Holoceno de

Hungría (ventiocho faunas conocidas cronológica y bivestratigráficamente. Montes Búkk: cueva “Muflon”,

cueva “Kajla-bérc”, cueva “Kólyuk II”, cueva “Rejtek I”, sima “Horvéti”, cueva “Petényi”, abrigo rocoso

“Csunya”-valley L, abrigo rocoso “Csunya”-valley IL, cueva “Háromágu”, cueva “Kajla-bérc”, valle “Sza-

lajka” travertino, “Monosbél” travertino, valle “Szentléleki” abrigo rocoso, pozo “Kálmán-rét”, pozo

“Kiskóhát”. Karst Aggtelek: cueva “Baradla”, pozo “Nagy-oldal”. Montes Bakony: Szentgál, Mecsek-

hill, cueva “Kó-lik”, sima “Rigo”. Las líneas representan la extensión y/o conexión bioestratigráfica y cro-

nológica de las localidades (mostradas en la Figura 1).

Iberus, 15 (2), 1997

Table I: Distribution of 81 molluscs species found the medium-high mountain ranges of the

Hungarian Holocene fauna, according to several authors. Authors, 1: Verney, Cameron and

Jungbluth; 2: Flasar; 3: Alexandrowicz; 4: Kórnig; 5: Frank; 6: Bába. Abbreviations, a: Alpian; adm:

Adriatic-Mediterranean; e: European; h: Holarctic; hm: Holomediterranean; k: Carpatian; ksz:

Capian-Sarmatian; m: middle; ma: Middle-Asiatic; med: Mediterranean; merid: meridional; p:

Palearctic; po: Pontomediterranean; s, n, o, w, the four cardinal points; sza: Siberian-Asiatic; ws:

West-Sibiric; sz: Siberian.

Tabla 1. Distribución de 81 especies de moluscos encontradas en media y alta montaña de la fauna del

Holoceno de Hungría, de acuerdo con distintos autores. Autores, 1: Verney, Cameron and Jungbluth; 2:

Flasar; 3: Alexandrowicz; 4: Kórnig; 5: Frank; 6: Bába. Abreviaturas, a: alpino; adm: adriatico-medi-

terráneo; e: europeo; h: holártico; hm: holomediterráneo; k: carpatiano; ksz: capiano-samartiano; m:

medio; ma: medioasiático; med: mediterráneo; merid: meridional; p: paleártico; po: pontomediterráneo;

s, n, o, w: los cuatro puntos cardinales; sza: siberiano-asíatico; ws: sibírico oeste; sz: siberiano.

Species 1 2 3 4 5 6

Achantinula aculeata (0. F. Miller, 1774) wp WSZ

Acicula polita (Harimann, 1840) e-0 me me me me-a po

Aegopinella minor (Stabile, 1864) some me-se soe some po

Aegopinella pura (Alder, 1830) e e e e WSZ

Bradybaena fruticum (0. F. Mller, 1774) moe-a e e-a 05Z

Bulgarica vetusta (Rossmíssler, 1836) s08 po

Carychium minimum (0. E. Múller, 1774) e-S7 e-sz e e-S7 057

Carychium tridentatum (Risso, 1826) e se e e hm

Cecilioides acicula (0. E. Múller, 1774) med-we wme

Cepea vindobonensis (Férussac, 1821) soe soe soe ksz

Chondrina clienta (Westwrlund, 1883) s08-0 s08-0 08-a po

Chondrula tridens (0. E. Múller, 1774) msoe pm msoe hm

Clausilia cruciata (Studer, 1820) ne-a ba b-a moe-a e

Clausilia dubia Draparnaud, 1805 me me me me me po

Clousilia pumila C. Pfeiffer, 1828 moe me oe ome moe po

Cochlicopa lubrica (0. E. Miller, 1774) h h ws h h h

Cochlicopa lubricella (Porro, 1837) h h h h ma

Cochlodina cerata (Rossm(ssler, 1836) k wk me

Cochlodina laminata (Montagu, 1803) e e ws e e me

Cochlodina orthostoma (Menke, 1830) moe 0€ me

Columella edentula (Draparnaud, 1801) h h h 0-Sz

Daudebardia brevipes (Draparnaud, 1805) mse mse po

Daudebardia helenae Fúkúh, 1985 me

Daudebardia rufa (Draparnaud, 1805) mse med-me sme po

Discus perspectivus (Múhlfeld, 1816) a-ok a-ok a-ok po

Discus rotundatus (0. E. Miller, 1774) wme wsm WS wme adm

Discus ruderatus (Férussac, 1821) p p p p 0-52

Ena montana (Draparnaud, 1801) me-k-a me me me-k e

Ena obscura (0. E. Múller, 1774) e e e hm

Euconulus fulvus (0. E. Miller, 1774) h h h h h

Evomphalia strigella (Draparnaud, 1801) me ome ome ome Ksz

Granaria frumentum (Draparnaud, 1801) med merid moe po

Helicella obvia (Menke, 1828) s08 smoe

Helicigona faustina (Rossmíssler, 1838) k me

Helicodonta obvoluta (0. E. Miller, 1774) me merid merid sme adm

EÚKOH: Mollusc fauna of the medium high mountain ranges of the Hungarian Holocene

Table 1. (Continuation).

Tabla I. (Continuación).

Species 1 2 3 4 5 6

Helicopsis striata (0. E. Miller, 1774) wmoe e

Helix pomatia Linné, 1758 smoe soe me soe msoe po

Isognomostoma isognomostoma (Schróter, 1784) a-k me a-k a-k a-k me

Laciniaria biplicata (Montagu, 1803) me-b some po

Laciniaria plicata (Draparnaud, 1801) moe me me moe po

Macrogastra latestriata (A. Schmidt, 1857) k

Macrogastra plicatula (Draparnaud, 1801) me e e e po

Macrogastra ventricosa (Draparnaud, 1801) me e me e e po

Monacha cartusiana (0. E. Múller, 1774) po

Nesovitrea hammonis (Siróm, 1765) p p h 0-SZ

Orcula doliolum (Bruguiére, 1792) s0€ me merid soe ma

Orcula dolium (Draparnaud, 1801) a-k a-wk a-k po

Oxychilus depressus (Sterki, 1880) a-k me a-k po

Oxychilus draparnaudi (Beck, 1837) med-we we

Oxychilus glaber (Rossm(ssler, 1838) sme smoe so€ a-se po

Oxychilus inopinatus (Ulicny, 1887) k

Oxychilus orientalis (Clessin, 1887) k k me

Perforatella incarnata (0. F. Miller, 1774) msoe mwe me msoe po

Perforatella vicina (Rossm(ssler, 1842) e

Punctum pygmaeum (Draparnaud, 1801) h p p h h

Pupilla muscorum (Linné, 1758) h h 0-57

Pupilla triplicata (Studer, 1820) 08-0 e-a adm

Pyramidula rupestris (Draparnaud, 1801) we-M oe m-a m-a ma

Ruthenica filograna (Rossm(ssler, 1836) 0€ oe moe moe me

Semilimax kotulai (Westerlund, 1871) a-k a-k a-k

Semilimax semilimax (Férussac, 1802) a-me a-me

Trichia unidentata (Draparnaud, 1805) a-k oa-wk oa-wk oa-k me

Trichia hispida (Linné, 1785) e e

Truncatellina claustralis (Gredler, 1856) m-sa m m hm

Truncatellina cylindrica (Ferrussac, 1807) se se se se hm

Vallonia costata (0. E. Múller, 1774) h h h h h h

Vallonia pulchella (0. F. Múller, 1774) h h h h h h

Vallonia enniensis (Gredler, 1856) mse me

Vertigo alpestris Alder, 1838 na p 0-Sz

Vertigo angustior Jeffreys, 1830 e e ksz

Vertigo parcedentata (A. Braun, 1847) sz0

Vertigo antivertigo (Draparnaud, 1801) p ksz

Vertigo pusilla (0. E. Múller, 1774) e e e-was hm

Vertigo pygmaea (Draparnaud, 1801) h h h WSZ

Vertigo substriata (Jeffreys, 1833) a 0-e

Vitrea contracta (Westwrlund, 1871) p hm

Vitrea crystallina (0. F. Miller, 1774) e e ws e e adm

Vitrea diaphana (Studer, 1820) a-k a-merid a-me po

Vitrina pellucida (0. E Múller, 1774) h p p p h

Zebrina detrita (0. F. Múller, 1774) s08 po

Zonitoides nitidus (0. E Miller, 1774) h h

Iberus, 15 (2), 1997

Table II. Distribution in number of 84 mollusc species in the Hungarian Holocene fauna of the

medium-high mountain ranges by Faunal-centres and Biozones. Abbreviations, 1: Vallonia costata

biozone; 2: Clausiliidae biozone; 3: Granaria frumentum biozone; 4: Helicigona faustina — Acicula

polita biozone.

Tabla 1. Distribución en número de las 84 especies de moluscos de la fauna del Holoceno de Hungría en

media y alta montaña agrupadas por “centros faunísticos” y biozonas. Abreviaturas, 1: biozona de

Vallonia costata; 2: biozona de Clausiliidae; 3: biozona de Granaria frumentum; 4: biozona de

Helicigona faustina — Acicula polita.

BIOZONES

FAUNAL-CENTRES No. of species 1 2 3 4

Siberian-Asiatic 21 15 19 16 16

Middle-Asiatic 3 3 2 7 2

Caspi-Sarmatian a) 3 4 2 3

Pontomediterranean 2 18 19 18 22

Adriato-Mediterranean 5 5 4 4 5

Atlanto-Mediterranean 1 0 1 0 0

Holomediterranean o o) 5 5 5

European -Mountain 7 4 3 2 4

Middle-Eur. -Mountain 11 8 o 6 9

TOTAL 84 62 63 5 66

Number of species

Faunal centres

Figure 3. Zoogeographical distribution in number of the mollusc fauna of the Hungarian medium

high mountain ranges considering Holocene and Recent species separately. 1: Pontomediterranean;

2: Siberian-Asiatic; 3: Middle-European-Mountain; 4: Holomediterranean; 5: Adriato-

Mediterranean; 6: European-Mountain; 7: Middle-Asiatic; 8: Caspi-Sarmatian; 9: Atlanto-

Mediterranean.

Figura 2. Distribución zoogeográfica en número de la fauna de moluscos de media y alta montaña de

Hungría, considerando tanto las especies del Holoceno como las recientes. 1: pontomediterráneo; 2: sibe-

riano asiático; 3: media montaña europea; 4: holomediterráneo; 5: adriático mediterráneo; 6: montaña

europeo; 7: medio asiático; 8: caspiano-samartiano; 9: atlanto-mediterráneo.

FÚKOH: Mollusc fauna of the medium high mountain ranges of the Hungarian Holocene

Table IM. Zoogeographical distribution of the Hungarian Holocene species by faunal-centres. (*)

Molluscan species not cited in Table 1.

Tabla 111. Distribución zoogeográfica de las especies del Holoceno de Hungría por “centros faunásticos”.

(%) Especies de moluscos no citadas en la Tabla 1.

1. Siberian-Asiatic faunal-centres 21species

Achantinula aculeata Euconulus fulvus Vallonia pulchella

Aegopinella pura Limax maximus Linné, 1758 (*) Vertigo alpestris

Bradybaena fruticum Nesovitrea hammonis Vertigo parcedentata

Carychium minimum Punctum pygmaeum Vertigo pusilla

Cochlicopa lubrica Pupilla muscorum Vertigo pygmaea

Columella edentula Vallonia costata Vitrina pellucida

Discus ruderatus Vallonia enniensis Zonitoides nitidus

2. Middle-Asiatic faunal centres 3 species

Cochlicopa lubricella Orcula doliolum Pyramidula rupestris

3. Caspian-Sarmatian faunal-centres 5 species

Cepaea vindobonensis Semilimax kotulai Vertigo antivertigo

Evomphalio strigella Vertigo angustior

4. Ponto-Mediterranean faunal-centres

25 species

Acicula polita Granaria frumentum Macrogastra ventricosa

Aegopinella minor Helix pomatia Monacha cartusiana

Bulgarica vetusta Helicella obvia Orcula dolium

Chondrina clienta Zebrina detrita Oxychilus depressus

Clausilia dubia Helicopsis striata Oxychilus glaber

Clousilia pumila Laciniaria plicata Perforatella incarnata

Doudebardia brevipes Laciniaria biplicata Vitrea diaphana

Daudebardia ruta Macrogastra plicatula Vitrea subrimata (Reinhardt, 1871)(*)

Discus perspectivus

5. Adriatic-Mediterranean faunal-centres 5 species

Cecilioides acicula Helicodonta obvoluta Vitrea crystallina

Discus rotundatus Pupilla triplicata

6. Atlantic-Mediterranean faunal-centres 1 species

Semilimax semilimax

7. Holomediterranean faunal-centres 6 species

Corychium tridentatum Ena obscura Truncatellina cylindrica

Chondrula tridens Truncotellina claustralis Vitrea contracta

8. European-Mountain faunal-centres 7 species

Clausilia cruciata Oxychilus inopinatus Trichia hispida

Ena montana Perforatella vicina Vertigo substriata

Macrogastra latestriata

9. Middle-European-Mountain faunal-centres 11 species

Cochlodina orthostoma Helicigona faustina Oxychilus draparnaudi

Cochlodina cerata Trichia unidentata Oxychilus orientalis

Cochlodina laminata Isognomostoma isognomostoma Ruthenica filograna

Daudebardia helenae Laciniaria turgida (Rossmassler, 1836) (*)

Iberus, 15 (2), 1997

The zoogeographical distribution of

species by Faunal-centres is summarised

in Table II.

The formation of the characteristic

zoogeographical conditions of the recent

fauna began after the last cold period of

the Pleistocene. This can be stated on the

basis of the relative abundance analyses

of the faunae (84 species) situated in nine

faunal-centres (Tables II and III). Prima-

rily, the abundance of Subatlantic species

has increased during the last ten thou-

sand years. Ponto-Mediterranean species

are the most important and can be obser-

ved almost in all every biostratigraphical

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ALEXANDROWICZ, S. W., 1984. Srodkowoholo-

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BABA, K., 1982. Eine neue zoogeographische

Gruppierung der Ungarische Landmollusken

und die Wertung des Faunabildes. Malaco-

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BABA, K. and FUKÓH, L., 1984. Holocén és re-

cens malakológiai adatok értékelése állatfol-

drajzi és 0kológiai módszerekkel a Búukkben.

Malakologiai Tájékoztató (Malacological News-

letter), 4: 42-53.

FLASAR, I., 1971. Zur Malakofauna des nordós-

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FRANK, C., 1988. Aquatische und Terrestrische

Mollusken der Osterreichischen Donau-auen-

gebiete und der Angrenzenden Biotope. Teil

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terreichisch-Deutschen Staatsgrenze bis Linz.

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FRANK, C., 1990. Pleistozáane und Holozáne Mo-

lluskenfaunen aus Stillfried an der March: Ein

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Stillfried und des Buhubergs nordlich von

Stillfried. Wissenschaftlichen. Mitteilung Nie-

derosterreichische Landesmus, 7: 7-272.

FRANK, C., 1992a. Spát-und postglaciale Gas-

tropoden aus dem Nixloch bei Losenstein-

Ternberg (Oberósterreich). Verbindung Os-

terreichisch Akademie der Wissenschaften, Mit-

teilung Kommission Quartarforschung, 8: 35-69.

FRANK, C., 1992b. Malakologisches aus dem

Ostalpenraum. Linzer biologische Beitrag, 24:

383-662.

zones (Vallonía costata biozone 18; Clausi-

liidae biozone 19; Granaría frumentum bio-

zone 18; Helecigona faustina-Acicula polita

biozone 22). Species of the Siberian-Asia-

tic faunal-centre follow them in impor-

tance (Vallonia costata biozone 15; Clausi-

Itidae biozone 19; Granaria frumentum bio-

zone 16; Helicigona faustina-Acicula polita

biozone 16). The Middle-European-

Mountain faunal-centres are located in

third place considering the relative

abundance of the species (Vallonia costata

biozone 8; Clausiliidae biozone 6; Granaria

frumentum biozone 6; Helicigona faustina-

Acicula polita biozone 9).

FUKOH, L., 1983. A búkki holocén Molluscák

állatfóldrajzi csoportositása (The Zoogeo-

graphical Grouppirung of the Holocene Mo-

lluscs in the Búkk). Malakologiai Tájekoztató

(Malacological Newsletter), 3: 37-39.

FUKOH, L., 1989. A Szilvasvárad: Szalajka-vólgy

(BNP) mésztufa úledekeinek malakosztrati-

gráfiai vizsgálata. Folia Historico-Naturalia

Musei Matraensis, 14: 39-42.

FUKOH, L., 1991. Examinations on Faunal-his-

tory of the Hungarian Holocene Mollusc

fauna (Characterization of the Succession

Phase). Folia Historico-Naturalia Musei Ma-

traensis, 16: 13-28.

FUKÓH, L., 1992a. Holocene Malacology in Hun-

gary. In Giusti, F. and Manganelli, G. (Eds.):

Abstracts of the eleventh International Malaco-

logical Congress. University of Siena: 115.

FuKOH, L., 1992b. The Holocene Mollusc fauna

of the Búkk Mountains. Abstracta Botanica,

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FUKOH, L., 1993a. Holocene malacofaunal as-

semblages in Hungary. Scripta Geologica.,

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FUKOH, L., 1993b. Main features of the deve-

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Newsletter), 12: 15-19.

FUKOÓH, L., KROLOPP, E. AND SUMEGI, P., 1995.

Quaternary malacostratigraphy in Hungary.

Malacological Newsletter. Suppl., 1: 113-198.

KERNEY, M. P., CAMERON, R. A. D. AND JUNG-

BLUTH, J. H., 1983. Die Landschnecken Nord-und

Mitteleuropas. P. P. Verlag Berlin: 63-284.

KORNIG, G., 1983. Studie zur Gastropodenfauna

der Westkarpatischen Kalk-und Karstsch-

luchten. Malakologische Abbhandlungen, 8: 131-

142.

O Sociedad Española de Malacología ——_—_—_—T— lIberus, 15 (2): 75-82, 1997

Aestivation responses of three populations of the giant Afri-

can snail, Achatina achatina Linne (Gastropoda: Achatinidae)

Respuestas a la estivación de tres poblaciones del caracol gigante afri-

cano Achatina achatina Linne (Gastropoda: Achatinidae)

Joseph R. COBBINAH*

Recibido el 9-X-1995. Aceptado el 7-1V-1997

ABSTRACT

The rates of aestivation and emergence from aestivation of three experimental populations

of the giant African snails Achatina achatina (Linne, 1758) were compared. The snails were

from three origins: Donyina in the Ashanti Region, Nkasem in the Brong Ahafo Region and

Apedwa in the Eastern Region of Ghana. The shortest aestivation period [4 weeks) was re-

corded for the Apedwa snails while the longest period (16 weeks) was recorded for the Don-

yina population. Data for pre-aestivation and post-aestivation growth rates show a decrea-

sing order: Apedwa > Nkasem > Donyina. The mean growth rates eight weeks before aesti-

vation were 3.6 g, 14.3 g and 19.2 g for the Donyina, Nkasem and Apedwa snails

respectively and differed significantly (P = 0.001). The variability in growth rates and dura-

tion of aestivation reflects the optimal sizes of the natural population of the three groups.

RESUMEN

Se comparan las tasas de estivación y de abandono de la misma de tres poblaciones ex-

perimentales del caracol gigante africano Achatina achatina (Linné, 1758). Los caracoles

eran de tres localidades diferentes: Donyina, en la región de Ashanti; Nkasem, en la re-

gión de Brong Ahafo, y Apedwa, al Este de Ghana. El periodo de estivación más corto (4

semanas) se registró en la población de Apedwa, mientras que el periodo más largo (16

semanas) se registró en la población de Donyina. Los datos de las tasas de crecimiento en

la pre-estivación y la post-estivación muestran un orden decreciente: Apedwa > Nkasem >

Donyina. Las tasas medias de crecimiento ocho semanas antes de la estivación eran 3,6

9, 14,3 g y 19,2 g para los caracoles de Donyina, Nkasem y Apedwa, respectivamente,

y diferían significativamente (P = 0,001). La variabilidad en las tasas de crecimiento y la

duración de la estivación reflejan los tamaños óptimos de las poblaciones naturales de los

tres grupos.

Key words: The giant African Snail, Achatina achatina, aestivation, Ecotypes.

Palabras clave: Caracol gigante africano, Achatina achatina, estivación, ecotipos.

INTRODUCTION

Achatina achatina (Linne, 1758) is which is maximal in the rainy season

found in the closed forest area in and minimal in the dry season. The

Ghana. It shows an annual activity snail burrows into the upper 10-15 cm

* Forestry Research Institute of Ghana. UST. Box 63, Kumasi, Ghana.

Iberus, 15 (2), 1997

COTE

d' IVOIRE

e Nkasem

_-“eKumasi

_- eDonyina

xx Moist Semi-deciduous

GULF OF GUINEA

Figure 1. Map of the closed forest area of Ghana showing the trial site (Kumasi) and the origins

(Donyina, Nkasem and Apedwa) of the snails used in trials.

Figura 1. Mapa del área de bosque cerrado en Ghana mostrando el lugar del experimento (Kumasz) y los

orígenes de los caracoles usados en el mismo (Donyina, Nkasem y Apedwa).

of soil during the dry season and

remains dormant for a period ranging

from three to five months (COBBINAH,

1993). This state of dormancy during

the dry season is referred to as aestiva-

tion. Circannual rythms are known to

be induced by such factors as light,

temperature, humidity and soil water

deficit (OwEnN, 1966). In the period

leading to the onset of aestivation, there

is a progressive decline in the snail's

metabolism. BARATOU (1988) asserts

that in order to maintain an equilibrium

between water in its tissue and the rela-

tive humidity of the immediate envi-

ronment, snails allow themselves to

dehydrate during this period. For

Achachatina marginata dehydration leads

to the loss of about 42% liveweight of

the non-shell tissues (STIEVENART, 1994).

Later, a mucous layer is secreted to

cover the shell opening; the fully

formed mucous layer is impervious to

both gases and water (HODASI, 1982).

Snails go into a state of dormancy whe-

, never conditions are too dry for their

liking. Whilst this behaviour is most

common in the dry seasons, COBBINAH

(1993) reported that even dry spells

during the wet seasons may induce

aestivation in A. achatina.

This is, however, in contrast to the

observation of HODASI (1982) that exten-

sive and persistent dry conditions are

necessary to induce aestivation, and that

aestivation normally does not occur

during the dry spells in the rainy

season. Extremes of temperature and

starvation is also reported to induce

aestivation (KONDO, 1964).

HODAsI (1982) suggested that not

every individual in the population aesti-

vates during the dry season. Accor-

dingly in certain localities within the

distribution range of A. achatina in

Ghana, fresh snails can be obtained

throughout the dry season. Here the

results of studies undertaken to deter-

mine the variability in aestivation res-

ponses of populations from three dis-

tinct enclaves of A. achatina in Ghana is

reported.

COBBINAH: Aestivation responses of the giant African snail, Achatina achatina Linne

MATERIALS AND METHODS

Sources of Snails: The snails for the

study were obtained from (a) Donyina in

the Ashanti Region, (b) Nkasem in the

Brong Ahafo Region and (c) Apedwa in

the Eastern Region. Donyina is 20 km from

our test site on the campus of the Univer-

sity of Science and Technology, Kumasi,

Ghana. Nkasem to the north west and

Apedwa to the south east of Donyina are

109 and 198 km respectively from our test

site. Figure 1 shows the origin of the th-

ree populations together with approxi-

mate isohyets. Donyina (6* 45' N and 2925'

W”, Nkasem (6* 15” N and 2* 20' W) and

Apedwa (6” 46" N and 1? 25 E) all fall wit-

hin the rainfall regime 1250 mm-1750 mm

typical of the moist semi-deciduous forest

type. Seasonal rainfall at the three origins

is influenced by meteorological Equator

(ME). Two weather systems are associa-

ted with the ME, the Intertropical Front

and the Intertropical Convergence Zone

which cause short heavy rain storms and

abundant continuous rain respectively and

result in bimodal annual rainfall pattern

with major rains falling between April-

June and minor period of rains between

September and October separated by two

months of less frequent rains (LEROUxX,

1988). The mean annual rainfall for the th-

ree locations are Donyina (1403 mm), Nka-

sem (1395 mm) and Apedwa (1561 mm).

The main dry season falls between De-

cember and March. The soils at the three

areas are of the forest ochrosols type. While

all three areas fall within one forest vege-

tation type, different levels of logging and

agricultural practices have resulted in var-

ying rates of deforestation. The Donyina

site has completely been converted into

farmland over the years. The Apedwa site

falls within the mountainous Atewa range

protection forest reserve where timber ex-

ploitation is prohibited. However, pockets

of illegal farms are not uncommon. The

Nkesem site is an off reserve area with

fairly dense vegetation interspersed with

farmlands.

Growth Rates: The snails were con-

ditioned in our research plot for four we-

eks on a diet consisting of pawpaw (Ca-

rica papaya) leaves and fruits, cocoyam

(Xanthosoma mofafa) leaves, and leaves of

the fameflower plant (Talinum triangulare).

Individuals that showed signs of inacti-

vity during this period were not used for

the trials. Twenty snails of each group

(ecotype) were placed in wooden boxes

measuring 0.6 x 0.6 x 0.35 m, filled to a

depth of 20 cm with sieved sterile silty

sandy soil obtained from an abandoned

rubbish dump. The snails in the boxes were

offered excess amounts of food but left

over foods were removed daily and soils

overturned weekly. Because of high sur-

vival rates of A. achatina (COBBINAH AND

OsEI-NKRUMAH, 1988) and insignificant

changes in shell size over short periods of

time, growth rates were measured by chan-

ges in live weights of snails. Each treat-

ment was replicated three times, with data

recorded for eight weeks prior to onset of

aestivation and eight weeks after emer-

gence from aestivation.

Aestivation patterns of the three

populations: The aestivation patterns of

individual snails used in the growth

studies described above were monito-

red. Snails were considered as having

aestivated when they covered the shell

opening with a white mucous layer.

Snails were recorded as emerged from

aestivation when they discarded the

epiphragm and resumed feeding.

A second trial was conducted to deter-

mine the effects of increased humidity.

Each group of snails was divided into two

lots. Twenty snails from one lot were

placed in a 0.6 x 0.6 x 0.3 m wooden box

as described above. A second set of 20

snails of the same ecotype was placed in

another box with humidity slightly incre-

ased by making a platform about 15 cm

above the soil level and placing a moiste-

ned fibre bag on the platform. The bag

was kept moist for two weeks before onset

of aestivation and throughout the aesti-

vation period. Similar sets were set up for

the other ecotypes and each treatment was

replicated three times.

All snails used in the studies were

labelled (paint marked) to enable obser-

vation of individual activities daily.

Aestivation responses (time of onset and

DÍ

Iberus, 15 (2), 1997

Table I. Growth rates of three populations of Achatina achatina cight weeks before and after aesti-

vation period.

Tabla I. Tasas de crecimiento de las tres poblaciones de Achatina achatina ocho semanas antes y después

del periodo de estivación.

Mean initial Mean weight gained Mean weight gained

weight (g) in 8 weeks before in 8 weeks after

Ecotype + s.e. aestivation + s.e. aestivation + s.e.

Donyina 46.6+1.25 3.60+0.58% 10.83 + 0.172

Nkasem 49.7+0.95 14.27+0.81? 12.47+ 0.359

Apedwa SS 19.20+1.37< 19.58 + 2.87*

Means within a row followed by the same letter are not significantly different (P= 0.05).

emergence from aestivation) were recor-

ded for all snails. The time required for

50% of each group to aestivate (TEsso) or

emerge from aestivation (TEms0o) was

estimated for the various treatments and

populations.

RESULTS AND DISCUSSION

Growth rates of the three popula-

tions: The pre-aestivation growth rates

for the Nkasem and Apedwa snails

were four and five times more than that

recorded for the Donyina snails. The

mean growth rates 8 weeks before aesti-

vation ranged from 3.6 g for Donyina to

19.2 g for Apedwa (Table 1).

Analyses of the data (ANOVA) in

Table 1 indicate that the growth rates

among the three populations differed sig-

nificantly for the pre-aestivation (F = 83.94;

df =2, 6; P< 0.001) and post-aestivation (F

= 7.73, df 2, 6; P< 0.02) periods. However,

Fisher's Multiple Range Test (LSD) did

not show significant difference in the post-

aestivation growth rates between the

Donyina and Nkasem snails. The very low

growth rates recorded for the Donyina

group 8 weeks before aestivation suggest

that, perhaps, aestivation in this group is

preceded by significantly longer period

of inactivity. Both the pre and post aesti-

vation growth rates of the three popula-

tions indicate that the Apedwa group

might be the most desirable group for

commercial snail farming.

Aestivation Patterns of the three

Populations: Figure 2A shows the aesti-

vation pattern of the entire snail popula-

tion. It took 10 weeks for all the snails in

the test to aestivate. Three peaks are

found in the second, sixth and tenth

week. The three peaks show the hetero-

geneity in the response of the entire

population to factors inducing aestiva-

tion.

Figure 2B shows the aestivation pat-

tern of the Donyina ecotype. Aestivation

commenced on 29 October 1993 and pea-

ked on 6th November, 1993, one week

after the beginning of aestivation. The

entire Donyina group aestivated in 5 we-

eks. The first observation of aestivation

for the Nkasem (Fig. 2C) and Apedwa

(Fig. 2D) groups was recorded on 6th

November, 1993 but peak aestivation for

these two groups was recorded on 4th

December, 1993 and 28th December,

1993, respectively. Aestivation for these

two groups spanned a period of 8 and 9

weeks. Time taken for 50% of each pro-

pulation to aestivate (TEss0) were 8, 28

and 52 days for Donyina, Nkasem and

Apedwa snails respectively.

The relatively longer periods required

for the Nkasem and Apedwa populations

to complete aestivation is a reflection of

the variability within these two popula-

tions. The peaks observed in Figures 2B-

D corresponded to the peaks in Figure 2A

and suggest that the heterogeneity in the

aestivation pattern of A. achatina observed

in our study was due mainly to the varia-

COBBINAH: Aestivation responses of the giant African snail, Achatina achatina Linne

ES 3

Percent aestivating

22)r0fa3 Sin wn 3h "h2 ap

Aestivation date

Percent aestivating

220 Sm am 32 M2 31h2

Aestivation date

Percent aestivating .

4, Dec —>

[27]

Percent aestivating

2/h0 sum 14n 3/2

Aestivation date

1712 31/12

50 28 Dec.—> D

2/0 sn. 19m 32 vh2 3112

Aestivation date

Figure 2. Aestivation patterns. A: entire experimental population; B: Donyina population; C:

Nkasem population; D: Apedwa population.

Figura 2. Patrones de estivación. A: toda la población experimental; B: población de Donyina; C: pobla-

ción de Nkasem; D: población de Apedwa.

tion in the responses of the three groups

to the factors inducing aestivation.

The emergence period for the entire

population covered a period of 7 weeks

from the end of January to mid-March,

1994. Again three peaks were evident (Fig.

3A). These were in the third, fifth and

seventh week and corresponded to peak

emergence periods for the Apedwa (Fig.

3D), Nkasem (Fig. 3C) and Donyina (Fig.

3B) respectively. The first snail to emerge

from aestivation was from the Apedwa

group on 31 January 1994 (Fig. 3D). By

mid-February 60% of this group had

emerged from aestivation. On the other

hand, not a single snail from the Donyina

group had emerged by mid-February,

three and half months after initiation of

aestivation (Fig. 3B). Twenty percent of

the Nkasem group had resumed normal

metabolic activities by mid-February (Fig.

3C). Peak emergence in the Donyina

group was recorded during the first week

in March. Time taken for 50% of popula-

tion to emerge from aestivation (T'Emso)

following the outset of emergence were 9,

Iberus, 15 (2), 1997

% Emergence from aestivation

6/1J94 30) az 2/2 vu 21

Emergence date

40 6Mar. —»

% Emergence from aestivation

6h 30h 13% 21 ad 7

Emergence date

40 20 Feb.

E 30

5 20

[|]

(5)

6h 30h 132 2/2 133 27/3

Emergence date

40F 9Feb. — D

% Emergence from aestivation

16h 30h 19k 27h

Emergence date

2/3 271

Figure 3. Emergence patterns. Á: entire experimental population; B: Donyina population; C:

Nkasem population; D: Apedwa population.

Figure 3. Patrones de salida de la estivación. A: toda la población experimental; B: población de

Donyina; C: población de Nkasem; D: población de Apedwa.

20 and 33 days for Apedwa, Nkasem and

Donyina groups respectively.

Humidity is considered a major factor

affecting aestivation behaviour of A acha-

tina. COBBINAH (1993) reported that when

the relative humidity falls during the dry

season A. achatina becomes inactive, seals

itself in its shell with a white calcareous

layer and aestivates in order to prevent

loss of water from the body. In this study

the enhanced humidity (3% above the

ambient condition) attained in the boxes

with moistened fibre bags did not

influence the overall aestivation pattern

of any of the three groups (see Table II).

Elmslie (pers. comm.) asserts that

aestivation / hibernation may be influen-

ced by a programmable regulation, reset-

table by environmental experience like cir-

cadian rhythm, but transmitted to offs-

pring in the case of parents that have

changed environment in a partially reset

state. It is possible that the enhanced

humidity in these boxes was not adequate

to destabilise the in-built mechanism

which sets in motion physiological

COBBINAH: Aestivation responses of the giant African snail, Achatina achatina Linne

Table II. Aestivation patterns of snails in boxes with or without moistened fibre bags.

Tabla II. Patrones de estivación de los caracoles en cajas con o sin bolsas de fibra humedecidas.

Mean Weekly % Aestivation Mean weekly % Emergence

Ecotype Dry (68-70% rh) Moist (68-74% rh) Dry (57-62% hr) Moist (56-66% rh)

Donyina 22.16 SEA 26.54 25.49

Nkasem 19.42 157 15772 16.52

Apedwa 2096 23.91 13.97 14.80

All differences are not significant at (P = 0.05).

changes resulting in aestivation during

periods of low atmospheric humidity.

Whilst aestivation has adaptive value

for the snail (HODASI, 1982; STIEVENART,

1994), for the snail farmer it represents the

loss of valuable growing time. The three

populations show significant differences

in growth rates and duration of aestiva-

tion. Shorter aestivation period mean

longer feeding period and ultimately

larger body sizes.

Based on peak aestivation and emer-

gence periods for the three groups, the

estimated duration of the dormant periods

were 4, 10 and 16 weeks for the Apedwa,

Nkasem and Donyina respectively. More-

over, data on growth rates clearly show a

decreasing order Apedwa > Nkasem >

Donyina among the three groups during

the pre and post aestivation periods. These

two factors acting in concert may explain

differences in adult sizes of A. achatina

from various areas of the country. The

Apedwa snails are usually twice the size

of the Donyina snails (COBBINAH, 1993).

The Nkasem snails are often intermediate

in size between the two populations.

Although all the three enclaves

where the snails originated from are

within the moist semi-deciduous forest

type and are characterized by similar soil

type, there are differences in mean

annual rainfall and vegetation cover. Soil

water regime are influenced by rainfall

gradient and evapotranspiration (VAN

ROMPAEY, 1993). Most studies of the soil

water regime in West African tropical

forest (HUTTEL, 1975; COLLINET, MONTE-

NEY AND POUYAUD, 1984) suggest that

seasonal soil water deficits increase with

decreasing annual rainfall. Mean annual

rainfall is highest at Apedwa (1561 mm)

but similar at Donyina (1403 mm) and

Nkasem (1359 mm). In the three enclaves

the Donyina area is the most degraded

due to logging and slash and burn agri-

culture practices over the years. Unlike

Apedwa and Nkasem where snails are

mainly gathered from forest reserves

and secondary forests outside reserves,

the Donyina snails are mainly gathered

from low vegetation farmlands. Alt-

hough Donyina and Nkasem have

similar mean annual rainfall, the relati-

vely poor vegetation cover at Donyina

would result in higher evapotranspira-

tion and longer duration of seasonal

drought. The snails from this area have

probably adapted to this relatively long

drought period through extension of

dormancy period.

All individuals in the three groups

aestivated in these studies. Neverthe-

less, a few individuals among the

Apedwa group had shorter periods of

aestivation than the four week group

average. Further studies are, however,

underway to determine whether some

individuals or groups normally remain

active throughout the dry season, and

also to better understand the physiologi-

cal, environmental and behavioural fac-

tors controlling aestivation. If the varia-

bility in the aestivation behaviour by the

different individuals or groups has a

significant genetic component, the resul-

ting information would be of potential

use for commercial snail farming.

Iberus, 15 (2), 1997

REFERENCES

BARATOU, J., 1988. Raising snails for food. llu-

minations Press, Calistoga, USA, 72 pp.

COBBINAH, J. R. AND OSE-NKRUMAH, A., 1988.

The effect of food on growth of Achatina acha-

tina. Snail Farming Research, 2: 20-24.

COBBINAH, J. R., 1993. Snail farming in West

Africa: a practical guide. Sayce Publishing Ltd.,

Exeter, U. K., 49pp.

COLLINET, J., MONTENEY, B. AND POUYAUD, B.,

1984. Le millieu physique. Recherche de amena-

gement en millieu forestier tropical humide: le pro-

jet taide Cote d'Ivoire. Notes Techniques du

MAB, 15.

HODASL J. K. M., 1982. Some aspects of the bio-

logy of Achatina (Achatina) achatina (Linne).

Bulletin de l'IFAN, 44: 100-114.

HUTTEL, CH., 1975. Recherches sur l'ecosys-

teme de la foret subequatoriale de basse Cote

d'Ivoire. IV. Estimation du Bilan hydrique.

La Terre et la Vie, 29: 192-202.

KONDO, L., 1964. Growth rates in Achatina fu-

lica Bowdich. Nautilus, 78: 6-15.

LEROUX, M., 1988. La variabilite des precipita-

tions en Afrique occidentale. Les compo-

santes aerologigues du probleme. Veille cli-

matique satellitaire, 22: 26-46.

OWEN, D. F., 1966. Animal Ecology in Tropical

Africa (1st Ed.). Oliver and Boyd, London, 122

STIEVENART, C., 1994. Artificial estivation of the

giant African snail Archachatina marginata sa-

turalis: Survival, weight loss and meat yield.

Snail Farming Research, 5: 23-28.

VAN ROMPAEY, R.S. A.R., 1993. Forest gradients

in West Africa. Ph. D. Thesis, Agricultural

University, Wagemningen, 98 pp.

O Sociedad Española de Malacología —————— IBERUS, 15 (2): 83-93, 1997

Snail communities associated to swampy meadows and

sedgy marshy meadows plant communities of the Great

Hungarian Plain

Comunidades de moluscos asociadas a comunidades vegetales de pra-

deras pantanosas y junqueras en la Gran Llanura Húngara

Károly BÁBA* and István BAGI'**

Recibido el 11-X-1995. Aceptado el 29-1V-1997

ABSTRACT

Simultaneous phytocoenological, malacological and pedological studies were carried out

in six successional plant community types characteristic on the Great Hungarian Plain.

Data were analyzed by multivariate statistical methods (PCA). Variation in the abundance

of ecological, habitat type and nutritional type species groups was also followed. In the

Succiso-Molinietum (swampy meadows) and Agrostio-Caricetum [sedgy marshy meadows)

plant communities, the distribution of constant and differential species is mostly influenced

by their range of pH tolerance. Habitat drying and salinization, and various human

impacts (draining, cutting and grazing by domestic animals) influence the succession of

vegetation. Changes in snail assemblages include altering proportion of living and dead

individuals and decreasing diversity (H'), both reflecting habitat drying and salinization.

Complementary changes in the abundance of riparian and steppe dweller species groups

indicate habitat drying, while swamp dwellers become more numerous as the topsoil

becomes muddy due to salt accumulation. Concerning nutritional types, the proportion of

omnivores decreases with habitat drying, whereas the frequency of herbivores increases in

a complementary manner. The increasing abundance of saprophagous snails reflects bio-

tope eutrophization caused by cutting and grazing.

RESUMEN

Se han desarrollado simultaneamente estudios fitocoenológicos, malacológicos y pedoló-

gicos en seis tipos de comunidades vegetales de la Gran Llanura Húngara. Los datos fue-

ron analizados mediante métodos estadísticos multivariantes. También se ha estudiado la

variación en los grupos de especies desde el punto de vista ecológico, de su hábitat y

tipo nutricional. En las comunidades vegetales Succiso-Molinietum (praderas pantanosas)

y Agrostio-Caricetum (¡junqueras), la distribución de las llamadas especies constantes y

diferenciales está mayoritariamente influenciada por su rango de tolerancia de pH. La

desecación del hábitat y su salinización, junto con un conjunto de alteraciones humanas

(desecación, segado y ramoneado por animales domésticos) afectan la suceción vegetal.

Los cambios en las comunidades de moluscos incluyen la variación en las proporciones

de individuos vivos y muertos y una menor diversidad (H'); ambos cambios reflejan la

desecación y salinización del hábitat. Cambios complementarios en la abundancia de los

grupos de especies ribereñas y de estepa son indicadores de la desecación, mientras que

las especies propias de zonas pantanosas se hacen más abundantes según la capa super-

* Department of Biology, Gy. Juhász Teacher Training College, Szeged, Hungary.

** Department of Botany, A. József University, Szeged, Hungary.

IBERUS IS NZ)IIH 7

ficial se vuelve fangosa debido a la acumulación de sal. Por lo que se refiere a los grupos

nutricionales, la proporción de omnivoros decrece con la desecación, mientras que la fre-

cuencia de herbivoros crece de manera complementaria. La creciente abundancia de

especies saprófagas refleja la eutrofización producida por el segado y ramoneo.

KEY WORDS: Gastropoda, drainage, salinization, species groups, succession, Hungary.

PALABRAS CLAVE: Gastropoda, desecación, salinización, grupos de especies, sucesión, Hungría

INTRODUCTION

Under the semiarid climate characte-

ristic to the Eupannonicum floristic

region (which the lowlands of the Car-

pathian Basin belong to), habitat drying

and salinization processes were studied

along a successional series spanning

from swampy to salt-affected meadows.

The Succiso-Molinietum association

represents the wet meadows on calcare-

ous swampy meadows soils. The phy-

siognomy of its vegetation is determi-

ned by tall grasses (Molinia caerulea, M.

arundinacea, Festuca pratensis, Deschamp-

sia caespitosa, Agrostis stolonifera). Its

stands have high species diversity and

are rich in protected rare species (Dacty-

lorrhiza incarnata, Orchis laxiflora

palustris, Cruciata pedemontana, Veratrum

album, Iris spuria, 1. sibirica, ...). The

Succiso-Molinietum is in successional

relation to the Agrostio-Caricetum asso-

ciation; namely, moderate habitat drying

and salinization involve its transforma-

tion into the latter. The Agrostio-Carice-

tum may occur independently, too. It is

characteristic of the solontschak-solonet-

zic alkaline meadows soils. Depending,

on the hydro- and haloecological condi-

tions the Agrostio-Caricetum forms

various vegetation units (eg. subassocia-

tions), that well reflect the environmen-

tal impacts. These units are extremely

diverse in its species composition and

their physiognomy. The two associa-

tions represent the meadow formation

of the Great Hungarian Plain in a signi-

ficant percentage. As the two associa-

tions have evolved under wet condi-

tions, the drought on their habitats

causes drastical transformation in their

vegetation structure, first of all in their

species composition. The changes in the

structure show close relationship to the

environmental conditions, therefore

every distinguished vegetation unit well

reflects a stage of the drought induced

vegetation transformation processes

(phytocoenological indication). A lower

proportion of data regards to other plant

associations, that are in successional

relationships with Succiso-Molinietum

and Agrostio-Caricetum.

The water management works in the

20 century and the more and more arid

climate of the last two decads endange-

red the vegetation of the wet meadows.

Their transformation into drier habitats

would have harmful consequences for

the whole ecosystem, particularly the

animal assemblages, that have no

enough mobility to change their habitat.

The terrestrial snails belong to a little

mobile group of animals, they are

bound to their habitats more firmly than

other ones; therefore the, however well-

known, successional and zonational re-

lationships of the vegetation units can

be examined on the basis of their snail

assemblages, too. If the transformation

processes of vegetation and their animal

assemblages show paralelism, the chan-

ges in the composition of this animal

group would have an indicative value

for nature conservation.

MATERIALS AND METHODS

In the South-eastern part of the

Great Hungarian Plain (Csongrád

County) snail assemblages were sam-

pled by the quadrat method. Ten plots

25 x 25 cm size were examined in para-

llel with phytosociological recording of

BÁBA AND BAGI: Snail communities in swampy and sedgy marshy meadows in Hungary

each plant subassociation encountered

(S0ó, 1964). A hight proportion of the

data regards to two plant communities:

Succiso-Molinietum and Agrostio-Carice-

tum; only some samples originated ot-

her successionally related association.

Altogether 30 collection sites were visi-

ted in six plant community types, while

the number of subassociations studied

was 22 (see Figure 3). In each quadrat, a

detailed soil analysis was conducted, in-

cluding measurements of relative per-

centage soil moisture, total organic mat-

ter content, CaCOz3 concentration, hy-

groscopy and pH.

The concepts applied in the coenolo-

gical characterization are the following;

Abundance (A) is the number of indivi-

duals of a snail species found in a plant

community regarding to one m? (A /m?);

Dominance (D) is the ratio of indivi-

duals of a species related to the total

individuals of every species; Species

density (SD) is the average species

number of 10 quadrats in a collection

site; Frequency (F) is the ratio of a

species in relation to the total number of

species in a collection site (consist of 10

quadrats); Constancy (K) is the ratio of a

species in relation to the number of all

species found in all collection sites

belonging to the same plant community.

When a species was found in all the

quadrats, it can be considered as an

absolute constant species. D, F and K are

expressed as percentages.

Data were analyzed by standardized

Principal Components Analysis (PCA,

PODANI, 1988). Shannon-diversity (H'),

SD and changes in the abundance

(A/m?) of living and dead individuals

were followed through examination of

the proportions of various species

groups. Ecological species groups were

defined as follows: S: sciophilous, P:

swamp dweller, Ph: photophilous, R:

riparian and OA: species of open areas.

They were obtained by applying the

block cluster method of FEOLI AND

ORLÓCZI (1979). A simplified version of

LoZEK'"s (1964) typology was used and

the following habitat type groups were

distinguished: riparian ubiquitous (RU),

bush forest dweller (B), hygrophilous

swamp dweller (HP) and steppe dweller

(ST). Nutritional type groups (O: omni-

vore, SP: saprophagous, H: herbivore)

were differentiated after the system of

FRÓMMING (1954). Species and their

group assignments are listed in Table 1.

RESULTS AND DISCUSSION

Species encountered: Field studies

yielded a collection of 3047 living and

3150 dead individuals belonging to 26

species (Table 1). The majority of the

specimens was found in the Succiso-

Molinietum (1062 + 1268) and Agrostio-

Caricetum (1496 + 1445) phytocoenoses,

while plant associations 1, 3, 4 and 6

altogether contained 489 + 440 alive and

dead individuals, respectively.

Two species new to the southern

part of the Great Hungarian Plain were

detected: Malacolimax tenellus (O. F.

Muller, 1774) and Deroceras sturanyi

(Simroth, 1894).

Characteristic species and their re-

quirements: On the basis of frequencies

of occurrence data, the constant, sub-

constant and accessorial species could be

determined for the two plant associa-

tions most rich in snails (Table ID). Cons-

tant and subconstant species reach low

levels of dominance in both communi-

ties. This is probably due to unfavoura-

ble changes in their environment caused

by either draining, drying, salinization

or grazing. Differential species are Coch-

licopa lubricella (Porro, 1938) and Cary-

chium minimum O. FE. Miller, 1774 in the

Succisio-Molinietum plant association,

and Pupilla muscorum (L., 1758) in the

Agrostio-Caricetum. The occurrence of

Truncatellina, Granaria and Helicopsis spe-

cies in the Agrostio-Caricetum association

indicates habitat drying. The distribution

of constant and differential species is

strongly influenced by the width of their

pH tolerance range (Figs. 1, 2), as it has

been shown earlier (BABA AND DOMON-

KOs, 1992). According to aur data diffe-

rential species of the Succisio-Molinietum

association have a narrow pH tolerance

range, in contrast with species occurring

IBERUS, 15 (2), 1997

Table I. Gastropod species found in the plant communities studied (1: Caricetum acutiformis-ripa-

riae, 2: Succiso-Molinietum; 3: Bolboschoenetum maritimae, 4: Astero-Agrostietum;, 5: Agrostio-

Caricetum distantis, 6: Achilleo-Festucetum pseudovinae) E: Ecological species groups (S: sciophilous;

P: swamp dweller; Ph: photophilous; R: riparian; OA: species of open areas); N: Nutritional type

(O: omnivore; SP: saprophagous; H: herbivore); H: Habitat type (RU: riparian ubiquitous; B: bush

forest dweller; HP: hygrophilous swamp dweller; ST: steppe dweller).

Tabla I. Especies de gasterópodos encontradas en las comunidades vegetales estudiadas (1: Caricetum acu-

tiformis-ripariae; 2: Succiso-Molinietum, 3: Bolboschoenetum maritimae; 4: Astero-Agrostietum;

5: Agrostio-Caricetum distantis; 6: Achilleo-Festucetum pseudovinae) E: Grupos ecológicos de las

especies (S: esciófilo; P: de marisma; Ph: fotófilo; R: ribereño; OA: de áreas abiertas); N: Tipo nutricio-

nal (O: omnívoro, SP: saprófago, H: herbívoro); H: Tipo de hábitat (RU: ribereño ubiquo; B: zonas

arbustivas; HP: marismeño higrófilo; ST: estepa).

E N H 1 2 3 4 5 6

SSP OR Carychium minimum (0. F. Miller 1774) 17 5

S SP HP Carychium tridentatum (Risso 1826) 8

0A HB Cepaea vindobonensis (Ferussac 1821) l 2 6

DA SP ST Chondrula tridens (0. E. Múller 1774) 135 l 388 50

Rs 0 Bb Cochlicopa lubrica (0. F. Múller 1774) 3 4

DA 0. ST Cochlicopa lubricella (Porro 1838) + 184

R 0 RU Deroceraslueve(0.F. Miller 1774) l

R 0. HP Derocerassturanyi(Simroth 1894) l

0A 0 B Euconulus fulvus (0. F. Miller 1774) 4

0A H ST Granaria frumentum (Draparnaud 1801) + 86

0A H ST Helicella obvia (Menke 1828) 3

MAS Helicopsis striata (0. E. Múller 1774) 1

Ph H B Helix pomatia (Linne 1758) 2

S 0 RU Malacolimax tenellus(0.P. Miller 1774) l

PH ST Monacha carthusiana (0. F. Miller 1774) 18 99 13 AS

R H RU Perforatella rubiginosa (A. Schmidt 1853) 3

0A H ST Pupilla muscorum (Linne 1758) l l 329 165

R 0 RU Succineaoblonga (Draparnaud 1801) 8 169 43 6 130 +

P 0 HP Succinea elegans (Risso 1826) 1

DA SP ST Irucatellina cylindrica (Ferrusac 1807) 3

0A H ST Vallonia costata (0. E Múller 1774) | 8 6

REO p Vallonia enniensis (Gredler 1856) 69 39 244

RS Vallonia pulchella (0. E. Múller 1774) 19 40 A

RES SRU Vertigo antivertigo (Draparnaud 1801) 1

A A | Vertigo pygmaea (Draparnaud 1801) 1 20 44 2

R 0 RU Zonitoides nitidus(0.F. Miller 1774) 12 1

Number of Individuals 151 1062 58 12 1496 268

Number of collection sites l 11 l l 15 l

Number of Species 11 17 3 3 15 7

Dead Individuals 155 1268 OD 42 1445 243

in both Agrostio-Caricetum and Succisio-

Molinietum, which tolerate a much wider

range of soil pH (Fig. 2).

Successional changes: The ordina-

tion of snail samples of collection sites

(1-30) clearly indicates (Fig. 3) the suc-

BÁBA AND BAGI: Snail communities in swampy and sedgy marshy meadows in Hungary

Table I1. The constant (above), subconstant (middle) and accesorial (below) species of the two plant

communities (Succiso-Molinietum, Agrostio-Caricetum) studied in more detail. K: constancy; D:

dominance.

Tabla 11. Las especies constantes (arriba), subconstantes (medio) y accesorias (abajo) en las dos comuni-

dades vegetales estudiadas (Succiso-Molinietum, Agrostio-Caricetum). K: constancia; D: dominancia.

Succiso - Molinietum K D

Monacha carthusiana 100 9.32

Succinea oblonga 90.9 15.91

Cochlicopa lubricella 81.81 17.32

Vallonia enniensis 1272 36.72

Chondrula tridens IDEA

Vertigo pygmaea 45.45 1.88

Carychium minimum 21.17 0.47

Vallonia pulchella 18.18 3.76

Cochlicopa lubrica 18.18 0.28

Carychium tridentatum 9.09 0.75

Perforatella rubiginosa 9.09 0.28

Cepaea vindobonensis 9.09 0.09

Deroceras laeve 9.09 0.09

Deroceras sturanyi 9.09 0.09

Malacolimax tenellus 9.09 0.09

Pupilla muscorum 9.09 0.09

Vertigo antivertigo 9.09 0.09

Agrostio - Caricetum K D

Chondrula tridens 100 25.93

Monacha carthusiana 100 13.03

Succinea oblonga 80 8.68

Pupilla muscorum 73.33 21.99

Vertigo pygmaea 33.33 2.94

Vallonia pulchella 26.66 3.67

Cepaea vindobonensis 20 0.4

Helix pomatia 13.13 0.13

Granaria frumentum 6.66 5.74

Vallonia costat 6.66 0.53

Cochlicopa lubricella 6.66 0.26

Iruncatellina cylindrica 6.66 0.2

Helicopsis striata 6.66 0.06

Zonitoides nitidus 6.66 0.06

cessional and zonational relationships of

the plant associations studied, and a

gradual habitat drying (Baci, 1988). The

main lines of the drying processes can

be outlined as follows: (A) Non-salinic,

wet line Caricetum acutiformis-ripariae (L,

1), Succiso-Molinietum typicum (IL, 2-10),

S. -M. poetosum (X, 11), S. -M. agrostieto-

sum (XI, 12). The latter is a connection to

a different, saline line: (B) Bolboschoene-

tum maritimae (IV, 14), Agrostio-Caricetum

bolboschoenetum (IV, 21), A. -C. fac. Juncus

(UL 16), Agrostio-Caricetum plantagineto-

sum maritimae (V, 17-20). Later, while the

drying procces continues, the two lines

originated a common line (C), whose re-

presentative associations are A-C. festu-

cetosum arundinacene (VI, 28), A. -C. poe-

tosum (Vlla, 22-24), A. -C. festucetosum

pseudovinae (VII, 15) and finally the

Achilleo-Festucetum pseudovinae (IX, 29).

Roman numbers indicates groups of co-

llection sites with similar features obtai-

ned from the analysis. The successio-

nally closely connected plant communi-

ties often form zonation systems in the

field. The drying processes have been

particularly accelerated since the sixties,

due to the draining of the region. The

three successionally related lines could

be distinguished in the process of drai-

ning-generated habitat drying by inves-

tigation of snails, too. The declining

density of dead shells and living indivi-

duals, the fall of individual density (ID)

and diversity (H”) indicate habitat dr-

ying (X, V, Vlla, b, IX and VIII) and

sometimes salinization (Fig. 4). Habitat

drying accelerates with draining, which

then leads to higher snail abundance

again at the end of the successional

xero-series at dry localities (Achilleo-Fes-

tucetum association). Chondrula and Pu-

pilla can become especially numerous.

Snail species groups were used to

evaluate these processes, for which the

IBERUS, 15 (2), 1997

2. axis

1.907 3

-2.043 5

Í | 1. axis

-2.038 2.23

Figure 1. Distribution of constant and subconstant species according to soil pH (standardized PCA).

1: Carychium minimum, 2: Succinea oblonga; 3: Cochlicopa lubricella; 4: Vertigo pygmaea; 5: Pupilla

muscorum; 6: Vallonia costata;, 7: Vallonia pulchella; 8: Vallonia enniensis, 9: Chondrula tridens.

Figura 1. Distribucion de las especies constantes y subconstantes de acuerdo con el pH del suelo (Análisis

de componentes principales estandarizado). 1: Carychium minimum, 2: Succinea oblonga; 3:

Cochlicopa lubricella; 4: Vertigo pygmaea; 5: Pupilla muscorum,; 6: Vallonia costata; 7: Vallonia

pulchella; 8: Vallonia enniensis; 9: Chondrula tridens.

9.2

No)

8.8

8.6

8.4

pH interval

(0,2)

159)

8 9 2 3) 4 3 7 6 l

Figure 2. Distribution of the pH tolerance ranges of constant and subconstant species. Numbers

refer to species as in Figure 1.

Figura 2. Distribución de los rangos de tolerancia al pH de las especies constantes y subconstantes. Los

números de las especies son idénticos a los de la Figura 1.

BÁBA AND BAGI: Snail communities in swampy and sedgy marshy meadows in Hungar;

2.axis

1.744

- 0.896

-1.329 1.258

Figure 3. Ordination of collection sites by using data of snails samples (standardized PCA). Roman

numbers indicate sample groups with similar phytological and malacological features; groups obtai-

nes by ordination of data. Arabic numerals in brackets indicate collection sites. Caricetum acutifor-

mis-ripariae 1 (1); Succiso-Molinietum a. typicum facies Veratrum album 1 (9), b. typicum facies

Phragmites 11 (10), c. typicum normale 1 (2-8), d. poetosum angustifoliae X (11), e. agrostietosum XI

(12); Bolboschoenetum maritimae YV (14); Astero - Agrostietum IV (30); Agrostio - Caricetum distan-

tis, a. agrostietosum IV (13), b. agrostietosum facies Juncus compressus YI (16), c. plantaginetosum

maritimae Va, b (17-20), d. poetosum angustifoliae Vlla (22-24), e. festucetosum arundinaceae VÍ

(28), £. festucetusum pseudovinae VII (15), g. bolboschoenetosum IV (20); Achilleo - Festucetum pseu-

dovinae IX (29). See the text for further details.

Figura 3. Ordenación de las estaciones de muestreo utilizando datos de muestras de moluscos (Análisis de

componentes principales estandarizado). Los números romanos indican grupos de estaciones con similares

características fitológicas y malacológicas; los grupos se obtuvieron por ordenación de los datos. Los núme-

ros arábigos entre paréntesis indican las localidades de muestreo. Caricetum acutiformis-ripariae / (1);

Succiso-Molinietum 4. typicum facies Veratrum album /7 (9), b. typicum facies Phragmites // (10),

c. typicum normale /7 (2-8), d. poetosum angustifoliae X (11), e. agrostietosum AX7 (12);

Bolboschoenetum maritimae /V (14); Astero - NIV (30); Agrostio - Caricetum distantis, 4. agros-

tietosum /V (13), b. agrostietosum facies Juncus N1I] (16), c. plantaginetosum maritimae Va, b (17-

20), d. poetosum angustifoliae V//a (22-24), e. festucetosum arundinaceae VI (28), f. festucetusum

pseudovinae VIT (15), g. bolboschoenetosum 7V (20); Achilleo - Festucetum pseudovinae /X (29).

Véase el texto para más detalles.

abundantial changes are shown (Figs. 5, quists (R, RU) monotonously decreases

6, 7). Habitat drying and salinization (collection sites 11 and 12 represent sta-

have different consequences in the two ges of ramification in the successional se-

plant associations. In the Succiso-Molinie- ries). As the wet terrain dries down gra-

tum one, the abundance of riparian ubi- dually, the abundance of species typical

IBERUS, 15 (2), 1997

1000

900

800

700

600

DA /m?

500

la)

/m?

VI Vila Vilb 1X vi

A B C

Figure 4. Variation in species density (1D), species diversity (H”), pH and density of living (A/m?)

and dead individuals (DA/m?) in groups of phytocoenologically similar samples (Roman numerals)

influenced by the dominant process of the habitat changes. A, B and C refer to the three succession

lines of the vegetation. A includes groups 1, II, X and XI; the main processes are drying (L, ll and

X) and salinization (XI). B includes groups IV, II! and V; the main processes are drying (IV and 111)

and an additional salinization with cutting (Va, 18-20) and degradation (Vb, 17). C includes groups

VI, VIL VII and IX; the main processes are drying (VI, Vlla 22-24 and IX) with cutting (VIIb 25-

27) and drainage (VII.

Figura 4. Variación en la densidad de las especies (ID), diversidad (H), pH y densidad de individuos

vivos (A/m*) y muertos (DA/m?) en los grupos de muestras con características fitocoenológicas similares

(números romanos) influenciadas por los procesos dominantes en los cambios de hábitat. A, B y C se

refieren a las tres líneas de sucesión de la vegetación. A incluye los grupos l, Il, X y XI; los principales pro-

cesos son la desecación (L, 1I y X) y la salinización (X1). B incluye los grupos IV; 11 y V: los principales

procesos son la desecación (IV y 11) y una salinización adicional con el segado (Va, 18-20) y degrada-

ción (Vb, 17). C incluye los grupos VI, VI, VII y LX; los procesos principales son la desecación (VI, Vlla

22-24 y IX) junto con el segado (VIIb 25-27) y el drenaje (VII).

in open areas (OA, ST) decreases, para-

lleled by a similar decline in the number

of species and individuals (Figs. 5, 6).

Concerning ecological species groups,

the abundance of swamp dwellers (P,

HP, Monacha) becomes higher with bio-

tope salinization (12). Na* accumulation

in the topsoil is responsible for its

muddy character.

Sciophilous (S) and bush forest dwe-

ller (B) snail species may also appear in

the tall and dense stands of Caricetum

acutiformis-ripariae community. The

abundance of OA and ST species groups

increases at the end of the successional

series in Agrostio-Caricetum and Achilleo-

Festucetum associations. Groups P and

HP are also more abundant there, due to

BÁBA AND BAGI: Snail communities in swampy and sedgy marshy meadows in Hung;

80 s F 600

20 A /

il L 500

- Ph j

60 le

==-==-=R 11

¡lp 400

===. OA 07 —

¡pl S

j! e

2 pa300 “

lle SE

J £

l. <

Ni l

N l F 200

iS 4

y e

Ñ US y

o S z Ni L 100

SUN N E

IN Ñ A 7%

IE ==

NEÁ E 0

T 1 T T e A pO

I u Xx XI Iv TI Va Vb VI Vila Vilb IX VI

Figure 5. Variation in the abundance (A/m?) of ecological snail species groups among plant asso-

ciations and subassociations. Abbreviations, S: sciophilous, P: swamp dweller, Ph: photophilous, R:

riparian and OA: species of open areas.

Figura 5. Variación en la abundancia (A/m?) de los grupos ecológicos de especies de moluscos en el con-

junto de las asociaciones y subasociaciones vegetales. Abreviaturas, S: esciófilo; P: de marisma; Ph: fotófi-

lo; R: ribereño; OA: de áreas abiertas.

120 RU L 700

le perio 0 coria

100 4 Í: ee E p E

: ' Pa En eS

DON IV IM Va Vb VI Vila Vb IX VIn

Figure 6. Variation in the abundance (A/m?) of snail habitat type groups among plant associations

and subassociations. Abbreviations, RU: riparian ubiquist, B: bush forest dweller, HP: hygrophilous

swamp dweller and ST: steppe dweller.

Figura 6. Variación en la abundandia (A/m”) de los tipos de hábitat en las especies de moluscos en el con-

junto de las asociaciones y subasociaciones vegetales. Abreviaturas, RU: ribereño ubiquo; B: zonas arbus-

tivas; HP: marismeño higrófilo; ST: estepa.

IBERUS, 15 (2), 1997

250

(0)

2 Sp

200 OSA

A 150

< 100

50 y

IU X Xi iv IM

: E 400

al 350

E 300

20)

; 250 po

200 au

150

100

Va Vb VI Vila Vilb Ix MIU

Figure 7. Variation in the abundance (A/m?) of snail nutritional type groups among plant associa-

tions and subassociations. Abbreviations, O: omnivore, SP: saprophagous, H: herbivore.

Figura 7. Variación en la abundancia (A/m?) de los grupos nutricionales de moluscos en el conjunto de

las asociaciones y subasociaciones vegetales. Abreviaturas, O: omnívoro, SP: saprófago, H: herbívoro.

the muddy topsoil formed under Na*

accumulation (Figs. 5, 6).

In both collection site groups of the

Agrostio-Caricetum association, habitat

salinization and drying, grazing and

cutting result in the decline of the pro-

portion of riparian ubiquists (R, RU:

Succinea, Vallonia pulchella), and a com-

plementary increase in the abundance of

steppe dwellers (OA, ST: Chondrula,

Pupilla). Stands of XL, Va and Vlla are

regularly cut, while site of VIII is both

cut and drained (Figs. 5, 6).

In terms of nutritional types, omni-

vorous snail species dominate in each

collection site group, as it was also

found elsewhere in willow-poplar fo-

rests (Bába, 1993). With habitat drying

the vegetation becomes denser, resulting

in a higher abundance of herbivores.

Subsequent cutting increases the pro-

portion of saprophagous species (Vallo-

nia, Chondrula, Vertigo). The omnivore

and herbivore-saprophagous groups

were found to change in a complemen-

tary manner (Fig. 7).

According to our data, the connec-

tion between the vegetation units and

the species groups of snails seems to be

very close. The composition of snail as-

semblages indicates the most important

environmental changes, such as drying

and salinization, and the human im-

pacts, eg. cutting, mowing and some ot-

her disturbances. The structural transfor-

mation of snail assemblages can be follo-

wed at a level of species groups and also

within these groups. The snail assembla-

ges indicate not only the differences bet-

ween plant communities but the diffe-

BABA AND BAGI: Snail communities in swampy and sedgy marshy meadows in Hungary

rent impacts of natural and anthropoge-

nic factors within a plant community as

well. The consequences of the habitat dr-

ying are the decrease in species number

and increase in abundance; the saliniza-

tion causes the decrease of species num-

ber and change of species groups in a

particular habitat. Mowing leads to the

decrease of species number and increase

of the ratio of saprophagous and steppe

dweller species groups. The changes can

be traced back to pedological reasons,

eg. habitat drying, increase in pH value,

REFERENCES

BABA, K. AND DOMOKOS, T., 1992. The occu-

rrence and ecology of Chilostoma banatica

(Rossmassler, 1838) in Hungary. Abstract of

the Eleventh International Malacological Con-

gress, Siena: 383-385.

BABA, K., 1993. Effect of the regions of the Tisza

Valley on the malacofauna. Tiscia, 18: 97-102.

Bacií, L, 1988. The role of water management

in the degradation processes of halophytic ve-

getation in Hungary. Environmental Conser-

vation, 15: 359-362.

FEOLI, E. AND ORLÓCZI L., 1979. Analysis of con-

centration and detection of underlying fac-

tors in structured tables. Vegetatio, 40: 49-54.

and accumulation of organic matter. The

investigations of the structural and com-

positional changes of snail assemblages

of the studied six plant communities

may provide a way to detect the conse-

quences of the salinization as a characte-

ristic successional process of Hungarian

Great Plain. The studies on the changes

of snail assemblages in meadow plant

communities can indicate the main pro-

cesses in this vegetatation type similar to

the investigations carried in forest

ecosystems (BABA, 1993).

FRÓMMING, E, 1954. Biologie der mitteleuropais-

chen Landgastropoden. Duncker-Huniblot, Ber-

lin 1-404.

LoZEk, V., 1964. Quartermollusken der Tschec-

hoslowakei. Tschechoslowakischen Akademie

der Wissenschaften. Praha, 374 pp.

PODANI, J., 1988. Syn-Tax III. Users Manual.

Abstracta Botanica, 12: 1-179.

S06, R., 1964. Synopsis systematico-geobotanica flo-

rae vegetationisque Hungariae I. Akadémiai

Kiadó, Budapest, 589 pp.

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De Ñ Fe 5

Caloabezá nt

VAT lee 20 0 a

lie

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pS de 1 o

$ dio 7

1999 ) M0

Ae. DUDA. al dhe E

e oso sabe e -

siniles 4

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bl Esos

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sembla vic

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Iberus, 15 (2)495-121, 1997

Morphological and biometrical researches on Austrian

Clausiliids. Shell morphology and variability in Clausilia

dubia Draparnaud, 1805

Investigaciones biométricas y morfológicas en Clausílidos de Austria.

Morfología y variabilidad de la concha de Clausilia dubia Drapar-

naud, 1805

Karl EDLINGER*

Recibido el 8-[-1996. Aceptado el 28-V-1997

ABSTRACT

Morphological and biometrical studies on shells of some Austrian populations of Clausilia

dubia Draparnaud, 1805, show a great variability in size and in the other morphological

features within the species as a whole and also within single populations. Investigations of

the variability of characters by metrical and statistical methods in some populations dis-

close impressive metrical divergences and morphological differences.

These results give rise to the question whether the characterization of Clausilia dubia as a

polytypic species, as suggested by Kemm (1960, 1973], is justified.

RESUMEN

Estudios morfológicos y biométricos en conchas de algunas poblaciones austriacas de

Clausilia dubia Draparnaud, 1805, muestran una gran variabilidad en el tamaño y en

otros caracteres morfológicos tanto en el conjunto de la especie como en poblaciones ais-

ladas. Investigaciones sobre la variabilidad de caracteres por medio de métodos métricos

y estadísticos de algunas poblaciones mestran importantes divergencias tanto métricas

como morfológicas.

Estos resultados originan la pregunta de si está ¡justificada la caracterización de Clausilia

dubia como una especie politípica, como sugiere KiemM (1960, 1973).

KEY WORDS: Clausilia dubía, subspecies, measures, distribution, morphological continuum.

PALABRAS CLAVE: Clausilia dubía, subespecies, medidas, distribución, continuidad morfológica.

INTRODUCTION

As in great parts of Europe also in

Austria, specially in the eastern parts of

the Alps and in the adjacent areas, Clau-

silia dubiía Draparnaud 1805 is a widely

diffused and in some localities a com-

mon species that is believed to be polyty-

pic (KLeEmMM, 1960, 1973; FECHTER AND

FALKNER, 1990; KEARNEY, CAMERON AND

JUNGBLUTH, 1983; NORDSIECK, 1990).

KLEMM (1960, 1973) gave a survey of di-

* Naturhistorisches Museum Wien, 3. Zoologische Abteilung, Burgring 7, A-1014 Wien, Austria.

Iberus, 15 (2), 1997

S ZP

$9 ] ] [TD

STD

[ID

C. d. dubia

C. d. speciosa C. d. huettneri C. d. schlechti

7 io

57) [TD

27 [>

S [NIP

PIIRAL La

2_/ [TD

|) Dm

7] [PI

— E

E [15

EI]

lA E

C. d. gracilior C. d. tettelbachiana C. d. floningiana C. d. bucculenta — C. d. runensis

C. d. kaeufeli

Figure 1. Distribution and vertical succession of Clausilia dubia subspecies as is suggested by

KLEMM (1960).

Figura 1. Distribución y sucesión vertical de las subespecies de Clausilia dubia, tal y como sugiere

KLEMM (1960).

verse “subspecies” of Clausilia dubia ac-

cording to the characters suggested by

several authors.

Traditional classification was based on

a subjective selection of characters belie-

ved to be important. This classification

depended on the preferences of the single

authors. In some cases, subspecies are

even described according to the presence

of typical features in the shell of few or sin-

gle specimens. HOLOYAK AND SEDDON

(1988) and NORDSIECK (1990) examined

some of these descriptions and offered re-

asonable revisions.

Some of these Clausilia dubia “subspe-

cies” are considered to be widely distri-

buted, others to be localized in small areas

or subdivided into isolated populations

that live in separated sites (KLEMM, 1960,

1973). Furthermore, KLeMM (1960) sug-

gested a vertical pattern of distribution

and an altitude dependent succession of

diverse subspecies at the eastern ranges of

the Alps (Fig. 1). He tried to explain the ob-

served distribution pattern by probable

re-immigration events in the alpine re-

gion after the Pleistocene and adaptation,

enforcing the role of the environment (al-

titude).

Examination and possible revision of

these interpretations have to critically con-

sider the definition and the meaning of the

term subspecies as it is used under va-

rious perspectives by several authors. For

the practical requirement of the collector

and for the taxonomic ordering, of collec-

tions the technical term “subspecies” as ba-

sed on peculiar morphological features

EDLINGER: Shell morphology and variability in Clausilia dubiía Draparnaud, 1805

Rax Schneeberg

Hohe Wand

Wienerwald

Figure 2. A profil of the estern edge of the Alps and the Wienerwald with the sample localities.

Figura 2. Perfil de la cara este de los Alpes y del Wienerwald con las localidades de muestreo.

may certainly be useful. However, it is

scientifically more important to consider

the conceptual background of the termi-

nology. In the given context the concept of

race or subspecies refers to biological units

that are groups of related populations

which are genetically characterized and,

in the case of very long isolation, may be

the forerunners of “valid species” (MaYr,

1967: 387; SUDHAUS AND REHFELD, 1992).

This view is highly important for tradi-

tional evolutionary biology, especially in

the frame of Darwinian models for evo-

lutionary change.

In respect to the biological relevance of

the subspecies concept, Sudhaus and REH-

FELD (1992) are suggesting that “geo-

graphic races” (subspecies) are allopatric

populations of a species, which can be

distinguished taxonomically. The diag-

nostic characters of races should be pre-

sent in 90 or more percent of individuals

of the population. “

OSCHE (1994) claims that 75 percent

or more members of a population must be

morphologically distinguishable from the

members of another population. Popula-

tions only in this case should be accepted

as valid subspecies. In this paper Osche's

definition is assumed as a very tolerant and

useful concept.

According to the presuppositions just

given, studies referring to the subspecies

problem have to treat large numbers of in-

dividuals and to apply methods of grea-

ter exactness than the usual descriptions

and discriminations of characters based on

a subjective and rather arbitrary approach.

It is therefore necessary to study as great

a number as possible of morphological

characters and to take measurements of the

greatest possible exactness to establish a

solid basis for statistical evaluations (NE-

MESCHKAL AND KOTHBAUER, 1988; KoTH-

BAUER, NEMESCHKAL, SATTMANN AND

WAWwRa, 1991; NEMESCHKAL, 1990, 1991,

1993; MYLONAS, KRIMBAS, TSIAKAS AND

AYOUNTANTI, 1990).

Such investigations may clear up,

whether the populations studied by

Klemm meet all requirements of a reliable

identification as subspecies (EDLINGER

AND FISCHER, 1997). It is also worth while

to attempt a critical revision of some sam-

ples of the Museum of Natural History in

Vienna (NHMW). This paper is first at-

tempt at elaborating a new basis for the

discussion of the Clausilia dubia problem

by morphological measurements. In future

the anatomy of the soft parts and geneti-

cal analyses might be also considered.

MATERIAL AND METHODS

606 specimens of Clausilia dubia from

different lots collected in various areas of

the “Wienerwald” (Lower Austria in the

southwest of Vienna) the massif of the

“Hohe Wand”, the “Schneeberg” and the

“Rax” were investigated. The localities

from which the samples came were si-

tuated at altitudes between 270 and 1850

m (Fig. 2).

Iberus, 15 (2), 1997

RA (in mm/10,

x factor 20)

Angle

BR (1-6)

GW 15)

B (in mm/10)

MH (in mm/10)

EA Ñ

MB Gan mm/10)<

LE (1-5)

Figure 3. Measures taken from the shells: shell-height (H), shell-width (B), height (MH) and width

(MB) of the aperture, distance of ribs (RA), number of whorls (WZ), angle between the spindle axis

and the upper palatal (left side) (A).

Figura 3. Medidas tomadas en las conchas: altura (H), anchura (B), altura (MH) y anchura(MB) de la

apertura, distancia de las estrías (RA), número de vueltas (WZ), ángulo entre el eje del huso y palatal

superior (lado izquierdo) (A).

Samples (Number of Sample (Sample

localities, altitude/ numbers of specimens.

R = Rax: R4 (Reichenau, 700 m/1 spec.),

R5 (Aufstieg z. Knappenhof, 730 m/5

spec.), R7 (Knappenhof, 800 m/24 spec.),

R10 (Thórlweg, 850 m/5 spec.), R11 (Thorl-

weg, 960 m/1 spec.), R12 (Thórlweg, 1120

m/29 spec.), R13 (Thórlweg, 1260 m/20

spec.), R14 (Thórlweg, 1320 m/12 spec.),

R15 (Jakobskogel, 1685 m/8 spec.); S =

Schneéberg; S1 (Puchberg, 560 m/15 spec.),

S2 (Schneebergbahn, 750 m/57 spec.), S3

(Schneebergbahn, 790 m / 5 spec.), S4 (Sch-

neebergbahn, 945 m/30 spec.), S5 (Sch-

neebergbahn, 1165 m/9 spec.), S6 (Schne-

ebergbaln, 1370 m/6 spec.), S8 (Waxriegl

11, 1820 m/4 spec.), S9 (Waxriegl 11, 1850

m/26 spec.), S10 (Schneebergbahn, 1650

m/11 spec.), H= Hohe Wand: H1 (Dreistet-

ten, 530 m/60 spec.), H2 (Einhornhóhle,

600 m/12 spec.), H3 (Drobilsteig, 700 m/24

spec.), H4 (Drobilsteig, 760 m/69 spec.), H5

(Auffahrt z. Plateau, 830 m/74 spec.), H6

(Plateau 1020 m/6 spec.), H7 (Plateau 1020

m /34 spec.), H8 (Plateau 1020 m/4 spec.);

W = Wienerwald: W1 (Anninger, 400 m/11

spec.), W2 (Anninger, 450 m/4 spec.), W3

(Moódling-Klause, 260 m/8 spec.), W4 (Hu-

sarentempel, 480 m/14 spec.), W5 (Auf-

gang Aminger, 270 m/11 spec.), W6 (Peils-

tein, 400 m/8 spec.),

Several individuals were dissected.

Dissections did not reveal significant dif-

ferences in the genital apparatus. 18

shells, syntypes of Austrian and South

Tyrolean (Italy) localities, given in loan

by the Natur-Museum Senckenberg as

typical representatives of various subspe-

cies were used for comparisons (Fig. 1).

They are:

1. Clausilia dubia dubiía Draparnaud,

1805 (SMF 163024a)

2. Clausilia dubia speciosa A. Schmidt,

1857 (SMF 163025a)

3. Clausilia dubia speciosa A. Schmidt,

1857 (SMF 163026a)

4. Clausilia dubia obsoleta A. Schmidt,

1857(SMF 163027a)

EDLINGER: Shell morphology and variability in Clausilia dubía Draparnaud, 1805

Figure 4. Shell forms (7 stages from club-shaped -left- to spindle-shaped -right-).

Figura 4. Formas de la concha (siete estadios desde forma de maza -izquierda- hasta abusada -derecha-).

5. Clausilia dubia huettneri Klemm,

1960 (SMF 1630248)

6. Clausilia dubia schlechti A. Schmidt,

1857 (SMF 163030a)

7. Clausilia dubia gracilior Clessin,

1887 (SMF 163031a)

8. Clausilia dubia tettelbachiana Ross-

massler, 1838 (SMF 163032a)

9. Clausilia dubia otvinensis H. Gallens-

tein, 1895 (SMF 163033a)

10. Clausilia dubia grimmeri L.

Pfeiffer, 1848 (SMF 163034a)

11. Clausilia dubia floningiana Tscha-

pek, 1886 (SMF 163035)

12. Clausilia dubia floningiana/gracilior

(SMF 163036)

13. Clausilia dubia bucculenta Klemm,

1960 (Holotypus, SMF 163037)

14. Clausilia dubia runensis Tschapek,

1883 (SMF 163039a)

15. Clausilia dubia moldanubica Klemm,

1960 (Holotypus, SMF 163040)

16. Clausilia dubia kaeufeli Klemm,

1960 (Holotypus, SMF 163042)

17. Clausilia dubia alpicola Clessin,

1878 (SMF 31969)

18. Clausilia dubia reticulata Pini, 1883

(SMF 31936)

The shells were measured under a bi-

nocular microscope; the measurements

were repeated three times. In the case of

different results a special check was

made. The height (H, Fig. 3) and the

width (B, Fig. 3) of the shell as a whole,

the height (MH, Fig. 3) and the width

(MB, Fig. 3) of the aperture, the form of

the aperture (MF, a series of 9 stages

from pear-shaped to deltoid form, the

angle between the spindle axis and the

edge of the upper palatal (0.5 degree

exactness), the mean of 5 rib distances

(RA, Fig. 3) on the last whorl, and the

number of whorls per shell (WZ, exact-

ness: 0.25) were recorded. By compari-

son with stencils the morphological cha-

racters of the form of the shells (GH, a

series of 7 stages from club-shaped, to

extremely spindle-shaped specimens,

Fig 4), the depth, and the thickness of the

basal groove (BR, 6 stages, Fig. 5), the la-

teral internal bulge (GW, on the left side

of the aperture, 5 stages of thickness (Fig.

5), and the incision in the columellar la-

mella (LF, 5 stages, Fig. 5) were recorded.

The measured values of the follo-

wing features were processed by a WIN-

DOWS-EXCEL 5.0 and a WINDOWS

SPSS 6.0 program (BROSIUS AND BROSIUS,

1995):

- Mean of shell heights in each spot

check

- Standard deviation of shell heights

in each spot check

- Mean of shell heights in each spot

check

- Standard deviation shell heights in

each spot check

- Correlation (Pearson s) Coefficient

of all 11 values:

Iberus, 15 (2), 1997

Figure 5. Basal groove (6 stages, upper row); lateral bulge (5 stages, middle row); incision in the

columellar lamella (5 stages, lower row).

Figura 5. Surco basal (6 estadios, arriba); protuberancia lateral (5 estadios, centro); incisión en la lame-

la columelar (5 estadios, abajo).

(R= Pearson“s Coefficient; N=

number of cases; X, Y= variables; Sx,

Sy= standard deviation of the varia-

bles).

By means of the WINDOWS SPSS

6.0 programs a factor extraction and a

principle component analysis were exe-

cuted. “Community” delivers informa-

tion about the quota of spreading of

one value that can be traced back to all

other values. “Eigenvalue” is a value of

the regression factors. It represents the

quota of spreading of all values as

interpreted by special regression

factors. A reduction process restricts the

numbers of factors in the final statistics

to that exceeding 1.0. The factor matrix

shows the influence of the regression

factors on every variable as a percen-

tage of 1.

The measured variables of the

samples in conjunction with the values

of the specimens described by KLEmMM

(1960) and the values of specimens of

Clausilia dubia alpicola and C. d. reticulata

were utilized for computing hierarchical

clusters as dendrograms. For purpose of

cluster analysis the measured values

were transformed to “z-values”, values

with a mean of 0 and a standard devia-

tion of 1. Hierarchical clusters result

100

from dissimilarities computed on the

basis of the sums of squared values of

distances of each character. Thereby the

spectrum of similarities and differences

between all individuals of a spot check

could be elaborated. The dendrograms

contain specimens of various clusters

according to their graduated similarity

(BROSIUS AND BROSIUS, 1995).

The formula of the general distances:

D*= xvi)

(D= distance; v= number of varia-

bles; X, Y= cases)

RESULTS

Means of shell height and shell

width: The means of the shell height

and shell width differ in all sampling

areas. The lowest value was found at the

“Hohe Wand” region, the highest in the

“Wienerwald” area. Comparisons of the

means at different altitudes reveal that

KLEMM's (1960) suggestion of a succes-

sion of different shell heights (according

to a succession of “races” resp. subspe-

cies, high values at low altitudes, low

values at high altitudes) is not generally

convincing (Fig. 7).

EDLINGER: Shell morphology and variability in Clausilia dubia Draparnaud, 18

Figure 6. Two spot-checks of the collection of the NHAMW (Naturhistorisches Museum, Wien). In

the upper row “Clausilia dubia schlechti”, Inv. Nr. 11. 229 NHMW. The specimen in the upper row

at the left belongs to Neostyriaca corynoides (Held, 1836). In the row below C. d. Eschlecht?”, Inv. Nr.

62. 348 NHMW. These spot-checks show us a high variability in the “subspecies”. Scale bar 1 cm.

Figura 6. Dos muestras de la colección del NAMW (Naturbistorisches Museum, Wien). En la fila supe-

rior Clausilia dubia schlechti ”, /nv. Nr. 11. 229 NAMW El especimen de la izquierda de la fila supe-

rior pertenece a Niostyriaca corynoides (Held, 1836). En la fila inferior C. d. “schlechti”, lnv. Nr. 62.

348 NAMW. Las fotografías muetran una alta variabilidad en las subespecies. Escala 1 cm.

Correlation coefficients: A very tude and the shell height (Table 1) but a

remarkable outcome of the study was a high correlation between altitude and

low positive correlation between the alti- the incicion in the columellar lamella.

101

Iberus, 15 (2), 1997

2000 —

AA A

se e.

——_—_—_—.

_ AAA

1000 El TAE dl

A Na —.—

===> =0

== RRRAXáÁ

—— —A—

—e—

5 -

—r— —A—

ai” SYMB

<X 0 T a JETS HA a Y

9 10 11 12 13 14

2000

PUE A

—_A— AAN

—r—

_-—— A _—

—A—

— A —

1000+ e. —e— E

—— EN

. ——

—a—

—a——

E E

O __

Se PARAR

JJ SYMB

< Jl TT E v A A Sano, mE A En)

2,4 2,5 2,56 2,7 2,8 2,9 3,0 3,1

Figure 7. Scatter plots of the means of the height and width of the various samples and the altitu-

de of sampling points.

Figura 7. Diagrama de puntos de las alturas y anchuras medias de las distintas muestras y la altitud de

los puntos de muestreo.

In all four regions a significant corre-

lation between the altitude and the

height of the shells could not be confir-

med. A maximum of correlation was

found between shell height and the

height of the aperture. Correlations bet-

ween shell height and heigth of the

aperture, shell height and number of

whorls, shell form and number of

whorls, shell form and shell height, shell

102

height and width of the aperture, and

between width of the shell and width of

the aperture, are more or less remarka-

ble. Correlation between altitude and

most of the shell variables with the ex-

ception of the columellar lamella (nega-

tive correlation: -0.5123) is low (Table I).

Primary factor analysis: For the

primary factor analysis all values were

EDLINGER: Shell morphology and variability in Clausilia dubía Draparnaud, 1805

Table 1. Correlation coefficients (bivariate) of altitude (ALT), breadth (B), basal groove (BR), shell-

form (GHE), internal bulge (GW), heighth (H), the incision in the columellar lamella (UL), width

of aperture (MB), angle between the upper palatal and the spindle axis (A), height of the aperture

(MH), distance of ribs (R) and number of whorls (WZ).

Tabla I. Coeficientes de correlación (bivariables) de altitud (ALT), anchura (B), surco basal (BR), forma

de la concha (GHE), protuberancia interna (GW), altura (H), incisión en la lamela columelar (UL),

anchura de la apertura (MB), ángulo entre el palatal superior y el eje del huso (A), altura de la apertu-

ra (MH), distancia entre las estrías (R) y número de vueltas (WZ).

A ALT B BR GHF GW

A 1.0000 -.0487 OO .0493 YA .0646

ALT .0487 1.0000 -.0032 ZOOL OOO AUS

B AD -.0032 1.0000 RIGA ZO .0438

BR .0493 SI ZOZS 2 OO00 .0148 SS O/ISS

GHF AY UE -.0036 PDA .0148 1.0000 UA

GW .0646 2082 .0438 SS 0/55 UA 1.0000

H ZO -.0021 OA ASI DUNA SiS

LF .0213 ZII ASIS ZO -.0542

MB .0822* SO; A693** .1606** .0446 ASIS

MH .0450 .0214 EA lA .1644** .0656 OZ

R AIS70SS EY ISS -.0704 LANA O

WZ BUS IZ UA .0127 .0745 IIA .0632

H LF MB MH R WZ

A LU .0213 .0822* .0450 SAO ALZA

ALT .0021 IZ OS .0214 Al Oj JANE

B SONAS AS AOS SATA OSI .0127

BR SMMES ESMAS US .1644** — -.0704 .0745

GHF UNS -.0741 .0446 .0656 LNGRA S/005 +

GW USAS -.0542 SSI OZ OS .0632

H 1.0000 .0780 .4968** YA ALE AS .6663**

LF .0780 1.0000 2200 .0913* Soles .0891*

MB A968** E2DO Oi 1.0000 OA OZ 0 II

MH 07M 07/13 OA OOOO -.0412 LAS

R AMI ISO -.0230 -.0412 1.0000 2LOTOSO

WZ .6603** .0891* SS ASS 200% 1.0000

*= Signif. LE .05;**= Signif. LE .01 (2-tailed)

used irrespective of the altitude. The

analyses result four factors with an

Eigenvalue of more than 1.0. One analy-

sis was done including the NMS speci-

mens (Table II), the other only with the

own samples (Table III). The results of

both analyses were corresponding at a

high degree.

Factor 1 has a significant influence on

the width of the shell, the shell height, the

width of the aperture, the height of the

aperture and the number of whorls.

Factor 2 has a significant influence on

the angle between the axis and the left

palatal, the width of the aperture, the rib

distance and the number of whorls, factor

103

Iberus, 15 (2), 1997

Table H. Primary component analysis factors of all samples including the NMS specimens.

Abbreviations as in Table 1.

Tabla II. Factores del análisis de componentes principales de todas las mustras incluyendo los ejemplares

NMS. Abreviaturas como en la Tabla 1.

Initial Statistics

Variable Communality Factor Eigenvalue Pctof Var Cum Pct

A 1.00000 l SADO 31.4 31.4

B 1.00000 2 1.48937 14.9 46.3

BR 1.00000 3 1.34787 13.5 59.8

GW 1.00000 4 1.09305 10.9 O)

H 1.00000 5 .81589 8.2 78.9

LF 1.00000 o 697772 7.0 85.9

MB 1.00000 7 34068 5.4 ONES

MH 1.00000 8 42572 43 0)

R 1.00000 9 .32418 3.2 98.8

WZ 1.00000 10 .12276 122 100.0

PC extracted 4 factors

Factor Matrix

Factor 1 Factor 2 Factor 3 Factor 4

A 24107 -.39349 /SZI -.01561

B .64730 .28942 -. 49248 -.01712

BR .19488 39543 32902 22231

GW .11069 .S0354 33469 -.34493

H .89714 .19205 .07467 -.16390

LF .25618 .22266 .08430 .85909

MB .75790 .21202 -.17180 .03663

MH . 85718 . 10931 -.19219 -.16564

R .24758 35811 .01044 III

WZ 61193 - 3340 36292 -.06089

Final Statistics

Variable Communality Factor Eigenvalue Pctof Var Cum Pct

A 54182 1 3.14275 31.4 31.4

B 74559 2 1.48937 14.9 46.3

BR IND 3 1.34787 138 59.8

GW .67067 4 1.09305 10.9 70.7

H .87418

LF .86036

MB .65022

MM .81108

R 49962

WZ 69772

Skipping rotation 1for extraction 1 in analysis 1

104

EDLINGER: Shell morphology and variability in Clausilia dubia Draparnaud, 180

Table III. Primary component analysis factors of all samples excluding the NMS specimens.

Abbreviations as in Table I.

Tabla II. Factores del análisis de componentes principales excluyendo los ejemplares NMS. Abreviaturas

como en la Tabla 1.

Initial Statistics

Variable Communality Factor Eigenvalue PctofVar Cum Pct*

A 1.00000 l 2.98596 DS 29

B 1.00000 2 1.52656 158 A5.]

BR 1.00000 3 1.27032 11247 57.8

GW 1.00000 4 1.19526 12.0 69.8

H 1.00000 S .79324 VS VUL)

LF 1.00000 o .70682 Je 84.8

MM 1.00000 7 36029 5.6 90.4

MM 1.00000 8 A5357 45 94.9

R 1.00000 9 .36803 3.7 98.6

WZ 1.00000 10 .13995 1.4 100.0

PC extracted 4 factors

Factor Matrix

Factor 1 Factor 2 Factor 3 Factor 4

A .19408 .61541 28529 -.11267

B .60129 -.52606 -.27656 .02153

BR .32508 -.17128 64145 42482

GW 25481 ISS 76411 -.07054

H .86917 22511 -.06761 -.24429

LF .28165 05486 -.10404 .82738

MM 74584 -.22587 -.11384 .04898

MM .83214 -.22388 -.10785 -.18500

R .18873 33391 -.27470 43010

WZ ALO .61465 .04084 -.17555

Final Statistics

Variable Communality Factor Eigenvalue PctofVar Cum Pct

A 51047 1 2.98596 DAS 29.9

B ASZS 2 1.52656 1578 45.1

BR .72695 3 1.27032 11227 57.8

GW .67784 4 1.19526 12.0 69.8

H .87038

LF UY UIN

MM .62266

MM 78843

R 38114

WZ .70730

Skipping rotation 1 for extraction 1 in analysis 1

105

Iberus, 15 (2), 1997

A r

' a ES

: ¡Y al. E

-,8 8 142

(pl

ZONA ASS ZA ES :6

¡O)

-2,8 -2,4 -2,0

FAC1_1

Count

FAC 2 _1

30 |

V E

p ll hi E

S ¡ q , Mr mv

10) -,8 38 1,2

20 24 28 36 4.0 4,8

Figure 8. Bar charts of factor 1 and 2 (spot checks differentiated).

Figura 8. Diagrama de barras de los factores 1 y 2 (pruebas diferenciadas).

on the basal groove and factor 4 has a sig-

nificant influence on the columellar

lamella.

Bar charts show us the distributions

and the maximum of the values of the

primary factors in the samples for com-

parison.

The values of factor 1 have a similar

distribution in the Schneeberg, the Rax

106

and the Hohe Wand area and another dis-

tribution pattern with another maximum

in the Wienerwald area (Fig. 8).

The values of factor 2 show us a very

similar distribution in the Rax and the Sch-

neeberg, area, but other patterns in the Hohe

Wand and the Wienerwald area (Fig. 8).

Factor 3 has similar distributions of

values in the samples of Rax, Schneeberg

EDLINGER: Shell morphology and variability in Clausilia dubiía Draparnaud, 1805

pun

[o]

DA 20 1 12

FAC3_1

[e)

0.0

DE DE A)

FAC4 1

ME a

¡Er

El Dom.

¿All A ¡A My

32. =2,8 2 -,8 -,4 0 4 8 1 1,6 2,0 2,4

Figure 9. Bar charts of factor 3 and 4 (spot checks differentiated).

Figura 9. Diagrama de barras de los factores 3 y 4 (pruebas diferenciadas).

and Hohe Wand, but another maximum

in the Sample of the Wienerwald area

(Fig. 9).

Diagrams of the values of factor 4

show us different distributions in all

samples (Fig. 9).

A two dimensional scatter plot of

factor 1 and factor 2 for a comparison of

the samples of the four areas (Fig. 10)

shows us, that by the positions of the spe-

cimens and by their pattern of distribu-

tion only two partially different groups

can be distinguished: one group consis-

ting of specimens of the Hohe Wand, the

Schneeberg and the Rax area at the one

side and a second group with the speci-

mens from the Wienerwald area at the

other. Both groups are overlapping.

107

Iberus, 15 (2), 1997

REGR factor score 2 for analysis 1

REGR factor score 1 for analysis 1

REGR factor score 4 for analysis 1

-4 3 S) aj

REGAR factor score 3 for analysis 1

Figure 10. Scatter plots of the samples with value 1 and 2, 3 and 4 (spot checks differentiated).

Figura 10. Diagramas de puntos de las muestras con los valores 1 y 2, 3 y 4 (pruebas diferenciadas).

A scatter plot with factor 3 and factor

4 (Fig. 10) shows us a wide distribution

of the specimens of the Schneeberg area

and a partial separation of the samples

of the Rax area at the one and the

samples of the Hohe Wand and the Wie-

nerwald area at the other side. Also these

distribution areas of the samples are

overlapping at a high degree.

108

Scatter plots of regression factor 1

and 2 of all samples including the SMF

specimens based on three morphologi-

cally important measures (Fig. 11) disclo-

ses a remarkable morphological isolation

of some SMF specimens, especially of

those specimens assigned to Clausilia du-

bia speciosa, C. d. dubia, C. d. floningiana

and C. d. floningiana/gracilor, C. d. graci-

EDLINGER: Shell morphology and variability in Clausilia dubiía Draparnaud, 1805

REGAR factor score 2 for analysis 1

4 E 0

REGR factor score 1 for analysis 1

REGR factor score 4 for analysis 1

4 ES E) E

REGAR factor score 3 for analysis 1

Figure 11. Scatter plots of the samples and the NMS specimens with value 1 and 2, 3 and 4 (points

represent samples, NMS specimens with abbreviations).

Figura 11. Diagramas de puntos de las muestras y los especímenes NMS con valores 1 y 2, 3 y 4 (los pun-

tos representan las muestras, los especímenes NMS son las abreviaturas).

lior, C. d. grimmeri, and also C. d. buccu-

lenta. These specimens are found outside

the central area of distribution of the two

dimensional coordinate system of the

scatter plot, where the bulk of specimens

appears in a high concentration.

A scatter plot of the factors 3 and 4

(Fig. 11) shows us more peripheral posi-

tions of Clausilia dubia moldanubica, C. d.

otvinensis, C. d. speciosa, C. d. kaeufeli, C.

d. gracilior, C. d. schlechti, C. d. grimmeri

and C. d. floningiana/gracilior.

109

Iberus, 15 (2), 1997

FAC1_2

FAC4_1

3 2) E

FAC3_1

Figure 12. Two-dimensional scatter plots with factors 1/2 and 3/4 of a spot check of the Schneeberg

samples (see the altitude of the sample areas at Material and methods).

Figura 12. Diagramas de puntos de dos dimensiones con factores 1/2 y 3/4 de las muestras de Schneeberg

(ver la altitud de las áreas de muestreo en Material y métodos).

Scatter plots of the factors 1 to 4 of

the various samples of the Schneeberg

area (from different altitudes) disclose

no remarkable dependence of the

factors 1, 3 and 4 on altitude (Fig. 12).

Factor 2 shows us a separate distribu-

tion of values of sample S9 (1850 m) and

110

S10 (1650 m). The distribution areas of the

other samples are covering one another.

They are overlapping distribution areas

of sample S9 and S10 insignificantly.

A scatter plot of the Rax samples

(Fig. 13) which were arranged in corres-

pondence to the altitude in three groups

EDLINGER: Shell morphology and variability in Clausilia dubiía Draparnaud, 1805

FAC2_1

FAC1_1

FAC4_1

FAC3_1

o 1000-1500m

o >1500m

“+. <1000m

e 1000-1500m

vo >1500m

. <1000m

Figure 13. Two-dimensional scatter plots with factors 1/2 and 3/4 of a spot check of the Rax sam-

ples, arranged in three groups in accordance with the altitude of the sample areas (less 1000 m,

1000-1500 m, more than 1500 m).

Figura 13. Diagramas de puntos de dos dimensiones con factores 1/2 y 3/4 de las muestras de Rax , orde-

nadas en tres grupos de acuerdo con la altitud de las áreas de muestreo (menos de 1000 m, 100-1500 m,

más de 1500 m).

(less than 1000m, 100-1500m, more than

1500m) results no altitude dependence

of the factors.

Cluster analysis: For the first cluster

analysis the measured values were ta-

ken from a group of specimens which

consisted of selected samples from diffe-

rent localities and altitudes; data from

the SMF specimens were also included

(Fig. 14). The hierarchical cluster which

is presented as a dendrogram contains

groups of different size which were co-

llected at different localities.

111

Iberus, 15 (2), 1997

Notable is the isolated position of

the SMF specimens of Clausilia dubia spe-

ciosa and C. d. dubia which constitute a

cluster of their own together with two

specimens of the samples. Almost the

same phenomena occur in dendrograms

of samples collected in the areas “Hohe

Wand”, “Schneeberg” and “Rax”. In

these cases specimens of the SMF were

also taken into consideration.

In general the clusters show remar-

kable segregations. Similarities of the

individuals coming out from the same

locality and appearing together in clus-

ters may be seen as indication of close

relationship.

At the other side the spot checks

from various areas are overlapping at a

high degree. Relevant portions of indivi-

duals, which present all the characters

of several “subspecies” (subspecies seen

in the traditional way) do not occur. It is

remarkable that most specimens from

the SMF which were considered to be

typical for specific regions, appear also

in the different branches of the dendro-

grams and in various clusters.

Only the SMF specimens of Clausilia

dubia speciosa, and in a most astonishing

way, the SMF specimen of C. d. dubia, an

individual belonging to the nominotypi-

cal subspecies” are seen at own separate

branches of the dendrograms. This

finding is in full agreement with the

above mentioned scatter plots of the

analysis of the main components. All

three above mentioned specimens are

characterized by strongly deviating

measures and stand in isolated posi-

tions. It is evident that this finding disa-

grees with the geographical distribution

shown in the literature (KLeMM, 1960).

Wienerwald: The samples from the

Wienerwald area come from altitudes

between 270 and 400 m and sites similar

in climate and ecological conditions (fir,

pine, and mixed forests).

The shells are rather similar and club

shaped, but differ considerably in shell

height and width, height and width of

aperture, and distance of ribs. The same

is true for all the other measures taken.

In the hierarchical cluster analysis of

samples from the Wienerwald area (Fig.

15) which was considered to be the type

locality of Clausilia dubia dubia, the posi-

tion of C. d. dubia, C. d. kaeufeli, and C. d.

speciosa is found to be extremely isolated

in a cluster of their own (Fig. 16). Within

the second cluster also other SMF speci-

mens appear in entirely isolated bran-

ches. Only C. d. runensis, C. d. tettelba-

chiana, C. d. grimmeri, C. d. schlechti and

C. d. moldanubica appear in a branch

together with the specimens of the

samples from the Wienerwald area.

Hohe Wand: Analyses of samples

taken from the Hohe Wand (altitude

between 560 m and 1080 m; Fig. 16) also

lead to results which are not in accor-

dance with generally held views. Clau-

silia dubia speciosa and C. d. runensis are

in an isolated position. They are in a

cluster of their own together with one

specimen of the local sample from H8.

Also C. d. bucculenta, C. d. floningia-

na/lgracilior, C. d. floningiana, C. d. reticu-

lata, C. d. obsoleta, C. d. gracilior, C. d.

(Right page). Figure 14. Hierarchical cluster of a spot-check of all samples and the SMEF specimens

(Wi-y= specimens of the Wienerwald area; i= sample; y= number of the specimen; Hi-y= specimens

of the Hohe Wand area; Si-y= specimens of the Schneeberg area; Ri-y= specimens of the Rax area;

Cd-= specimens of the SME: du= dubia; sp= speciosa; ob= obsoleta; hn= huettneri, sc= schlechti, gr=

gracilior, te= tettelbachiana, ot= otvinensis, gi= grimmeri, l= floningiana; flgr= floningianal gracilior,

bu= bucculenta; ru= runensis, mo= moldanubica; ki= kaeufeli, al= alpicola; re= reticulata).

(Página derecha). Figura 14. Cluster de todas las muestras y los especimenes SMF (Wi-y= especímenes del

área de Wienerwald; ¡= muestra; y= número del especimen; Hi-y= especímenes del área de Wand; Si-y=

especímenes del área de Schneeberg; Ri-y= especímenes del área de Rax; Cd-= especímenes SMF: du=

dubia; sp= speciosa; 0b= obsoleta; /n= huettneri; sc= schlechti; gr= gracilior; te= tetrelbachiana; 01=

otvinensis; g/= grimmeri, fl= floningiana, flgr= floningiana/gracilior; b4= bucculenta; 7u= runensis;

mo= moldanubica; Rá= kaeufeli; 4/= alpicola; re= reticulata).

112

EDLINGER: Shell morphology and variability in Clausilia dubía Draparnaud, 180

Cdob

Cdflgi

Cdbu

cafl

Cdsp

Cddu

Rescaled Distance Cluster Combine

EJE

113

Iberus, 15 (2), 1997

Rescaled Distance Cluster Combine

CASE 0 5 10 15 20

Label Num —+-=-=--=-=-=--- Moscas MiS E O o ==

w1 6

wl 7 ia]

wl1 3

w1 4 z

W4 33

w1 5 E]

wW2 12

W4 27 y]

W3 23

w1 2

W4 30 poa

W4 29 ——

W3 24

Cdmo 73 Al

wW3 18 op po

W3 20 —

W2 14 ==

wW4 3/9) —

W3 119 —

wW4 38 =>

W2 16 =

w2 157) 4 —

W3 22 ==

W4 31

W4 32 7

w4 28 —

W4 34 a)

W3 21 A

Wa 36 ——

WS 50 Á

W3 25 ==

wW4 37 — >

W4 35 —

wW2 13 ——

WS 42

WS 40 =>

W5 41 — HZ

W2 15 ———— 7

w1 de

WS 44 0 ¡FR

WS 45

W5 48 —

W6 52

WS 47 — 0)

W6 a) ==

W6 56 —

W6 53

W6 54 -=)

WS 43

W4 26 ===

WS 46

W5 49 — HH

W6 Sl A [EU

wl1 8

w1 11 iS

Wl 10

w Sparta

Cdsc 64 =——,

Cdgi 68 EIA ===)

Cdte 66

Cdru 72 SIS y)

W6 57

W6 58 A ¡E

Cdflgi 70 =>

Cdbu NS EA E

Cdga 65

Cdre 76 A

Cdob 62 _—A

Cdhn cp AS

Cdot 67

Cdal IS pe

Cdká 74 E EE

cdf1l 69

Cddu 59

Cdsp 6l ] '

Cdsp 60 Ea

Figure 15. Hierarchical cluster of a spot check of all samples of the Wienerwald area and the SMF

specimens. Abbreviations as in Figure 14. |

Figura 15. Cluster de todas las muestras del área de Wienerwald y los especímenes SME Abreviaturas

como en la Figura 14.

114

EDLINGER: Shell morphology and variability in Clausilia dubía Draparnaud, 1805

Rescaled Distance Cluster Combine

CASE 0 5 10 15 20 25

Label Num +--------- OS ASFALTO ERARIO a E

H5 39

HS 43

HS 41

H5 42

H6-2 46

H6-4 48

H5 40

H2 9

H6- 3 47

H5 35 —

Cdká 76

H4 24

Cdsc 66

H1 1 [asta

H8-2 58

HS 34

H8-4 60

Cdhn 65

HS 44

H6-6 50

H7-25 53

H8-3 s9

H7-24 52

H7-23 51 —y

H1 3

Cdmo 75 Tf)

H3 14

H3 22

H1 5

H2 8

H7-27 55

H7-26 54 ——

H4 26 ES

do ¿E A

H4 23

H4 25

H1 6

H2 10

H1 7 |

H4 29

H3 19

H7-28 56 J

H3 15

H3 18

H3 21

H5 38

H6-1 45

H2 12

H4 32

pitos

H3 20

HS 36 ———

H4 27

H4 30

H1 2

H3 16

H3 17

H4 28

H2 11

H3 13

H5 37 F

Cdgi 70

Cdot 69

Cdal 27

HS 33 ae

Cdga 67 1] -

Cdob 64 PR

Cdre 78

Ccdfl 71

Cdflgi 72

Cdbu 73 E

Cdsp 62

Cdsp 63 En _———

H8-1 57

Cddu 61 Ertasto de a]

Figure 16. Hierarchical cluster of a spot check of all samples of the Hohe Wand area and the SMF

specimens. Abbreviations as in Figure 14.

Figura 16. Cluster de todas las muestras del área de Hohe Wand y los especímenes SME Abreviaturas

como en la Figura 14.

115

Iberus, 15 (2), 1997

alpicola, C. d. otvinensis and C. d. grim-

meri occur in separated branches and

only C. d. tettelbachiana, C. d. moldanu-

bica, C. d. huettneri and C. d. kaeufeli are

integrated in branches together with

the major part of the specimens from

the Hohe Wand area. There are no indi-

cations of the succession postulated by

former workers (KLeMM, 1960) when

one takes into consideration altitudes of

the localities from which they came and

the similarities that appear in the den-

drogram.

Schneeberg: The studied material

was collected at altitudes between 700 to

1850 m. The cluster analysis (Fig. 17)

reveals morphologically isolated posi-

tions for Clausilia dubia speciosa, C. d.

dubia, C. d. kaeufeli, C. d. floningiana, C. d.

reticulata, C. d. obsoleta, C. d. bucculenta,

C. d. floningiana / gracilior, C. d. alpicola, C.

d. otvinensis, C. d. tettelbachiana, C. d.

runensis and C. d. moldanubica. Only C. d.

grimmeri, C. d. gracilior and C. d. schlechti

occur in branches together with most of

the specimens of the Schneeberg spot

checks. Remarkable is that specimens

with close morphological relations to C.

d. kaeufeli, as expected for the peak of

the mountain, don't occur in the clus-

ters. The SMF specimen of C. d. kaeufeli

is isolated in the dendrogram.

Rax: The studied material was

collected at altitudes between 700 to

1685 m. As in the Clusters analyses of

the other areas Clausilia dubia dubia, and

C. d. speciosa are isolated in a Cluster of

ist own (Fig. 18). Also C. d. kaeufeli, C. d.

floningiana, C. d. bucculenta, C. d. flonin-

giana / gracilior, C. d. runensis, C. d. alpi-

cola, C. d. otvinensis, C. d. otvinensis, C. d.

obsoleta, C. d. reticulata and C. d. gracilior

occur very isolated in branches together

with only few specimens of the local

samples. Only C. d. grimmeri, C. d. mol-

danubica, C. d. tettelbachiana, C. d. sch-

lechti and C. d. huettneri can be seen as

well integrated in clusters with the

major part of the local samples. No

remarkable position of C. d. kaeufeli or a

succession as suggested by Klemm was

visible.

116

DISCUSSION AND CONCLUSION

The conclusions presented here must

be seen as being valid only for samples

of Clausilia dubia collected in restricted

areas. Only further research might lead

to conclusive results on possible geo-

graphic variation in C. dubia.

The results of the present research as

well as a critical evaluation of those of

earlier researchers show us the limits

inherent the intuitive, subjective method

used in the analysis of characters.

Earlier studies by EDLINGER AND

MILDNER (1979) on Clausilia dubia in

Carinthia, using traditional morphologi-

cal methods based on a number of shell

characters, also showed a high variabi-

lity of most of these characters within

each population. This may be a common

phenomenon when analyses are based

on a large number of characters. In this

Case, techniques of measurement, as

well as applied statistical methods

might present potential sources of error.

Nevertheless, all observations are

indicative of high variability within the

species. Similar observations can be

made in many of the collection samples

(Fig. 6). This can be clearly discerned, in

spite of the fact that many of these

samples have been classified as belon-

ging to various subspecies.

To gain a better understanding of

shell variability, we must also consider

the influence of ecological factors, and

the life history of the specimens. Morp-

hological features may be influenced by

non hereditary factors too (GOODFRIEND,

1986).

Against the background of low

correlation coefficients between altitude

and most characters, except columellar

lamella, altitude in itself cannot be con-

ceived as an substantial ecological

factor, because the measures of the

shells except the columellar lamella

don't vary significantly in correlation

with altitude. A reason for the remarka-

ble correlation of altitude with the colu-

mellar lamella might be that the Clausi-

lia dubia shells of the Wienerwald area

commonly have very pregnant incisions

of the columellar lamella.

EDLINGER: Shell morphology and variability in C/ausilia dubia Draparnaud, 1805

Rescaled Distance Cluster Combine

Cdmo

Cdru

Cdte

s10

Cdot

Cdal

Cdflgi

Cdbu

Cdob

Cdre

Ccdf1

Cdká

Cddu

Cdsp

Num

7/3

0 5 10 15 20 25

gol

ll ejes

==.

UTE

Ip uu

Figure 17. Hierarchical cluster of a spot check of all samples of the Schneeberg area and the SMF

specimens. Abbreviations as in Figure 14.

Figura 17. Cluster de todas las muestras del área de Schneeberg y los especímenes SME Abreviaturas como

en la Figura 14.

117

Iberus, 15 (2), 1997

Rescaled Distance Cluster Combine

DIU

SBPBRRA An e

VwWWWVWVWA NN

POUOWWVWWWW ny

>bBIAOIWADIOBONIW

R7 15

R10 20

R13 31

R14 44

R14 46

R14 47 =

R14 41

R14 42

R14 43

R15 50

R15 Sp

R7 Y aa

R7 13

R10 21

R14 38

R12 23 JA

R13 29

R12 24 NA

R12 25

cal

RS 6

Cdhn 60 a)

R7 el

R7 12

R7 9 a

R14 40

Cdsc 61

R7 19

Cdte 63

R14 45

R14 49

R14 48 —

Cdmo 70

Cdgi 65

RS 2

R7 17

R11 22

Cdga 62

Cdre ve )

Cdob 59

Cdot 64

Cdal 72 AN

Cdru 69

R4 1

Cdflgi 67

AS pa

cdf1 66

Cdká 71

Cdsp 57

Cdsp 58

Cddu 56

Figure 18. Hierarchical cluster of a spot check of all samples of the Rax area and the SMF speci-

mens. Abbreviations as in Figure 14.

Figura 18. Cluster de todas las muestras del área de Rax y los especímenes SME Abreviaturas como en la

Figura 14.

118

EDLINGER: Shell morphology and variability in Clausilia dubiía Draparnaud, 1805

Other ecological factors may be of

relevance, but we must consider them as

arranged in ensembles and affecting

upon animals collectively.

Certainly, individual living condi-

tions, as well as hereditary dispositions,

result in specific modifications of cha-

racters. nevertheless we can reject the

hypothesis of EDLINGER AND MILDNER

(1982) that the characteristics of Clausilia

dubia runensis might be a result of hel-

minth parasitism.

A representation of variability of

species or populations by character

analysis also has to take into considera-

tion the fact that many characters act as

correlated variables. In these cases one

can speak of character complexes.

Arrangements in complexes restrict

variability. The question arises, whether

there are other factors influencing the

variables mentioned above, which are

not subjects of the researches presented

above. Shells, for example, function as

hard skeletons and as mechanical abut-

ments for musculature. Hard skeleton

and musculature must be suited to each

other. So it is evident, that morphologi-

cal relations between various measures

of the shell are indirectly caused by

constructional needs of the animal as a

whole and also of the soft body in parti-

cular (GUTMANN, 1989; EDLINGER, 1991).

Considering the local patterns of dis-

tribution of values and primary compo-

nents inside the species and its popula-

tions, these patterns must be seen as

varying locally and gradually. The

variation of patterns must be seen as the

result of local variation, following from

step by step changes of frequencies of

characters. When different frequencies

are developed under similar ecological

conditions, we may assume that these

changes are a result of genetic drift.

Certain local accumulation of special

characters resp. values of variables may

result from genetic drift too. Typical

characters of the so called “Clausilia

dubia runensis”, C. d. speciosa (shell form)

or “C. d. otvinensis” (distances between

shell ribs), for example, occur in sepa-

rate areas with large distances between

them. Other characters of the same

animals very often don't differ from that

of surrounding populations in adjacent

areas (EDLINGER AND FISCHER, 1997).

This leads us to the conclusion that

frequencies of special characters cannot

automatically be taken as a reason for a

common origin of separated popula-

tions resp. of a heterogeneous origin of

contiguous populations. Above all this

is true, when other characters are identi-

cal with those of contiguous popula-

tions.

So it may be that genetic drift or

special environmental factors have an

influence on single characters. With

regard to these characters we may con-

ceive of some populations as homoge-

nous and “pure bred”. At the other side

there cannot arise populations being

homogeneous and “pure bred” in all or

most characters and containing such a

high number of “typical specimens”

homogenous in most or in all characters

by simple environmental influences.

In any case, “typical” or “pure bred”

specimens were recognized mostly by

earlier researchers, and are extremely

arbitrary. So, the natural populations of

Clausilia dubia which were investigated

do not match the preconceived expecta-

tions of well established races or subs-

pecies occurring in well delimited areas

with rare interbreeding occurrences.

Therefore, we must question, if a morp-

hologically uniform population of Clau-

silia dubia which can be defined as a

subspecies or race, might ever have

existed in the eastern Alpine region.

KLEMM (1960) who was of the same

opinion, believed that the transitional

stages between the races and subspecies

might be the consequence of (post

glacial) re-immigration of pure bred

populations, and their subsequent

mixing with other local forms.

Shell characters, and their distribu-

tion patterns do not support this hypot-

hesis. Additionally, cluster analysis

shows that very variable samples come

from regions where the forerunners of

present populations must definitely

have lived and survived during the

Pleistocene. Contrary to KLemMM (1960),

we must state that a reconstruction of

119

Iberus, 15 (2), 1997

events after the Pleistocene gives us no

evidence to support this view. Why

should areas, glaciated during the Pleis-

tocene be resettled by the descendants

of single, pure bred populations after

the disappearance of the ice?

A new interpretation of character

distribution seems to be more adequate

for the case presented, and discussed

here. This interpretation requires the

(for the case discussed here) use of theo-

rems which are accepted by most anth-

ropologists (CAVALLI-SFORZA, MENOZZI

AND PIAZZA, 1994; KLEIN, TAKAHATA

AND AYALA, 1994).

According to their theoretical guide-

lines, the definition of subspecies and

race depends on subjective argumenta-

tions, and does not mirror objective,

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CAVALLI-SFORZA, L. L., MENOZZI, P. AND

PIAZZA, A., 1994. The History and Geography

of Human Genes. Princeton, NJ.

EDLINGER, K., 1991. The mechanical constraint's

in mollusc constructions. The function of the

shell, the musculature, and the connective tis-

sue. In Schmidt-Kittler, N, and Vogel, K.

(Eds.): Constructional morphology and evolution.

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EDLINGER, K. AND MILDNER, P., 1979. Mono-

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GOODFRIEND, G. A., 1986. Variation in land-

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120

concrete facts. They also accept diversity

within given populations. Therefore,

distribution and frequency of characters

are subject to constant change.

Thus, in any investigated snail

population, only the distributions of

characters, and the frequency of their

occurrence can be reasonably recorded.

This entirely altered interpretation also

corresponds better with the results of

genetic analysis in various species of

Albinaría (MYLONAS, KRIMBAS, TSIAKAS

AND AYOUNTANTI, 1990; SCHILTHUIZEN,

1994). The results of these studies show

that there is an overlap of genetic featu-

res in some of the traditionally recogni-

zed subspecies, and even of species.